MONOGRAPH OF THE AMPHISIELLIDAE AND TRACHELOSTYLIDAE (CILIOPHORA, HYPOTRICHA)
MONOGRAPHIAE BIOLOGICAE VOLUME 88
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H. J. Dumont
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Monograph of the Amphisiellidae and Trachelostylidae (Ciliophora, Hypotricha) by HELMUT BERGER Consulting Engineering Office for Ecology Salzburg, Austria and University of Salzburg Department of Organismal Biology Salzburg, Austria
Helmut Berger Consulting Engineering Office for Ecology Radetzkystrasse 10 5020 Salzburg Austria and Department of Organismal Biology University of Salzburg Hellbrunnerstrasse 34 5020 Salzburg Austria
[email protected]
ISBN: 978-1-4020-8916-9
e-ISBN: 978-1-4020-8917-6
Library of Congress Control Number:2008933595
© 2008 Springer Science + Business Media B.V. No part of this work may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, microfilming, recording or otherwise, without written permission from the Publisher, with the exception of any material supplied specifically for the purpose of being entered and executed on a computer system, for exclusive use by the purchaser of the work. Cover Illustration: Amphisiella annulata, illustration by Berger H. (2004): Amphisiella annulata (Kahl, 1928) Borror, 1972 (Ciliophora: Hypotricha): morphology, notes on morphogenesis, review of literature, and neotypification. – Acta Protozool. 43: 1–14. Printed on acid-free paper. 9 8 7 6 5 4 3 2 1 springer.com
Dedication This book is dedicated to my mentor and friend Wilhelm (“Willi”) Foissner (University of Salzburg, Austria) on the occasion of his 60th birthday. Willi is a sedulous worker who provided many significant contributions to the systematics of the amphisiellids
Contents Preface. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xi Acknowledgements and Permissions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xv A General Section. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 1 Morphology, Biology, and Terminology. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 1.1 Size and Shape. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 1.2 Nuclear Apparatus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 1.3 Contractile Vacuole and Cytopyge. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 1.4 Cytoplasm, Cortex, and Colouring. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 1.5 Cortical Granules. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 1.6 Movement. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 1.7 Somatic Ciliature and Ultrastructure. . . . . . . . . . . . . . . . . . . . . . . . . . 10 1.8 Oral Apparatus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 1.9 Silverline System. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 1.10 Life Cycle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 1.10.1 Cell Division. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 1.10.2 Conjugation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 1.10.3 Cyst. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 1.10.4 Reorganisation, Regeneration, Doublets. . . . . . . . . . . . . . . . . . 22 2 Phylogeny. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 2.1 Notes on Nomenclature. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 2.2 The Ground Pattern of the Hypotricha Stein, 1859. . . . . . . . . . . . . . . 23 2.2.1 Apomorphies of the Hypotricha. . . . . . . . . . . . . . . . . . . . . . . . 23 2.2.2 Plesiomorphies of the Hypotricha. . . . . . . . . . . . . . . . . . . . . . . 29 2.2.3 Features not Considered in the Ground Pattern. . . . . . . . . . . . . 38 2.3 Comments on the Evolution within the Spirotricha. . . . . . . . . . . . . . . 42 2.4 Comments on the Evolution within the Hypotricha. . . . . . . . . . . . . . . 44 3 Previous Classifications and Revisions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48 4 Parasitism. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51 5 Ecology, Occurrence, and Geographic Distribution. . . . . . . . . . . . . . . . . . . 51 6 Collecting, Culturing, Observing, and Staining of Hypotrichous Ciliates. . 53 6.1 Collecting and Culturing. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53 6.2 Observing Living Hypotrichs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54 6.3 Staining Procedures. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55 6.3.1 Feulgen Nuclear Reaction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55 6.3.2 Supravital Staining with Methyl Green-Pyronin. . . . . . . . . . . . 55 6.3.3 Protargol Methods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56 6.4 Preparation for Scanning Electron Microscopy. . . . . . . . . . . . . . . . . . 65 7 Species Concept and Nomenclature. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65 7.1 Species Concept. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65 7.2 Nomenclature. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66 vii
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7.3 Summary of New Taxa and Nomenclatural Acts. . . . . . . . . . . . . . . . . 67 7.4 Deposition of Slides. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69 B Systematic Section. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71 Key to the Taxa Treated in Present Book. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71 Amphisiellidae (59 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79 Group I: Marine Amphisiellids (9 species). . . . . . . . . . . . . . . . . . . . . . . . . . 84 Amphisiella (5 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84 Caudiamphisiella (2 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129 Maregastrostyla (1 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136 Spiroamphisiella (1 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148 Group II: Terrestrial Amphisiellids with six (I–VI) Frontal-ventral-transverse Cirri Anlagen (25 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161 Lamtostyla (12 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161 Lamtostyla lamottei-group (8 species). . . . . . . . . . . . . . . . . . . . . . . 167 Lamtostyla granulifera-group (2 species). . . . . . . . . . . . . . . . . . . . 205 Lamtostyla longa-group (2 species). . . . . . . . . . . . . . . . . . . . . . . . . 218 Uroleptoides (8 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 224 Hemiamphisiella (5 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 288 Group III: Terrestrial Amphisiellids which lack Frontal-ventral-transverse Cirri Anlage IV (7 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 322 Lamtostylides (5 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 322 Paramphisiella (2 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 351 Incertae sedis in the Amphisiellidae (18 species). . . . . . . . . . . . . . . . . . . . 370 Afroamphisiella (2 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371 Cossothigma (1 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 382 Hemisincirra (10 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 387 Mucotrichidium (1 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 440 Terricirra (3 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 447 Tetrastyla (1 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 463 Taxa not Considered in the Amphisiellidae. . . . . . . . . . . . . . . . . . . . . . . . 466 Trachelostylidae (6 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 471 Trachelostyla (3 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 474 Spirotrachelostyla (3 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 502 Taxa not Considered in the Trachelostylidae. . . . . . . . . . . . . . . . . . . . . . . 512 Taxa of Unknown Position in the Hypotricha (7 species). . . . . . . . . . . . . . 514 Apourosomoida (2 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 514 Bistichella (5 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 532 Taxa of Unknown Position in the Non-oxytrichid Dorsomarginalia (12 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 560 Nudiamphisiella (2 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 560 Erimophrya (4 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 577 Vermioxytricha (2 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 596 Hemiurosoma (4 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 614
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Supplement to the Urostyloidea (2 species). . . . . . . . . . . . . . . . . . . . . . . . . . 636 Supplemented Key to Anteholosticha species. . . . . . . . . . . . . . . . . . . . . . . 636 Supplement to Anteholosticha (2 species). . . . . . . . . . . . . . . . . . . . . . . . . . 640 Supplement to the Oxytrichidae (3 species). . . . . . . . . . . . . . . . . . . . . . . . . 651 Amphisiellides (1 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 651 Pseudouroleptus (1 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 658 Ponturostyla (1 species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 672 Addenda. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 683 References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 695 Systematic Index. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 723 Table Index. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 737
Preface The present book is a monograph about two groups of hypotrichous ciliates, namely the Amphisiellidae and the Trachelostylidae. It is the third of six volumes which review the Hypotricha, one of the three major taxa of the spirotrichs. The first volume is about the Oxytrichidae, a rather large group, many species of which have 18 highly characteristically arranged frontal-ventral-transverse cirri and, much more importantly, a comparatively complex dorsal ciliature due to (oxytrichid) fragmentation of dorsal kineties during cell division (Berger 1999). The second volume deals with the Urostyloidea, which are characterised by a zigzag-arrangement of the ventral cirri (Berger 2006). Although this pattern is often very impressive, it is a relatively simple feature originating by a more or less distinct increase of the number of frontal-ventral-transverse cirri anlagen. These anlagen produce cirral pairs which are serially arranged in non-dividing specimens. Some users are likely astonished that the monograph on urostyloids does not include Uroleptus, a group of tailed species, which also have a distinct zigzagging cirral pattern. However, morphological and molecular data indicate that the zigzag pattern of Uroleptus evolved independently, that is, convergently to that of the urostyloids. Thus, Uroleptus was excluded from the urostyloid review. A zigzag pattern is also known from some oxytrichids, for example, Neokeronopsis, Territricha, Pattersoniella, showing that this pattern evolved several times independently (Berger 1999, 2006, Foissner et al. 2004). The present volume reviews the Amphisiellidae and the Trachelostylidae. The Amphisiellidae are characterised by a more or less distinct ventral file, termed amphisiellid median cirral row. Amphisiellids produce their frontal-ventral-transverse cirri from six anlagen (I–VI), a feature taken over from the ground pattern of the hypotrichs. The amphisiellid median cirral row usually originates from the two rightmost anlagen (V and VI); in some taxa, anlage IV forms the middle portion of the row. The anterior portion is formed from anlage VI and can therefore be easily homologised with the frontoterminal cirri of the 18-cirri hypotrichs. The amphisiellids have, like the urostyloids and some other taxa, taken over the simple dorsal ciliature from the ground pattern of the hypotrichs, that is, they basically have three bipolar kineties which divide by intrakinetal proliferation. There is of course some variation in the number of dorsal kineties within the amphisiellids. In some “amphisiellid” taxa, the formation of the dorsal kineties is not known, or they do not form a distinct amphisiellid median cirral row. They are preliminarily classified as incertae sedis in the amphisiellids, unless I could find a more parsimonious solution. Few species previously assigned to the amphisiellids have dorsomarginal rows, that is, dorsal kineties which originate from/near the anterior end of the right marginal row primordia. Dorsomarginal rows are characteristic for the oxytrichids, but also for Uroleptus and some other taxa. To include these taxa, the Dorsomarginalia have been established (Berger 2006, p. 38). Consequently, the few “amphisiellids” xi
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which possess dorsomarginal kineties are very likely misplaced in this group. Since their true position is not yet known, they are treated in the present book as nonoxytrichid Dorsomarginalia. Few “amphisiellids” very likely have a dorsal kinety fragmentation characteristic for the oxytrichids, namely Pseudouroleptus and Amphisiellides. Thus, they are treated in a supplement to the oxytrichids. Trachelostyla, the eponymous type of the Trachelostylidae, has been assigned to various higher taxa. It is an 18-cirri hypotrich which lacks dorsomarginal rows, but shows a multiple fragmentation in dorsal kinety 1 and forms two complete bipolar kineties from kinety 6 (Shao et al. 2007). The phylogenetic positions estimated from molecular date are varying (Schmidt et al. 2007, Shao et al. 2007), but indicate that Trachelostyla branched off rather early in the hypotrich tree. The trachelostylids are a small marine group possibly related to Gonostomum-like hypotrichs – a mainly terrestrial group previously mistakenly assigned to the oxytrichids (Berger 1999) – because the oral apparatus is similar and the postoral ventral cirri are displaced anteriad. Amphisiellids and the other taxa reviewed in the present volume are common only in marine and terrestrial habitats, that is, only very few species inhabit running waters, lakes, or ponds. The first and last detailed illustrated guide to these groups of hypotrichs was provided by Kahl (1932). Of course, Kahl’s book is outdated, especially as concerns the amphisiellids, because most species of this group are from soil and have been discovered mainly in the last five decades. Thus, it is not too early for a monographic treatment. As in the monographs on the oxytrichids and urostyloids, almost all available data on morphology, ontogenesis, ecology, and faunistics have been included. For each species, a detailed list of synonyms is provided, followed by a nomenclature section. In the remarks, all important data concerning systematics, synonymy, phylogeny, and similar taxa are considered. The morphology section contains a thorough description, following the same sequence in every species. If the data on various populations or synonyms do not agree very well, then they are kept separate so that even workers who do not agree with the synonymy proposed can use the revision. For several species, cell division data are available. They are also included because the ontogenesis is often very important to understand the interphasic cirral pattern correctly. The occurrence and ecology section contains a description of the type locality and all other localities where a species was recorded. In addition, almost all illustrations published so far have been included. Thus, with the present book the general microscopist need not refer back to the widely scattered original literature. Specialists, however, should always check both the present treatise and the original description or authoritative redescription when redescribing a known species. The most prominent and productive workers dealing with amphisiellids and trachelostylids are, in chronological order, Kahl, Foissner, Hemberger, Eigner, Song, and Hu. However, several other authors also wrote important papers on the
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alpha-taxonomy of these taxa. 59 amphisiellid, six trachelostylid, and 24 “other” species are treated as valid in the present revision. Details about synonymy rates will be provided in the last volume of the monographic series. The oxytrichids and the urostyloids are groups characterised by rather good apomorphies, like dorsal kinety fragmentation or the presence of a distinct midventral complex (Berger 1999, 2006). Other higher taxa, including the amphisiellids, are much more difficult to characterise, that is, the assignment of the non-oxytrichid and non-urostyloid species and genera to a certain higher level group is a difficult task, as indicated by the rather different classifications and molecular trees published so far. During the revision of the outstanding genera I will certainly find species which should have been treated in a previous volume. These species will be reviewed in supplements at the end of each book, as already done in Berger (2006) and the present revision. The last volume of the series will contain a key and a systematic index to all species so that the reader can find all hypotrichs very easily within the various volumes of the monographic series. The next group which will be treated in the monograph series are the Kahliellidae, also a moderately large taxon. Fortunately, the Austrian Academy of Sciences is sponsoring a major part of the series so that the monographic treatment of the Hypotricha can be completed in the foreseeable future. I hope that many ciliate-lovers benefit from the series on hypotrichs. Salzburg, April 2008
Helmut Berger
Acknowledgements and Permissions This book could not have been written without the assistance of many colleagues. I am especially gratefully to Wilhelm Foissner (University of Salzburg, Austria) for supplying original micrographs, faunistic data, and fruitful discussions; to Alois Lametschwandtner (head of the Department of Organismal Biology, University of Salzburg) for institutional support; to Martin Schlegel, Stefanie Schmidt, and Detlef Bernhard (University of Leipzig, Germany) for providing molecular biological data; to Erna Aescht (Upper Austrian Museum, Linz), Ulrich Buitkamp (Stawa Lippstadt, Germany), Horst Hemberger (Germany), Weibo Song (Ocean University Qingdao, P. R. China), and Norbert Wilbert (University of Bonn, Germany) for supplying information, data, and literature. Thanks to the staff at the Salzburg University Library, for bibliographic and interlibrary loan. Many thanks to Eric Strobl (Salzburg) for improving the English – I take full responsibility for any mistakes that remain. I also wish to acknowledge the generosity of the Springer Publisher, especially Tamara Welschot (Senior Publishing Editor Paleo-Environmental Sciences) and Judith Terpos (Senior Assistant to Publishing Editor), and the editor of the Monographiae Biologicae Henri J. Dumont (The State University of Ghent, Belgium), for printing the monographic series of the Hypotricha. The present volume is supported by a generous three-year research grant from the Austrian Academy of Sciences, Vienna (APART, Austrian Programme for Advanced Research and Technology; Project 10940), based on independent reviews. Many thanks to Peter Schuster (president of the Academy), Herbert Mang (former president), Lottelies Moser, Birgit Distler, Eva Gutknecht (Department for Grants and Awards), their colleagues from the Austrian Academy of Sciences, and to the anonymous reviewers. Eric Strobl (native speaker) was funded by the Stiftungs- und Förderungsgesellschaft der Paris-Lodron-Universität Salzburg (executive director Alfred Rinnerthaler). As in my other books, I have to thank my wife, Elisabeth, and my daughters, Magdalena, Eva, and Helena, for giving up time that belonged to them. The figures are either originals or reproductions from the literature of the past 120 years. My sincere thanks to the following publishers and authors who freely granted permission to use published drawings and photographs: Alekperov Ilham: An atlas of free-living ciliates - Publishing House Borcali, Baku. Asociación Latinoamericana de Microbiología, Cuernavaca (http://www.medigraphic. com/espanol/e-htms/e-lamicro/em-mi.htm): Revista Latinoamericana Microbiología. Cambridge University Press, Cambridge (http://www.cambridge.org): Bulletin of the British Museum of Natural History. Chinese Academy of Sciences, Beijing (http://zss.ioz.ac.cn): Acta Zootaxonomica Sinica. xv
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ACKNOWLEDGEMENTS AND PERMISSIONS
Duncker & Humblot GmbH, Berlin (http://www.duncker-humblot.de): Zoologische Beiträge. Elsevier, Amsterdam (http://www.elsevier.com): Archiv für Protistenkunde; European Journal of Protistology; Zoologische Jahrbücher Anatomie; Zoologische Jahrbücher Systematik; Zoologischer Anzeiger. Fernandez-Leborans Gregorio: Proceedings of the Biological Society of Washington, vol. 105 (1992), pp. 165–179. Hemberger Horst: Revision der Ordnung Hypotrichida Stein (Ciliophora, Protozoa) an Hand von Protargolpräparaten und Morphogenesedarstellungen - Dissertation University of Bonn, Bonn. Hungarian Academy of Sciences, Balaton Limnological Research Institute, Tihany (http://tres.blki.hu/BLRI.htm): Acta Biologica Hungarica. Instituto de Biologia, UNAM, Coyoacán (http://www.ibiologia.unam.mx): Anales del Instituto de Biologia de la UNAM, Series Botánica y Zoologia; Cuadernos del Instituto de Biologia. Naturhistorischer Verein der Rheinlande und Westfalens E.V., Bonn (http:// www.nhv.uni-bonn.de): Decheniana. Nencki Institute of Experimental Zoology, Polish Academy of Sciences, Warszawa (http://www.nencki.gov.pl/ap.htm): Acta Protozoologica. Oberösterreichisches Landesmuseum Biologiezentrum, Linz (http://www.biologie zentrum.at): Denisia; Stapfia. Scientific and Technical Research Council of Turkey (http://journals.tubitak.gov. tr/zoology/index.php): Turkish Journal of Zoology. Springer Science and Business Media, Berlin (http://www.springer.com): Biodiversity and Conservation; Biology and Fertility of Soils; Carey - Marine Interstitial Ciliates. An illustrated key; Hydrobiologia. Station Biologique de Roscoff, Roscoff (http://www.sb-roscoff.fr/BibDoc/travaux. php): Travaux de la Station Biologique de Roscoff. Taylor & Francis Group Ltd, Oxford (http://www.tandf.co.uk/journals): Journal of Natural History (London). Wiley-Blackwell, Hoboken, Malden (http://www.wiley.com; http://www.blackwell publishing.com): The Journal of Eukaryotic Microbiology (previously The Journal of Protozoology). Specific acknowledgements are made in the list of synonyms and the figure legends where the authors of the papers and the journals/books, in which the illustrations originally appeared, are named. All sources are cited in the reference section.
A General Section In the following chapters the general external and internal morphology of the hypotrichs treated in the present volume and terms specific to these taxa are described and explained. The focus is of course on the Amphisiellidae (= amphisiellids)1 because their cirral pattern deviates most obviously from that of the 18-cirri hypotrichs (Fig. 2a–c). For terminology relating to the Trachelostylidae, which are 18-cirri hypotrichs, and other taxa reviewed in the present book, see Fig. 4a, b and Berger (1999, 2006). However, all illustrations of the individual species described in the systematic section are labelled in great detail so that even inexperienced workers will understand the morphology of hypotrichs easily. For explanation of other terms, see Corliss (1979), Corliss & Lom (1985, 2002), Lynn & Corliss (1991), Hausmann & Bradbury (1996), Hausmann & Hülsmann (1996, 1996a), and Hausmann et al. (2003). Further information about the morphology can be found in the phylogeny chapter, which discusses the ground pattern of the hypotrichs, that is, I tried to reconstruct the last common ancestor of the hypotrichous ciliates (Fig. 7a, b). The ground pattern of the amphisiellids and trachelostylids is explained in the systematic section. Other topics, for example, ecology and distribution, life cycle, are briefly discussed in chapters 2–7.
1
Morphology, Biology, and Terminology
1.1 Size and Shape Amphisiellids are usually medium-sized hypotrichs, that is, the majority is between 100 µm and 200 µm long. The ratio of body length to body width ranges from 3–4:1 (e.g., Amphisiella annulata; Fig. 17g) to about 9:1 in Lamtostyla procera (Fig. 33a). Thus, the body outline is basically elliptical to elongate elliptical. A relatively high percentage of species is narrowed tail-like posteriorly or almost vermiform. The ventral side of the amphisiellids is, as in most other hypotrichs, usually flat, the dorsal side more or less distinctly vaulted (Fig. 1b, 2c). The body is flexible (supple) and usually acontractile or only slightly contractile. No rigid species is described. In the Hypotricha, a rigid cortex/body is only known from the Stylonychinae (for review, see Berger 1999, p. 499) and some species of uncertain phylogenetic position, for example, Rigidothrix Foissner & Stoeck, 2006 or Urospinula Corliss, 1960 (Foissner 1983b). The adoral zone of membranelles, the most prominent part of the oral apparatus, is, as is usual, in the left anterior body portion and usually less than 40% of body length, in many species around 30%. For some general terms used in the descriptions, see Fig. 1a–c.
1
For names of higher taxa used in this book, see Fig. 6a, 9a and Table 3.
1
2
GENERAL SECTION
MORPHOLOGY
3
1.2 Nuclear Apparatus The species in the present volume have an ordinary nuclear apparatus, that is, two or several macronuclear nodules and one or more micronuclei (Fig. 1a, 2b, c, 3b, Table 1). A considerable number of species has many small macronuclear nodules. Usually, the nuclear apparatus is left of body midline (Fig. 1a). As in other ciliate species, the nuclear pattern is very important for species identification. The macronucleus is – as in most other ciliates – homomerous and polyploid. Homomerous means that there is no distinct differentiation into DNA-rich and DNA-poor parts, as is the case in the heteromerous macronuclei characterising groups like the Chlamydodontidae and Dysteriidae (Raikov 1969). For detailed reviews on the nuclear apparatus of hypotrichs and ciliates in general, see Raikov (1969, 1982 1996), Klobutcher & Prescott (1990), and Prescott (1994, 1998). The development of the amphisiellid nuclear apparatus during ontogenesis is the same as in many other hypotrichs. The micronuclei divide mitotically, whereas the fused macronucleus makes one or more rapid, successive amitotic divisions to produce the species-specific number of nodules in each filial product (Prescott 1994). Of course, the macronuclear nodules of the amphisiellids posses a replication band, a feature which evolved in the stem-line of the spirotrichs. For documentation of the division of the nuclear apparatus in an amphisiellid, see, for example, Lamtostyla australis (Fig. 31h, j, t, v, x, y, z; Voß 1992).
1.3 Contractile Vacuole and Cytopyge The contractile vacuole is, as is usual for the hypotrichs, near the left cell margin at about 40–50% of body length or somewhat ahead of it; usually it is not ahead of the level of the proximal end of the adoral zone of membranelles (Fig. 1c, 7b). In some species (e.g., Lamtostylides edaphoni, Lamtostyla perisincirra) it is displaced somewhat inwards (Fig. 64b, 39c). Many species have distinct collecting canals extending
b
Fig. 1a–h Schematic illustrations to explain some general terms used in the species descriptions (original). a: Ventral view. Asterisk marks buccal cavity. b: Left lateral view. c: Dorsal view. d: The cirri of a true row or row are formed by the same anlage. Examples: marginal row; anterior portion of amphisiellid median cirral row. e: The cirri of a pseudorow are formed by different anlagen. Examples: transverse cirri; longitudinal rows formed by midventral pairs of urostyloids (previously, these two rows where designated as midventral rows; for new urostyloid terminology, see Berger 2006). f: A mixed row is basically a pseudorow made of true rows. Example: amphisiellid median cirral row. g, h: To symbolise the origin of cirri, respectively, cirral rows two different auxiliary lines are used. A = distal (= frontal = collar) portion of adoral zone of membranelles, AZM = adoral zone of membranelles, B = proximal (= ventral = lapel) portion of adoral zone of membranelles, C = gap in adoral zone of membranelles (only present in some taxa), CC = caudal cirri, CV = contractile vacuole with collecting canals, DE = distance between anterior body end and distal end of adoral zone, E = endoral, MA = macronuclear nodules, MI = micronucleus, P = paroral, 1, 2, 3 = dorsal kineties with bristles (1 = leftmost kinety; kineties not shown in full length in [a, c]).
4
GENERAL SECTION
Table 1 Nuclear apparatus of amphisiellid and trachelostylid ciliates and other species reviewed in this monograph Nuclear apparatus
Species a
Two macronuclear nodules; two or more micronuclei or number of micronuclei not known
Afroamphisiella abdita; Amphisiella annulata (Fig. 17b, g); Amphisiella australis sensu Foissner (Fig. 32a, f); Amphisiella capitata (Fig. 16c); Amphisiella milnei (Fig. 19a); Amphisiella oscensis (Fig. 24a); Apourosomoida halophila (Fig. 108a, e); Apourosomoida natronophila (Fig. 112a); Bistichella namibiensis (Fig. 114a, d); Bistichella procera (Fig. 116a, d); Bistichella terrestris (Fig. 117a, b); Caudiamphisiella antarctica (Fig. 23a, g); Erimophrya arenicola (Fig. 122a, f); Erimophrya glatzeli (Fig. 121a, c); Erimophrya sylvatica (Fig. 123a, c); Hemiamphisiella granulifera (Fig. 61a, e); Hemiamphisiella wilberti (Fig. 60a, g); Hemiurosoma goertzi (Fig. 131a, f); Hemiurosoma similis (Fig. 132b); Lamtostyla australis (Fig. 31a, e); Lamtostyla decorata (Fig. 41a, c); Lamtostyla granulifera (Fig. 40a, l); Lamtostyla islandica (Fig. 37a, c); Lamtostyla lamottei (Fig. 30a, b); Lamtostyla longa (Fig. 43a, b); Lamtostyla procera (Fig. 33a, c); Lamtostyla raptans (Fig. 44a, b); Lamtostylides edaphoni (Fig. 64a, e); Lamtostylides halophilus (Fig. 66a, c); Lamtostylides hyalinus (Fig. 69a); Lamtostylides kirkeniensis (Fig. 65a, e); Lamtostylides pori (Fig. 68a, b); Maregastrostyla pulchra (Fig. 26a, h); Nudiamphisiella illuvialis (Fig. 120a, c); Nudiamphisiella interrupta (Fig. 119a, h); Pseudouroleptus caudatus (Fig. 136a, b, d); Spiroamphisiella hembergeri (Fig. 27a, e); Spirotrachelostyla simplex (Fig. 106a, b); Spirotrachelostyla tani (Fig. 105a, l); Stichochaeta pediculiformis sensu Kahl (1928; Fig. 103a); Tetrastyla oblonga (Fig. 95a); Trachelostyla rostrata (Fig. 102a, b); Trachelostyla sp. sensu Kahl (Fig. 76e); Uroleptoides binucleatus binucleatus (Fig. 50a, b, h); Uroleptoides binucleatus multicirratus (Fig. 51a, f); Uroleptoides magnigranulosus (Fig. 52a, c); Uroleptoides polycirratus (Fig. 55a, d); Uroleptoides terricola (Fig. 54b, d)
Two macronuclear nodules and one micronucleus in between
Cossothigma dubium b (Fig. 76a–d); Lamtostyla perisincirra (Fig. 39a–f); Lamtostylides hyalinus (Fig. 69d); Mucotrichidium hospes (Fig. 91a, j)
Four macronuclear nodules (in some species the nodules are arranged in pairs)
Afroamphisiella abdita (Fig. 75a, c); Amphisiella turanica (Fig. 20a); Bistichella buitkampi (Fig. 113a, f); Erimophrya quadrinucleata (Fig. 125a, b, g); Hemiamphisiella quadrinucleata (Fig. 62a, e); Hemisincirra namibiensis (Fig. 82a, c–p); Hemisincirra quadrinucleata (Fig. 83a, b); Hemiurosoma terricola (Fig. 130a–c); Lamtostyla elegans (Fig. 36a, b); Lamtostyla quadrinucleata (Fig. 35a, e); Lamtostyla vitiphila (Fig. 34a, d); Ponturostyla enigmatica (Fig. 138b, f, k); Terricirra matsusakai (Fig. 93a, e); Uroleptoides longiseries (Fig. 46a, e)
Eight macronuclear nodules
Anteholosticha verrucosa (Fig. 134a, c); Hemisincirra gellerti (Fig. 85a); Hemisincirra octonucleata (Fig. 84a, b); Hemiurosoma polynucleatum (Fig. 132a); Terricirra viridis (Fig. 92a, d); Uroleptoides raptans (Fig. 47a, b)
More than eight macronuclear nodules
Afroamphisiella multinucleata (Fig. 74a, j); Amphisiella ovalis (Fig. 21a); Amphisiellides atypicus (Fig. 135a, b); Anteholosticha hetero-
#
MORPHOLOGY
5
Table 1 Continued Nuclear apparatus
Species a
More than eight macronuclear nodules
cirrata (Fig. 133a, b), Bistichella humicola (Fig. 118a); Hemiamphisiella terricola qingdaoensis (Fig. 59a, d); Hemiamphisiella terricola terricola (Fig. 56a, f); Hemisincirra buitkampi (Fig. 77a, b); Hemisincirra gellerti (Fig. 85g); Hemisincirra inquieta (Fig. 78a, b, 79a, f, 80a); Hemisincirra interrupta (Fig. 87a, c); Hemisincirra rariseta (Fig. 86a, b); Hemisincirra vermicularis (Fig. 88a, b); Hemisincirra wenzeli (Fig. 89a, g); Paramphisiella acuta (Fig. 70a, e); Paramphisiella caudata (Fig. 71a, b); Spirotrachelostyla spiralis (Fig. 104a, b); Terricirra livida (Fig. 94a, e); Trachelostyla caudata (Fig. 101a); Trachelostyla pediculiformis (Fig. 96e, h); Uroleptoides kihni (Fig. 45a); Uroleptoides multinucleatus (Fig. 48a, g); Vermioxytricha arenicola (Fig. 126a, m); Vermioxytricha muelleri (Fig. 129a, d)
a
For details of the nuclear apparatus, see individual descriptions. Note that the nuclear apparatus is not illustrated for all species.
b
Single micronucleus in between macronuclear nodules uncertain (see text).
near the left body margin during diastole (e.g., Fig. 48a). For many marine species (e.g., some Amphisiella spp.) or taxa adapted to very high salinities (e.g., Apourosomoida spp.) no contractile vacuole is described or illustrated, possibly because it is lacking. However, it is known that in marine species the vacuole contracts in rather long intervals so that one cannot exclude that this organelle has sometimes been overlooked. The excretory pore is, likely as in the other hypotrichs, on the dorsal side above the contractile vacuole. The cytopyge of the amphisiellids is a little-known organelle which is, as is usual, located in the posterior body portion near the left cell margin (e.g., Fig. 62b, 66e).
1.4 Cytoplasm, Cortex, and Colouring The cytoplasm of the amphisiellids and the other species reviewed here is more or less colourless and contains the ordinary inclusions, for example, greasily shining globules, rod- and/or Y-shaped cytoplasmic crystals, and food vacuoles. However, note that some species are coloured due to cortical granules (see chapter 1.5). Some species (e.g., Amphisiella annulata and A. milnei) have ring-shaped structures (“hollow” spheres? lithosomes?) in the cytoplasm, making these species rather easily determinable (Fig. 17b, g, 19a). Likely the same structures occur in the anterior and posterior body portion of Tachysoma pellionellum (see Fig. 135c, d, 136i in Berger 1999). Species with symbiotic algae are not known. The cortex of the amphisiellids is supple, that is, the body is flexible when freely motile. Consequently, it is very unlikely that your specimen/population belongs to a
6
GENERAL SECTION
MORPHOLOGY
7
species described in the present book if its body is rigid and moves like a board when freely swimming; if you find such a specimen/population you have to look at the stylonychines (Berger 1999, p. 499). No study about the ultrastructure of amphisiellids is available.
1.5 Cortical Granules Cortical granules occur in a relatively high number of amphisiellids (Table 2). Their size, shape, colour, and arrangement are very important features, which cannot usually be seen after protargol impregnation. Consequently, live observation is absolutely necessary for a reliable identification of a hypotrich (e.g., Stein 1859, Berger & Foissner 1987a, Foissner et al. 2002, Hu et al. 2004). Note that the “correct” colour can only been seen at well-adjusted bright-field illumination; the presence or absence of cortical granules should be checked with differential interference contrast and by staining with methyl-green pyronin.
1.6 Movement The taxa in this book are – like the vast majority of the other hypotrichs – usually thigmotactic, that is, they adhere more or less strongly to the substrate whenever the opportunity arises. They creep on their flattened ventral side by means of the cirri. Usually, the specimens move to and fro rather hastily. All amphisiellids have a supple body which bends to varying degrees. Thus, when you see a rigid, freely motile
b
Fig. 2a–c Terminology of amphisiellid ciliates (figures from Berger 2004). Infraciliature, that is, cirral and kinety pattern after protargol impregnation of Amphisiella annulata, a typical amphisiellid. a: Ventral side showing, inter alia, oral apparatus – composed of adoral zone of membranelles, undulating membranes (endoral and paroral), and cytopharynx –, cirral pattern with the amphisiellid median cirral row (ACR), the main morphological apomorphy of the amphisiellids, and the two rightmost dorsal kineties. Long arrow marks buccal vertex, short arrow marks a supernumerary transverse cirrus, a characteristic feature of A. annulata. Double arrow marks area where the anterior portion and the posterior portion of the amphisiellid median cirral row fuse so that a continuous, mixed row is formed. Broken lines connect cirri which originate from same anlage (only shown for anlagen I–IV). The dimension arrowheads show how the length of the adoral zone is measured. b: Dorsal side showing bipolar dorsal kineties and nuclear apparatus. c: Left lateral side showing, inter alia, nuclear apparatus and adoral zone. Note that this specimen is almost not flattened dorsoventrally. ACR = anterior and posterior end of amphisiellid median cirral row, AZM = adoral zone of membranelles (in [a] AZM marks the distal end of the zone), BC = buccal cirrus (= cirrus II/2), DB = dorsal bristles (= dorsal cilia), FC = right frontal cirrus (= cirrus III/3; forms a pseudorow with middle and left frontal cirrus as indicated by dotted line), LMR = left marginal row, MA = macronuclear nodule, MI = micronuclei, PF = pharyngeal fibres, PTVC = pretransverse ventral cirri (cirri V/2 [left] and VI/2 [right]; = PT in Berger 2006), RMR = right marginal row, TC = transverse cirri (form hook-shaped pseudorow in this species), I–IV = frontal-ventral-transverse cirri anlagen, III/2 = cirrus “behind” right frontal cirrus, 1–6 = dorsal kineties (1 = leftmost kinety, 6 = kinety closest to right marginal row).
8
GENERAL SECTION
hypotrich you can exclude that it is an amphisiellid or other species described here. No detailed studies on the movement of amphisiellids exists. Table 2 Amphisiellidae and other species with cortical granules. Species arranged alphabetically Species
Granules Size (in µm)
Shape
Colour
Arrangement and Remarks a, b
Afroamphisiella multinucleata (Fig. 74b, c, m, n)
1.0 × 0.5
ellipsoidal
yellowish, highly refractive
in longitudinal, rather widely spaced rows; do not impregnate with protargol, become red when MGP is added (details see text)
Amphisiella annulata (Fig. 17g, l, 18d)
0.8–1.0 and 0.3–0.5
globular
colourless
larger one in patches between dorsal bristles; smaller one throughout cortex
Amphisiella milnei (Fig. 19a)
two size classes, but size not indicated
both globular?
smaller ones colourless; larger ones yellowish
smaller granules are dustshaped protrichocysts; larger granules loosely arranged between smaller granules
Caudiamphisiella 10 antarctica (Fig. 23b)
globular
colourless
sparsely distributed on dorsal side
Hemiamphisiella granulifera (Fig. 61b, c)
1.0–2.0
globular
colourless
arranged in longitudinal rows
Hemiamphisiella quadrinucleata (Fig. 62b)
0.5
globular
colourless
arranged in longitudinal rows; stain red when MGP is added
Hemiamphisiella terricola terricola (Fig. 56c, 57c)
1.4
globular
colourless; make cells brownish
irregularly distributed; impregnate with protargol
Hemisincirra gellerti (Fig. 85c)
1.5
bean-shaped
colourless
arranged in about 30 longitudinal rows
Hemisincirra inqui- 1.0 eta (Fig. 79b, g, 81e–h)
globular
bright yellow
around cirri and dorsal bristles
Hemisincirra wenzeli (Fig. 89c, d)
1.2 × 0.8
ellipsoidal
colourless
along cirral rows and dorsal bristles
Lamtostyla decorata (Fig. 41e–g, n)
0.3
globular
colourless
form conspicuous plaques around dorsal bristles and scattered around bases of cirri
Lamtostyla granulifera (Fig. 40d, h–k)
1.0–4.0, usually 2.0
globular
colourless, hyaline and bright
arranged in narrowly spaced rows; do not stain with MGP
Lamtostylides halophilus (Fig. 66e, 67e)
0.5–1.0
globular
colourless
loosely arranged; usually impregnate with protargol
MORPHOLOGY
9
Table 2 Continued Species
Granules Size (in µm)
Shape
Colour
Arrangement and Remarks a, b
<1.0
globular
likely colourless
arranged in short rows
Nudiamphisiella in- 0.5–1.0 terrupta (Fig. 119b)
globular
almost colourless
arranged in widely spaced rows; stain red with MGP
Terricirra livida (Fig. 94d)
1.5
globular
blue-green
arranged in short rows
Terricirra matsu1.0 sakai (Fig. 93c, f–h)
globular
dark green
about six longitudinal rows each on ventral and dorsal side
Terricirra viridis (Fig. 92c, i)
0.8
globular
dark green
arranged in short rows, mainly along cirral rows and dorsal kineties
Uroleptoides binucleatus binucleatus (Fig. 50f)
>1.0
globular
colourless
arranged around cirri and dorsal bristles; do not stain with MGP
Uroleptoides binucleatus multicirratus (Fig. 51d)
0.8–1.0 (around cirri); 0.5–1.0 and 3.0 (around bristles)
globular
colourless
arranged around cirri and dorsal bristles; occasionally impregnate with protargol
Uroleptoides longiseries (Fig. 46b, c)
0.5 (around cirri); 0.5 and up to 1.5 (around bristles)
globular
colourless
arranged around cirri and dorsal bristles; granules around bristles usually impregnate with protargol
Uroleptoides magnigranulosus (Fig. 52f–h, 53a–e)
<1.0 and up to 3.0
globular
colourless
arranged in clusters around cirri and dorsal bristles (details see text)
Uroleptoides multinucleatus (Fig. 48b–d)
0.8 (around cirri); 0.5 and 3.0 (around bristles)
globular
colourless
arranged around cirri and dorsal bristles; occasionally some impregnate with protargol
globular
yellowish
mainly around cirri and dorsal bristles
Maregastrostyla pulchra (Fig. 26b, c, g)
Vermioxytricha are- 1.0 nicola (Fig. 126h, i) a b
MGP = methyl-green pyronin. See text for a more detailed description of the cortical granulation.
10
GENERAL SECTION
1.7 Somatic Ciliature and Ultrastructure As in other hypotrichs, the somatic ciliature of the amphisiellids, trachelostylids, and other taxa reviewed in the present book consists of rows and localised groups of cirri on the flattened ventral side, and several (around six in Amphisiella; usually three in some other major taxa, e. g., Lamtostyla) rows of more or less widely spaced, usually short (2–5 µm), stiff cilia (bristles) on the vaulted dorsal side (Fig. 1a–c, 2a–c, 3a, b, 4a, b). Caudal cirri are, if present at all, part of the dorsal ciliature because formed at the end of bipolar kineties (Fig. 1b, c, 3b). A cirral row in hypotrichs belongs to one of the following three types, namely, (i) true rows or rows; (ii) pseudorows; or (iii) mixed rows (Fig. 1d–f). True row or row: all cirri of such a row originate from the same anlage. Examples are the marginal rows or the row of postoral ventral cirri in Apourosomoida halophila (Fig. 1d, 108e). Pseudorow: each cirrus of such a row is formed by a different anlage. Examples are the transverse cirri, the frontal cirri, the anterior or posterior bow of a bicorona, or the row formed by the left cirri of a midventral complex composed of cirral pairs (Fig. 1e). This term was introduced by Berger (2006, p. 11). Mixed row: this new term is introduced for “rows” like, for example, the amphisiellid median cirral row, which are composed of two or three more or less long and linearly arranged Fig. 3a–d Terminology of amphisiellid ciliates (figures from Foissner 1997 and Foissner 1984. Protargol impregnation). a: Infraciliature of Lamtostyla granulifera, an 18-cirri amphisiellid. In such 18-cirri amphisiellids, the amphisiellid median cirral row is composed of only four cirri (from posterior: V/3, V/4, VI/3, VI/4 [Fig. 3d]; the latter two cirri are also termed frontoterminal cirri); the cirri left of the anterior portion of the amphisiellid median cirral row are cirri III/2, VI/3, and IV/2 (this cirrus is homologous to the postperistomial cirrus). Note that this cirral pattern is identical to that of many 18-cirri oxytrichids, except that the postoral ventral cirri are behind the rear end of the adoral zone. Cirri which originate from the same anlage are connected by a broken line; cirri which form pseudorows are connected by dotted lines. The numbering system introduced by Wallengren (1900) for 18-cirri hypotrichs designates the frontal-ventral-transverse cirri anlagen (= streaks, primordia) with I–VI (from left to right) and the cirri within an anlage with 1–4 beginning from the rear. b: Infraciliature of dorsal side of Hemiamphisiella quadrinucleata. Long arrow marks the rear end of the distinctly shortened dorsal kinety 4; this shortening and the lack of a caudal cirrus at the end of the kinety strongly indicates that it is a dorsomarginal row (originates from/near anterior end of right marginal row primordium); however, this has to be verified by ontogenetic data; hypotrichs with a dorsomarginal kinety do not belong to the amphisiellids. Short arrow marks chromatin bodies (previously termed nucleoli). Caudal cirri originate at the rear end of bipolar dorsal kineties (broken lines). c: The last common ancestor of all hypotrichs very likely had 18 frontalventral-transverse cirri. d: Frontoventral and postoral ventral cirri of the specimen shown in (a) to demonstrate the amphisiellid median cirral row in an 18-cirri hypotrich. The cirri “left of the anterior portion of the amphisiellid median cirral row” are circled. AZM = adoral zone of membranelles, CC = caudal cirri (at end of bipolar kineties), BC = buccal cirrus (= cirrus II/2), E = endoral (endoral and paroral are the undulating membranes), FC = frontal cirri (= cirri I/1, II/3, III/3), FVC = frontoventral cirri (= cirri III/2, IV/3, VI/3, VI/4), LMR = anterior end of left marginal row (usually left of proximal portion/end of adoral zone), MA = macronuclear nodule, MI = micronucleus, P = paroral (paroral and endoral are the undulating membranes), PVC = postoral ventral cirri (= cirri IV/2, V/3, V/4; cirrus IV/2 is identical with the postperistomial cirrus of some other amphisiellids), PTVC = pretransverse ventral cirri (= cirri V/2, VI/2), RMR = right marginal row, TC = transverse cirri (= cirri II/1, III/1, IV/1, V/1, VI/1), I–VI = frontal-ventraltransverse cirri anlagen, 1–4 = dorsal kineties (kinety 4 is possibly a dorsomarginal kinety).
d
MORPHOLOGY
11
12
GENERAL SECTION
Fig. 4a, b Terminology of trachelostylid ciliates (illustration of infraciliature from Gong et al. 2006. Protargol impregnation). a: Infraciliature of ventral side of Trachelostyla pediculiformis, type of Trachelostyla and thus also type of the whole group. Cirri which (very likely) originate from same anlage are connected by a broken line. Ellipse encircles postoral ventral cirri, rhomboid encloses the frontoventral cirri. b: A second interpretation of the origin of the postoral ventral cirri. I–VI = frontal-ventral-transverse cirri anlagen.
true rows (Fig. 1f). Often the borders of the individual true rows forming a mixed row are (almost) not recognisable in morphostatic specimens; thus, ontogenetic data are needed to recognise the row type. Sometimes, however, the elements are clearly separated, for example, in the frontoventral row of Nudiamphisiella interrupta (Fig. 119g). The arrangement of cirri and dorsal kineties is a very important feature for the systematics. Consequently, as in other groups of hypotrichs, an unambiguous terminology is needed to understand the morphology of the amphisiellids and the other taxa treated (Fig. 1a–h, 2a–c, 3a, b, 4a, b, 5a, b). The paragraphs below describe the individual cirri and cirral groups. Many cirri of the various higher taxa of the hypotrichs (e.g., Urostyloidea, Oxytrichidae, Amphisiellidae, Trachelostylidae) can be homologised and therefore have, of course, the same designation in these taxa. A detailed discussion of the confusing terminology of some cirri is provided in the Monograph of the Oxytrichidae (Berger 1999). As in the first two volumes of the revision of the Hypotricha (Berger 1999, 2006), I use the well-established numbering system introduced by Wallengren (1900) to designate the individual cirri or cirral rows and the anlagen from which these cirri originate (Fig. 3a); however, note that this system was basically established for the characterisation of 18-cirri hypotrichs. In the following, the cirral groups and structures are explained in the same sequence as they are usually treated in the individual species descriptions. Frontal cirri (FC). These cirri are near the anterior end of the cell (Fig. 2a, 3a, 4a). All taxa discussed have – like the oxytrichids and many urostyloids – three more or less distinctly enlarged frontal cirri which usually form a slightly oblique pseudorow. They are undoubtedly homologous in all groups. The left frontal cirrus (= cirrus I/1) is usually ahead of the par-
MORPHOLOGY
13
oral. It is formed from the same anlage (= anlage I) as the undulating membranes during cell division. The middle cirrus is homologous to cirrus II/3 of, for example, the 18-cirri hypotrichs (Berger 1999) or urostyloids with three frontal cirri (Berger 2006). It is produced, like the buccal cirrus, from anlage II during ontogenesis. The right frontal cirrus (homologous to cirrus III/3) is usually behind/close to the distal end of the adoral zone of membranelles. Buccal cirrus (BC). This cirrus (= cirrus II/2) is usually right of the paroral (Fig. 3a); in some species it is ahead of the undulating membranes (Fig. 2a, 4a). For a discussion of the confusing terminology, see Berger & Foissner (1997) and Berger (1999). Almost all amphisiellid species have one buccal cirrus which is certainly the plesiomorphic state. All Bistichella species have more than one buccal cirrus (e.g., Fig. 113e, 116c). Usually, the buccal cirrus has an ordinary size. In some species, however, it is composed of two basal bodies/cilia only (Fig. 60c); then it is difficult to recognise in life because easily misinterpreted as paroral cilia. Parabuccal cirrus/cirri (PC; cirrus III/2). Usually, at least one parabuccal cirrus is present in amphisiellids. It is homologous to cirrus III/2 (preferred term in present and previous books), which is the cirrus behind the right frontal cirrus (Fig. 2a). In the 18-cirri hypotrichs, cirrus III/2 is part of the four frontoventral cirri, together with cirri IV/3, VI/3, and VI/4 (Fig. 3a; Fig. 6a in Berger 1999). Frontoventral cirri (FVC). This group comprises four cirri (III/2, IV/3, VI/3, VI/4) between the anterior portion of the right marginal row and the paroral. They are arranged in various patterns, usually in a V-shaped one (Berger 1999). In Hemiurosoma and Urosoma, cirrus III/2 is in front of the other three cirri, which thus form a roughly longitudinal row (Fig. 130f, v, 131h; Berger 1999). Another term for the frontoventral cirri VI/3 and VI/4 is frontoterminal cirri (see next entry). Frontoterminal cirri (FT). This term was introduced by Hemberger (1982, p. 11) for the frontoventral cirri VI/3 and VI/4 because they migrate to near the anterior body end. Borror & Wicklow (1983) thus designated these cirri, which never form primordia during morphogenesis, migratory cirri. In most amphisiellids not two, but more frontoterminal cirri are present; they form the anterior portion of the amphisiellid median cirral row (Fig. 2a, 5a). Frontoventral row. This is a general term for a cirral row formed from one or more frontal-ventral-transverse cirri anlagen. If such a row is formed from two or three anlagen then it is an amphisiellid median cirral row (see next entry), assuming that the other features (dorsal kinety pattern!) also show that the species is an amphisiellid. In discocephalids each cirrus of the frontoventral row is formed from a different anlage, that is, it is a pseudorow (Fig. 8c, d); Eigner & Foissner (1994, p. 244) therefore introduced the term discocephalid median cirral row. Amphisiellid median cirral row (ACR). This term was introduced by Eigner & Foissner (1994, p. 244) for the mixed cirral row (see above for explanation of the term “mixed row”; Fig. 1f) so characteristic for most amphisiellids. The amphisiellid median cirral row is formed at least by (true) rows originating from cirral anlage V (forms posterior portion) and anlage VI, which produces the anterior portion (Fig.
14
GENERAL SECTION
MORPHOLOGY
15
5a; Borror & Wicklow 1982, Wicklow 1983, Eigner & Foissner 1994). In some cases, the amphisiellid median cirral row comprises a third, middle portion which is formed by anlage IV (Fig. 58f). The individual parts of the amphisiellid median cirral row can be easily homologised with cirri of the 18-cirri hypotrichs; for example, the anterior portion corresponds to the frontoterminal cirri (cirri VI/3, VI/4) of the 18-cirri hypotrichs or urostyloids (Fig. 3a, d). However, since the number of cirri formed in anlagen V and VI is often distinctly higher than in 18-cirri hypotrichs, the homology is often not clearly recognisable. Usually, the amphisiellid median cirral row is rather homogenous, that is, it looks like a true row, and therefore morphogenetic data are needed to show that it is a mixed row. Only in few cases are the individual parts clearly recognisable because the elements are not in line. Note that even 18-cirri hypotrichs have an “amphisiellid median cirral row”: however, it is composed of only four cirri (from anterior; Fig. 3a, d): VI/4, VI/3, V/4, V/3. The first two cirri (= anterior portion of amphisiellid median cirral row) are the frontoterminal cirri, the posterior two cirri (= posterior portion) are the postoral ventral cirri formed by anlage V. Cirri left of anterior portion of amphisiellid median cirral row. All amphisiellids have one or more cirri left of the anterior portion of the amphisiellid median cirral row (Fig. 2a). This is not a specific feature of the amphisiellids with a distinct amphisiellid median cirral row, as it is also present in the 18-cirri hypotrichs. However, in these species cirri III/2 and IV/3 are part of the frontoventral cirri and cirrus IV/2 is part of the postoral ventral cirri; therefore, the present cirral group is less conspicuous than in the amphisiellids with a distinct amphisiellid row (Fig. 3a). Cirrus III/2 is usually present in all amphisiellids; only rarely, anlage III forms a second such cirrus. The other cirri of the present cirral group are formed by anlage IV, for example, in Amphisiella annulata (Fig. 5a). If only one cirrus is present left of the anterior end of the amphisiellid median cirral row, it is almost cer-
b
Fig. 5a, b Terminology of amphisiellid ciliates (figures from Berger 2004. Protargol impregnation). a: Infraciliature of ventral side of a late reorganiser of Amphisiella annulata. This illustrations very impressively shows the main morphological apomorphy, namely the amphisiellid median cirral row which is formed from two portions: (i) the front portion is made of the anteriorly migrating cirri (= frontoterminal cirri) of anlage VI, that is, the number of frontoterminal cirri increased from two (state in ground pattern of Hypotricha) two 12 in present specimen; (ii) the rear portion of the amphisiellid median cirral row is the anterior portion of anlage V; in 18-cirri hypotrichs this portion is composed of only two cirri, namely V/3 and V/4 (Fig. 3a, d). Only in very late reorganisers or dividers do the two portions fuse to the amphisiellid median cirral row (S-shaped arrow); when the reorganisation or division process is finished the amphisiellid median cirral row is a continuous, mixed cirral row (Fig. 2a). The so-called “cirri left of the anterior portion of the amphisiellid median cirral row” are circled. The marginal rows are formed, as is usual, within the parental rows. Arrow marks an additional transverse cirrus; this is a specific feature of A. annulata. b: Infraciliature of dorsal side and nuclear apparatus of a late divider of A. annulata. Within each parental dorsal kinety two anlagen occur (arrows). Note that amphisiellids do not form a dorsomarginal kinety an lack dorsal kinety fragmentation. MA = dividing macronucleus, MI = dividing micronucleus, I–VI = frontal-ventral-transverse cirri anlagen.
16
GENERAL SECTION
tain that the species lacks anlage IV. However, this only applies when the species also lacks the postperistomial cirrus or the postoral ventral cirri. Postperistomial cirrus (cirrus IV/2). This term was introduced by Eigner & Foissner (1994, p. 244) for an isolated cirrus close behind the buccal vertex, present in only a small number of amphisiellids (Fig. 56e). Ontogenetic data show that it is formed from anlage IV (Fig. 58e, f) and I have no doubt that this cirrus is homologous to cirrus IV/2 of the 18-cirri hypotrichs because both cirri have the same position and origin (Fig. 7a). Consequently, the presence of a postperistomial cirrus has to be interpreted as plesiomorphy because it is already present in the last common ancestor of the hypotrichs (see chapter 2). By contrast, the lack of this cirrus is a novelty. Unfortunately, it is not known whether this cirrus was lost only once during the evolution of the amphisiellids, or twice or several times independently. In 18cirri hypotrichs the postperistomial cirrus is one of the three postoral ventral cirri (Fig. 3a). Postoral ventral cirri (PVC). This term is commonly used for the cirri IV/2, V/3, and V/4, which are behind the proximal end of the adoral zone (Fig. 7a). In few taxa, for example, the trachelostylids, some Lamtostyla-species, or Gonostomum-like hypotrichs, the postoral ventral cirri are displaced anteriad right of the proximal portion of the adoral zone of membranelles (Fig. 3a, 4a). Perhaps this is the plesiomorphic state within the hypotrichs (see end of chapter 2.4). In some amphisiellids, only postoral ventral cirrus IV/2 is present behind the buccal vertex because the other two cirri (V/3, V/4) form, together with many other cirri of anlage V, the posterior portion of the amphisiellid median cirral row (Fig. 58e); in such cases it is usually designated as postperistomial cirrus (see above). Pretransverse ventral cirri (PTVC; PT in Berger 2006). This term was introduced by Berger & Foissner (1997) for two cirri of 18-cirri hypotrichs. Accessory transverse cirri is an older, synonymous term introduced by Wicklow (1981, p. 348). They are usually arranged immediately ahead of the transverse cirri V/1 and VI/1 and are designated V/2 and VI/2 according to Wallengren’s scheme (Fig. 3a). They are present in only a limited number of amphisiellids, for example, in Amphisiella annulata (Fig. 2a) or Lamtostyla granulifera (Fig. 3a). Trachelostylids also have these cirri because they are 18-cirri hypotrichs. In species with a reduced number of pretransverse ventral and transverse cirri they are often difficult to distinguish because it is not known which cirri have been lost; in addition, the size of cirri of these two groups is often rather similar. In such cases, ontogenetic data are needed for a correct designation. Transverse cirri (TC). These cirri, which often form a distinct pseudorow, are usually in the posterior quarter of the cell (Fig. 2a, 3a, 4a). Transverse cirri are present in most hypotrichs and therefore also in most amphisiellids and other species reviewed in the present book. A transverse cirrus is, per definition, the rearmost cirrus produced by a frontal-ventral-transverse cirri anlage (Fig. 3a, 5a). It forms – usually together with other rearmost cirri – a “transverse” pseudorow which is typically more or less obliquely arranged. They often form a hook-shaped, or in some Lamto-
MORPHOLOGY
17
styla species U-shaped pattern (Fig. 41b). In many hypotrichs (e.g., Amphisiella, Stylonychia), the transverse cirri are rather prominent because they are distinctly larger than the other cirri nearby. In many amphisiellids, however, they have about the same size as, for example, marginal and pretransverse ventral cirri and therefore the correct designation is often difficult, especially when they are reduced in number and exactly in the gap formed by the rear end of the marginal rows. In tailed species it is sometimes also difficult to distinguish transverse cirri from caudal cirri, which are, however, located on the dorsal side (Fig. 1c). The loss of transverse cirri commences, as is usual, from left to right, that is, a species with, for example, only one transverse cirrus forms it from anlage VI, and not, for example, from anlage II or III. Whether or not transverse cirri are completely lacking is often difficult to tell and usually needs ontogenetic data, especially when the posterior body portion is narrowed. True, longitudinal rows between the rear portion of the marginal rows should not be designated as transverse cirri (Fig. 56e). 18-cirri hypotrichs. The last common ancestor of the Hypotricha very likely had 18 frontal-ventral-transverse cirri arranged in a relatively highly characteristic pattern and originating from six (I–VI) anlagen (Fig. 3a, c; see chapter 2). Consequently, this pattern occurs at just about all sites of the Hypotricha tree. Eigner (1997, p. 553) supposed that this pattern has evolved several times independently. However, this is almost impossible because the pattern, including its formation, is too complex to evolve convergently. Marginal cirri (RMR, LMR). These cirri run along the left and right body margin. All species discussed here have one left and one right marginal row (Fig. 2a, 3a, 4a), except for the marine Spiroamphisiella hembergeri, which has two right marginal rows (Fig. 27a, d, g, 29d). Marginal rows are true cirral rows because all cirri originate from the same anlage. The right marginal row often commences near the distal end of the adoral zone; in some species it is distinctly shortened anteriorly or it extends onto the dorsolateral surface. The left row usually begins left of the proximal portion of the adoral zone. In most species the marginal rows are slightly shortened posteriorly, that is, they do not extend to the posterior tip of the cell so that the rows are distinctly separated posteriorly. However, the gap is sometimes difficult to recognise because it is seemingly occupied by the caudal cirri, which, however, insert on the dorsal side. Dorsal cilia (DB; 1, 2, 3, ...). The dorsal side of all hypotrichs and euplotids is covered by a more or less high number of kineties, which are therefore named dorsal kineties or dorsal bristle rows (e.g., Fig. 2a–c, 3b). Many amphisiellids have, like many urostyloids, three bipolar kineties, which is likely the state in the stem line of the hypotrichs. Amphisiella, a marine group, has about six kineties (Fig. 2b, c). The bristle rows of the amphisiellids are basically bipolar, that is, they extend from near the anterior to near the posterior body end. Dorsomarginal kineties, which originate from/near the right marginal primordium, and fragmenting kineties (one [usually kinety 3] or more kineties fragment into an anterior and [usually one, sometimes more] posterior portion) are lacking in the amphisiellids. The fragmentation is characteris-
18
GENERAL SECTION
tic for the oxytrichids1 (for review, see Berger 1999), whereas dorsomarginal kineties are probably the main morphological apomorphy of the Dorsomarginalia (Berger 2006). Thus, I exclude “amphisiellids” with a dorsomarginal kinety (e.g., Nudiamphisiella interrupta, Fig. 119i) from the Amphisiellidae, and place them as incertae sedis in the Dorsomarginalia. Pseudouroleptus – according to the ventral cirral pattern a typical amphisiellid – has very likely an oxytrichid kinety fragmentation (Fig. 136c). Therefore, I included it in the monograph of the Oxytrichidae (Berger 1999, p. 888; see also present book). So far, the importance of the dorsal kinety pattern in elucidating the phylogeny of the hypotrichs has been underestimated. New data indicate that groups based on dorsal features are confirmed by molecular data. Neokeronopsis spectabilis (Kahl, 1932) Warren, Fyda & Song, 2002, a typical “urostyloid” hypotrich according to the “misleading” (in present case) ventral cirral pattern, can be easily recognised as oxytrichid by the presence of the oxytrichid dorsal kinety fragmentation (see Berger 2006, p. 1190ff). The two higher taxa based on features of the dorsal infraciliature are the Oxytrichidae Ehrenberg, 1831 (with [oxytrichid] dorsal kinety fragmentation), a subgroup of the Dorsomarginalia Berger, 2006 (dorsomarginal kinety present). Consequently, the exact description of the dorsal kinety pattern is an absolute prerequisite for a serious classification of a hypotrich.2 Unfortunately, ontogenetic data are often needed to know whether or not dorsomarginal rows and/or kinety fragmentation are present. Caudal cirri (CC). These cirri originate at the posterior end of the bipolar dorsal kineties, that is, the caudal cirri are part of the dorsal infraciliature (Fig. 1c). Dorsomarginal kineties, present in some species previously classified in the amphisiellids, are never associated with caudal cirri (Berger 1999), that is, in amphisiellids the number of caudal cirri (if present at all) is usually equal to the number of dorsal kineties, assuming that only one cirrus per kinety is formed. However, in some species only a part of the bipolar kineties form a caudal cirri; thus, fewer caudal cirri than dorsal kineties does not always indicate that a dorsomarginal row and/or kinety fragmentation are/is present (e.g., Paramphisiella caudata, Fig. 73n). The caudal cirri are always arranged at, or very close to, the rear body end, frequently above the gap formed by the rear end of the marginal rows. Thus, live and silver preparations must be studied with care to avoid a misinterpretation of caudal cirri as marginal or transverse cirri or vice versa. In vermiform species it is sometimes impossible to decide whether cirri at the rear cell end are caudal cirri or transverse cirri; in such cases, ontogenetic data are needed for a correct interpretation. The caudal cirri of the amphisiellids are nearly always inconspicuous, that is, neither very long and/or strong. The presence of caudal cirri is a plesiomorphy in the amphisiellids because they are already present in the stem line of the hypotrichs. Many taxa lack caudal cirri (e.g., 1
Note that the fragmentation present in Apourosomoida halophila (Fig. 110q) is very likely not homologous to the oxytrichid fragmentation. Recently, Shao et al. (2007) described a multiple fragmentation in kinety 1 of Trachelostyla pediculiformis; however, dorsomarginal rows are lacking making the estimation of the phylogenetic position of this marine species rather difficult (details see description of species). 2 For the determination of hypotrichs it is usually not necessary to know the exact dorsal kinety pattern.
MORPHOLOGY
19
Lamtostyla, Uroleptoides); very likely the loss, a rather simple feature, occurred several times independently. Fine structure of cirri and membranelles. There is no paper available dealing with the ultrastructure of the somatic ciliature of species reviewed. Likely the ultrastructure is rather similar to that of oxytrichids and urostyloids (for reviews, see Berger 1999, 2006).
1.8 Oral Apparatus The oral apparatus is composed, as in the remaining hypotrichs, of an adoral zone of membranelles, two undulating membranes (paroral and endoral), the buccal cavity (buccal field, oral field), and associated fibres including the cytopharynx. For a detailed characterisation and terminology of the various oral types present in the hypotrichs, see Berger & Foissner (1997), Berger (1999, 2006), and Foissner & ALRasheid (2006). Likely, details of the oral apparatus will provide further differences among various taxa. However, these features are generally very sophisticated and known only for a very low number of species. The adoral zone of membranelles, the most prominent part of the oral apparatus, extends from the anterior body end along the left body margin to near midline of the cell and usually terminates at about 20–35% of body length. Very often it is roughly the shape of a question mark and the distal end does not extend far posteriorly, that is, the so-called DE-value1 is often less than 0.11 (Berger 2006, p. 18). Some amphisiellids and other species treated here have a more or less distinct gap (break) in the adoral zone at the left anterior body corner (e.g., Fig. 1a, 74i, k). The anterior, often transversely arranged portion is termed distal, frontal, or collar portion and thus largely on the dorsal side of the frontal scutum, the anteriormost part of the body (Fig. 17g–i); the posterior part of the zone is termed proximal, ventral, or lapel portion. The membranelles of the amphisiellids have the ordinary fine structure, that is, each membranelle is composed of (i) two long kineties, (ii) one moderately long kinety, and (iii) one rather short kinety (Fig. 2a). Hypotrichs have two undulating membranes, the paroral and the endoral (Fig. 1a, 2a, 3a, 4a). For a detailed discussion of various patterns formed by the membranes, see Berger & Foissner (1997) and Berger (1999). In general, the paroral extends between two usually inconspicuous cytoplasmic lips at the right outer margin of the buccal cavity, that is, on the cell surface, while the endoral is on the bottom and right wall of the cavity (Foissner & AL-Rasheid 2006). This means that the membranes are at different levels, but when the cell is viewed from the ventral side, they appear to lie side by side or to intersect, depending on their shape and arrangement. The diversity of the undulating membranes pattern is lower in the amphisiellids than in the oxytrichids (Berger 1999). The buccal cavity is covered by a very fine mem1
DE-value = distance DE divided by length of adoral zone of membranelles (Fig. 1a).
20
GENERAL SECTION
brane, the so-called phago-assistant membrane (Sui et al. 2001) or buccal seal (Foissner & AL-Rasheid 2006). The buccal cavity is of different size and shape, usually described by the terms flat or deep and wide or narrow. Flat means that the cavity is only slightly hollowed, whereas a deep cavity extends to near the dorsal side of the cell, making the field conspicuously bright. In species with a wide cavity, the right margin of the cavity is in the midline of the cell, whereas in a narrow cavity it is close to the right margin of the adoral zone (further details, see Berger & Foissner 1997, Berger 1999, Foissner & AL-Rasheid 2006).
1.9 Silverline System The silverline system of the hypotrichs is composed of small (1–2 µm) polygonal meshes. It has no systematic value in the hypotrichs, although it is successfully used to characterise euplotids (e.g., Curds & Wu 1983, Foissner et al. 1991, Borror & Hill 1995) and other ciliate taxa (e.g., Foissner 1993, Foissner et al. 1995).
1.10 Life Cycle The species reviewed in the present book have, like most other hypotrichs, (very likely) a normal life cycle, that is, the theronts feed, become trophonts and divide, encyst, or conjugate. Literature about the non-morphostatic parts of the life cycle is scarce, as compared to the oxytrichids and urostyloids (for review of these groups, see Berger 1999, 2006).
1.10.1 Cell Division The species treated in the current volume divide by isotomic transverse fission, like many other ciliates (for review, see Foissner 1996). For a detailed description of a cell division, see, for example, Lamtostyla australis (Fig. 31g–z; Voß 1992). The anterior filial product is the proter, the posterior the opisthe. During very early phases of division, a replication (= reorganisation) band traverses each macronuclear nodule. The two to several nodules fuse to a single mass in early and middle dividers. By contrast, the micronuclei divide mitotically (details on nuclear apparatus, see chapter 1.2). The principles of the formation of the frontal-ventral-transverse cirri in the amphisiellids proceeds basically as in many oxytrichids, simply because both groups evolved from a hypotrich which very likely produced 18 cirri from six (I–VI) anlagen. Consequently, many plesiomorphic features occur in both taxa, for example, the migration of the frontoterminal cirri (= anterior portion of amphisiellid median cirral row). Even the long amphisiellid median cirral row dominating the ventral infraciliature of many amphisiellids can be easily homologised with the corresponding cirri of
MORPHOLOGY
21
the 18-cirri hypotrichs (Fig. 3a, d, 5a) and even the origin of the cirral anlagen (e.g., anlage III of proter from cirrus III/2) is largely identical. Of course, deviations exist, otherwise the taxa would not be different! The amphisiellid median cirral row, the apomorphy of the amphisiellids, is formed from three anlagen in some taxa (e.g., Hemiamphisiella, Spiroamphisiella, Maregastrostyla), namely from anlage VI (forms anterior portion), IV (middle portion), and V (posterior portion; details see Fig. 58f). In most amphisiellids, however, the amphisiellid median cirral row is formed from two anlagen, namely, anlage VI (anterior portion) and V ( posterior portion; details see Fig. 5a). The new marginal rows are usually formed within the parental rows, that is, two anlagen are produced within each row, one at the anterior end and one roughly in mid-body. Generally, the anlagen for the marginal rows occur at about the same level as the anlagen for the dorsal kineties (Fig. 31s, t). In Maregastrostyla the marginal rows are formed de novo and in the oxytrichid Ponturostyla the many marginal rows per side originate from a single anlage. Note that in Amphisiella, type of the whole group, and many other related taxa no dorsomarginal kineties are formed at/near the anterior end of the right marginal row anlage. If such a dorsomarginal kinety occurs in a certain taxon, a close relationship to the amphisiellids is very unlikely. I have radically cleared up the amphisiellids in that I removed all species with a dorsomarginal kinety (see next paragraph). Dorsal morphogenesis proceeds rather simple. Briefly, two anlagen occur within each parental kinety (Fig. 5b); dorsomarginal kineties and an oxytrichid kinety fragmentation do not occur. The same pattern is known from the urostyloids (Berger 2006). Consequently, I excluded all species which have either a dorsomarginal kinety (e.g., Nudiamphisiella) or an oxytrichid kinety fragmentation (e.g., Pseudouroleptus) from the amphisiellids. Apourosomoida halophila shows a dorsal kinety fragmentation which is, however, (very likely) not homologous to that of the oxytrichids. Since A. halophila lacks frontal-ventral-transverse cirri anlage V it is also (very likely) not an amphisiellid because in this group anlage V forms a major part (rear portion) of the amphisiellid median cirral row, the main amphisiellid apomorphy. Of course I cannot exclude that the amphisiellids are Dorsomarginalia which have lost the dorsomarginal kinety, or even oxytrichids, which have lost the dorsomarginal kinety and the oxytrichid kinety fragmentation. However, these assumptions are not very parsimonious and therefore I prefer the assumption that amphisiellids are hypotrichs which branched off outside the Dorsomarginalia (Fig. 9a). Further details, see chapter 2. Caudal cirri originate at the rear end of bipolar dorsal kineties (Fig. 1c), but not at the end of dorsomarginal rows. The presence of caudal cirri within the Hypotricha is interpreted as plesiomorphic state (see chapter 2). Many taxa have lost the caudal cirri, a rather simple feature which very likely evolved several times independently in the amphisiellids and hypotrichs in general.
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GENERAL SECTION
1.10.2 Conjugation Basically nothing is known about conjugation in the amphisiellids, the trachelostylids, and the other taxa reviewed here. However, the principles of this part of the life cycle are similar in all ciliates so please refer to reviews dealing with other groups of hypotrichs (Berger 1999, 2006) or ciliates in general (e.g., Dini & Nyberg 1993, Miyake 1996).
1.10.3 Cyst Only very little information about this stage of the life cycle is available and no paper deals with the ultrastructure of amphisiellid resting cysts (for review, see Gutiérez et al. 2003, Foissner 2005). If data are available then they are mentioned in the description of the individual species. Reproductive cysts are not known for taxa discussed here.
1.10.4 Reorganisation, Regeneration, Doublets Like other hypotrichs, the amphisiellids produce ciliature not only during cell division or other normal parts of the life cycle (conjugation, excystment), but also during physiological reorganisation. However, only very few data are available about this process (e.g., Fig. 18n–q). Generally, the formation of the ciliature during reorganisation proceeds very similarly to that during cell division (Fig. 5a).
2
Phylogeny
Little is known about the phylogeny of the taxa discussed. Chapter 2.1 briefly explains the nomenclature of the hypotrichs and spirotrichs because at present two systems exist. In chapter 2.2 the ground pattern of the Hypotricha is described. This is important for the estimation of the phylogeny within this group and within subgroups, for example, the amphisiellids. Of course, this pattern is not final because new data about the morphology, cell division, ultrastructure, and molecular biology and hypotheses are constantly becoming available. Chapter 2.3 covers some ideas about the phylogenetic position of two taxa (Diophrys, discocephalids) previously usually assigned to the euplotids. For some comments about the phylogenetic relationships within the hypotrichs, including the rough estimation of the position of the amphisiellids, see chapter 2.4.
PHYLOGENY
23
2.1 Notes on Nomenclature The naming of the major taxa of the Spirotricha is not uniform at present (for review, see Berger 2006, p. 29). To avoid confusion, the names used in the present book and in other revisions are mapped in Table 3, and their hierarchy and supposed relationships are shown in Fig. 6a, c. For authorship of hypotrich and euplotid taxa not discussed here, see Berger (2001) and the permanently updated online-version of this nomenclator at http://www.protozoology.com. For authorship of other taxa, see, for example, Corliss (1979), Aescht (2001), and Lynn & Small (2002).
2.2 The Ground Pattern of the Hypotricha Stein, 1859 The ground pattern of a monophyletic group (evolutionary unit) is the combination of features of the stem-species from which the monophylum evolved, that is, it is a summary of apomorphies and plesiomorphies present in the last common ancestor (Ax 1995). Of course, only more or less young plesiomorphies are included in the ground pattern. Old plesiomorphies – for example, the presence of cilia in the Hypotricha – are not included. According to estimates by Wright & Lynn (1997) using a small subunit rRNA molecular clock, Oxytricha separated from Onychodromus + Stylonychia about 350 million years ago, and Protocruzia branched off from the Oxytricha + (Onychodromus + Stylonychia) group in the interim period of Mesoproterozoic to Neoproterozoic, that is, about 109 years ago. Consequently, the last common ancestor of the hypotrichs likely lived in the period before 400 million years. For a detailed description of the morphological features see chapter 1. Note that new data and hypotheses will change the situation more or less distinctly. In addition, misinterpretations of data and states of features can modify the situation. For details on the oligotrichs, which are very likely the next relatives of the hypotrichs, see, for example, Agatha (2004) and Agatha & Strüder-Kypke (2007) and references therein. For a schematic illustration of the supposed last common ancestor of the Hypotricha see Fig. 7a, b.
2.2.1 Apomorphies of the Hypotricha The following features (very likely) characterise the hypotrichs (Fig. 7a, b), that is, they evolved in the stem-line and were present in the last common ancestor of this taxon (Fig. 6a, 9a). The main character is the highly characteristic pattern formed by the 18 frontal-ventral-transverse cirri. 18-frontal-ventral-transverse cirri in highly characteristic pattern. For detailed designation of cirri and origin of cirri see Fig. 3a, c, 4a in this book and Fig. 6a in Berger (1999). This is a rather complex feature because it is a combination of morphological and ontogenetic traits. Consequently, a convergent evolution of this pat-
24
GENERAL SECTION
Table 3 Comparison of names of some higher spirotrich taxa used in four recent books and in the present review (from Berger 2006, supplemented) Present book a
Corliss (1979)
Small & Lynn (1985)
Tuffrau & Fleury (1994)
Lynn & Small (2002)
Spirotricha Bütschli, 1889 (spirotrichs)
Polyhymenophora Jankowski, 1967 b
Spirotrichea Bütschli, 1889
Spirotricha Bütschli, 1889
Spirotrichea Bütschli, 1889
Euplota Ehrenberg, 1830 (euplotids)
Euplotidae Ehrenberg, 1838; Aspidiscidae Ehrenberg, 1838; Gastrocirrhidae Fauré-Fremiet, 1961
Hypotrichia Stein, 1859 d
Euplotia author? f
Hypotrichia Stein, 1859
Oligotricha Bütschli, 1889 i (oligotrichs)
Oligotrichida Bütschli, 1889
Choreotrichia n. subclass.
Oligotrichea Bütschli, 1887 g
Choreotrichia Small & Lynn, 1985; Oligotrichia Bütschli, 1889
Hypotricha Stein, 1859 (hypotrichs)
Hypotrichida Stein, Stichotrichia n. 1859 c subclass. e
Oxytrichia author? f Stichotrichia Small & Lynn, 1985
Amphisiellidae Jankowski, 1979 (amphisiellids)
-
Amphisiellidae n. fam.
Amphisiellidae Jankowski, 1979
Amphisiellidae Jankowski, 1979
Oxytrichidae Ehrenberg, 1838 h (oxytrichids)
Oxytrichidae Ehrenberg, 1838
Oxytrichidae Ehrenberg, 1838
Oxytrichidae Ehrenberg, 1838
Oxytrichidae Ehrenberg, 1838
Trachelostylidae Small & Lynn, 1985 (trachelostylids)
-
Trachelostylidae fam. n.
-
Trachelostylidae Small & Lynn, 1985
Urostyloidea Bütschli, 1889 (urostyloids)
Urostylidae Bütschli, 1889; Holostichidae Fauré-Fremiet, 1961
Urostylina Jankowski, 1979
Urostylida Jankowski, 1979
Urostylida Jankowski, 1979
a
Note that in the present book the names are usually used as originally introduced (see Berger 2001). Vernacular names in brackets.
b
With the single subgroup Spirotricha Bütschli, 1889 (for a brief discussion of the redundancy of the name Polyhymenophora, see Berger & Foissner 1992).
c
Also includes the Euplotidae, the Aspidiscidae, and the Gastrocirrhidae.
d
With the single subgroup “Euplotida n. ord.”
e
With the single subgroup Stichotrichida Fauré-Fremiet, 1961.
f
According to Berger (2001) the authors of this subclass are Tuffrau & Fleury (1994).
g
Incorrect year.
h
Incorrectly spelled “Oxytrichidea” in Berger (1999, p. 102).
i
Comprises the orders Oligotrichida and Choreotrichida (Agatha & Strüder-Kypke 2007). Whether the halterids (Halteria, Pelagohalteria, Meseres) are oligotrichs or hypotrichs is not yet known.
PHYLOGENY
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Fig. 6a Names and supposed relationships of the main taxa of the Spirotricha (original; some small taxa, for example, Phacodinium, not considered). Note that (i) I usually use the same spellings as proposed in the original descriptions, and (ii) I do not use categories (e.g., family, order; see chapter 7.2). Diophrys is, according to various data, very likely not a euplotid; the same is true for the discocephalids. The extent of the new taxon Perilemmaphora (Oligotricha + Hypotricha or Diophrys to Hypotricha) is not yet certain. Further details see chapter 2.3. Main apomorphies (squares 1–9; for discussion of features, see chapter 2. ? = apomorphy not known or uncertain): 1 – replication band present. 2 – high support by molecular data. 3 – two undulating membranes?; perilemma present? (see also 7). 4 – ? 5 – two frontoterminal cirri; two pretransverse ventral cirri; two distinct marginal rows. 6 – more than six frontal-ventral-transverse cirri anlagen form a discocephalid median cirral row, which is a pseudorow; distinct cephalisation; more than two macronuclear nodules. 7 – perilemma? (see also 3); high support by molecular data. 8 – planktonic life; loss of cirri; high support by molecular data 8 (more data see Agatha 2004 and Agatha et al. 2007). 9 – 18 frontal-ventral-transverse cirri; three dorsal kineties; three caudal cirri; contractile vacuole at left body margin; high support by molecular data. Well-known representatives: Hypotricha (hypotrichs): Amphisiella, Holosticha, Oxytricha, Sterkiella, Stylonychia, Uroleptus, Urostyla Oligotricha (oligotrichs): Codonella, Limnostrombidium, Lohmanniella, Strobilidium, Strombidium Discocephalidae (discocephalids): Discocephalus, Marginotricha, Prodiscocephalus Diophrys: Diophrys Euplota (euplotids): Aspidisca, Euplotes, Certesia, Uronychia?
tern, as supposed by Eigner (1997, p. 553), is highly unlikely. Previously we assumed that this pattern is an apomorphy of the Oxytrichidae (Berger & Foissner 1997; Berger 1999, p. 76, 102). Later we came to the conclusion that this pattern is older and that even the zigzagging midventral pattern of the urostyloids and other, non-related taxa (e.g., Neokeronopsis) can be derived from an 18-cirri hypotrich using the CEUU-hypothesis (Foissner et al. 2004). However, in the monograph on urostyloids I fixed the 18-cirri pattern, which originates from six (I–VI) cirral anlagen, as apomorphy of the Hypotricha (Berger 2006, p. 33). Six cirral anlagen are already
26
GENERAL SECTION
Fig. 6b–d The three possibilities to arrange the three major taxa of the spirotrichs (from Berger 2006). Protocruzia, Phacodinium, Diophrys, and discocephalids not considered.
present in the euplotids (e.g., Wallengren 1900, Curds & Wu 1983), but they form less than the 18 frontal-ventral-transverse cirri (Fig. 16a in Berger 2006). And even Diophrys, which already has two undulating membranes like the hypotrichs, does not have the full number of 18 cirri (e.g., Curds & Wu 1983, Song & Packroff 1993; Fig. 8a). The discocephalids, which also posses two undulating membranes, have increased the number of cirral anlagen and therefore form a somewhat deviating cirral pattern (Wicklow 1982b). However, specific elements of the 18-cirri pattern, for example, the anteriorly migrating frontoterminal cirri or the two pretransverse ventral cirri ahead the two rightmost transverse cirri, are already present in this little known taxon (Fig. 8c, d; Wicklow 1982b, Lin et al. 2004). This indicates, that the 18-cirri pattern did not evolve at once, but over a long period. Of course, one cannot exclude that the 18-cirri pattern was already present in the last common ancestor of the Perilemmaphora (oligotrichs + hypotrichs; Fig. 6a, square 7). However, in the stem-line of the oligotrichs all cirri vanished without a trace and therefore we can postulate that the 18-frontal-ventral-transverse cirri pattern (Fig. 7a) occurred for the first time in the stem-line of the Hypotricha. Using the 18-cirri pattern as a starting point, very many cirral patterns occurring in the hypotrichs can be explained rather easily, for example, the midventral pattern, which evolved several times independently (e.g., urostyloids, uroleptids, some oxytrichids like Neokeronopsis and Territricha) by insertion of few to many anlagen (see Berger 2006), or the amphisiellid median cirral row which is often so distinct because the anlagen IV, V, and VI do not form few (basically 2–4), but many cirri (Fig. 2a, 5a). In addition, the present hypothesis also explains the paraphyly of the Oxytrichidae briefly discussed – but not explained – in some molecular works (e.g., Schmidt et al. 2004, 2007), because the 18-cirri pattern is a plesiomorphy within the Hypotricha and therefore cannot be used to estimate the evolution within the hypotrichs. The Oxytrichidae are now defined mainly via the oxytrichid dorsal kinety fragmentation (Berger 1999, 2006). Consequently, 18-cirri hypotrichs without such an oxytrichid fragmentation and without dorsomarginal kineties, for example, Trachelostyla pediculiformis (Gong et al. 2006; Fig. 4a; see Addenda for details on the cell division of T. pediculiformis), may not be classified in the oxytrichids, but obviously branched off outside the Dorsomarginalia Berger, 2006, as so-called nondorsomarginalian hypotrichs (Fig. 9a). Such a basal position is also indicated by gene sequence data (e.g., Schmidt et al. 2007). Further details, see chapter 2.3.
PHYLOGENY
27
Fig. 7a, b Schematic illustration (original) of the ventral and dorsal side of the supposed last common ancestor of the Hypotricha (Fig. 6a, square 9; Fig. 9a, square 1). The 18 frontal-ventral-transverse cirri and the three caudal cirri are grey, the marginal cirri are black. Broken lines connect cirri which originate from the same anlage; dotted lines in (a) connect or surround cirral groups. Perhaps the three postoral ventral cirri have been right of the proximal portion of the adoral zone in the last common ancestor (dotted cirri). Detailed explanation of structures see chapter 1; details about the phylogeny see chapter 2. AZM = adoral zone of membranelles, BC = buccal cirrus, CC = caudal cirri, CV = contractile vacuole, FC = frontal cirri, FVC = frontoventral cirri, LMR = left marginal row, PTVC = pretransverse ventral cirri, PVC = postoral ventral cirri, RMR = right marginal row, TC = transverse cirri, I–VI = frontal-ventral-transverse cirri anlagen, 1–4 = cirri within anlage (Fig. 7a), 1–3 = dorsal kineties (Fig. 7b).
28
GENERAL SECTION
Contractile vacuole near left cell margin(?). The contractile vacuole of hypotrichs is, if present at all, near the left cell margin (Fig. 1c), indicating that the vacuole was at this site in the last common ancestor of the Hypotricha (e.g., Kahl 1932, Berger 1999, 2006). Euplotids (e.g., Euplotes, Aspidisca, Cytharoides, Uronychia, Certesia) have the contractile vacuole subterminally near the right body margin or terminally (e.g., Kahl 1932, Foissner et al. 1991, Petz et al. 1995, Lin & Song 2004). In the oligotrichs the situation is rather diverse because the vacuole is either lacking, terminal, subterminal, or – as in the hypotrichs – at the level of the oral apparatus in tintinnids (e.g., Foissner et al. 1999). Agatha & Strüder-Kypke (2007) did not use this feature in their phylogenetic analyses. In the halterids (e.g., Meseres corlissi; Petz & Foissner 1992, Foissner et al. 1999) the vacuole is in the same position as in the hypotrichs. In addition, it empties via a permanent excretion pore between the second and third somatic kinety, which is reminiscent, like some other morphological features and the molecular data, of the hypotrichs (for review, see Foissner et al. 2004a). I did not find data about the excretion pore in tintinnids so I do not know whether the vacuole empties via the dorsal side as in the hypotrichs or, for example, into the buccal cavity as in the peritrichs (e.g., Schröder 1906, Ruiz & Anadón 1987). Diophrys and discocephalids likely lack a contractile vacuole (Kahl 1932, Wicklow 1982b, Lin et al. 2004). Consequently, the arrangement of the vacuole at the left cell margin could be an apomorphy of the Hypotricha. Usually the vacuole of the hypotrichs is at the left margin between the level of the buccal vertex and mid-body. Rarely, it is displaced distinctly posteriad, for example in Holosticha pullaster or Anteholosticha pulchra (Berger 2006, Li et al. 2007). Whether this position shows us the “migration” of the contractile vacuole from the right subterminal position in the euplotids to the left margin is not known. Three dorsal kineties. Three bipolar kineties are widely distributed in the hypotrichs, for example, in the stem-line of the urostyloids (Berger 2006), but also in many amphisiellids (Table 22). Oxytrichids also have three bipolar kineties, but kinety 3 fragments, which is the major apomorphy of this group (Berger 1999, 2006; Fig. 9a). The rightmost kineties in oxytrichids and in the Dorsomarginalia, to which the oxytrichids belong, are dorsomarginal kineties, that is, they are of different origin and a novelty for the Dorsomarginalia (Berger 1999, 2006; Fig. 9a). Euplotids, Diophrys, and discocephalids have usually more than three kineties (e.g., Wicklow 1982b, Curds & Wu 1983, Voss 1989, Song & Packroff 1993, 1997, Lin et al. 2004, Song et al. 2007), indicating that three dorsal kineties – each with one caudal cirrus at its distal end (see next feature) – are a novelty for the hypotrichs (Berger 2006, p. 78). Of course this pattern was heavily modified within the hypotrichs. Note that the dorsal kinety pattern is as important as the ventral cirral pattern for the classification of the hypotrichs! Three caudal cirri. Caudal cirri originate at the end of the bipolar dorsal kineties (Fig. 1c). Dorsomarginal kineties and the anterior fragment(s) of a splitting dorsal kinety are never associated with such cirri. The loss of some or all caudal cirri occurred many times independently within the hypotrichs; in some taxa the number of caudal cirri per kinety increased significantly (for review, see Berger 1999, 2006, present book). However, caudal cirri in general are older because euplotids, Diophrys, and dis-
PHYLOGENY
29
cocephalids also form such cirri at the end of some dorsal kineties (e.g., Wicklow 1982b, Voss 1989, Song & Packroff 1993, Lin et al. 2004).
2.2.2 Plesiomorphies of the Hypotricha In the following paragraphs the most important, more or less young plesiomorphies of the Hypotricha are discussed. Plesiomorphic means that a feature was already present before the oligotrichs, the supposed sister-group of the hypotrichs, branched off (Fig. 6a). Note that many features agree with the ground pattern of the urostyloids described by Berger (2006). In addition, the list below is not complete (Berger 2007). Body length about 100 µm. There is no evidence that the last common ancestor of the Hypotricha was very large or very small. Many species are, like many oligotrichs (Kahl 1932), discocephalids (Wicklow 1982b, Lin et al. 2004), and euplotids (Curds & Wu 1983), around 100 µm long, indicating that the last common ancestor of the Hypotricha had roughly such a length. Within several groups (e.g., Urostyloidea, Stylonychinae; Berger 1999, 2006) some very large (500 µm or more) species evolved. Very small species (50 µm or less) are also rare. Body elongate elliptical. The body outline of many hypotrichous ciliates is elongate elliptical (body length:width ratio roughly around 2–3:1; e.g., Kahl 1932, Berger 1999, 2006). By contrast, the euplotids and Diophrys usually have a distinctly lower value (usually below 2:1, often almost 1:1; of course exceptions exist; e.g., Curds & Wu 1983). Discocephalids, which very likely branched off later than Diophrys, have – like the hypotrichs – an elongated body, indicating that this feature is a young plesiomorphy for the Hypotricha. Body distinctly dorsoventrally flattened. The body of almost all hypotrichs is distinctly flattened dorsoventrally (Fig. 1b). Discocephalids, euplotids, and even Phacodinium are also strongly flattened, indicating that this is a plesiomorphy for the Hypotricha (e.g., Kahl 1932, Wicklow 1982b, Curds & Wu 1983, Lin et al. 2004). If we assume that the oligotrichs are the sister group of the hypotrichs (Fig. 6a, c) then this feature was obviously lost in the last common ancestor of the oligotrichs because this primarily planktonic group is not flattened (e.g., Kahl 1932, Foissner et al. 1999, Agatha & Riedel-Lorjé 2006), that is, a globular or obconical body is an apomorphy for the oligotrichs (Agatha & Strüder-Kypke 2007). Thus, the dorsoventral flattening of some benthic oligotrichs is likely a novelty for these species (e.g., Xu et al. 2006). Body flexible. The majority of hypotrichs has a flexible body when freely motile (if cells are squeezed, then even the body of rigid species becomes more or less flexible). Within the hypotrichs only the Stylonychinae – a comparatively small subgroup of the Oxytrichidae – and some species of uncertain position (e.g., Urospinula, Rigidothrix) have a rigid body (e.g., Foissner 1983b, Berger 1999, Foissner & Stoeck 2006). The oligotrichs, the supposed sister group of the hypotrichs (Fig. 6a), also have a soft body, indicating that the last common ancestor of the hypotrichs was flexible. Likely all
30
GENERAL SECTION
euplotids have an inflexible body (e.g., Stein 1859, Petz et al. 1995, Lei et al. 2002), indicating that this is an apomorphy of the euplotids. Within the discocephalids the situation is not uniform; most species are rigid, but at least Marginotricha faurei has a supple body (Wicklow 1982b). Macronuclear nodules fuse to a single mass during cell division. In ciliate species, including hypotrichs, with two or more macronuclear nodules, these pieces fuse to a single mass during cell division (e.g., Czapik 1981, Raikov 1982, Petz & Foissner 1993, Berger 1999, Foissner et al. 2002, Shao et al. 2007). Consequently this is a rather old plesiomorphy in the last common ancestor of the Hypotricha. The individual division of the many nodules in the Pseudokeronopsinae is an apomorphy for this group (for review, see Berger 2006). Replication band of macronucleus. In this band the DNA is replicated locally and sequentially along the macronucleus (e.g., Raikov 1982, 1996, Olins & Olins 1994). It is likely the most important morphological apomorphy of the spirotrichs (Lynn 2003). Thus, the presence of such a band is a plesiomorphy in the stem-line of the Hypotricha. Two replication bands are present in species with a C-shaped macronucleus, for example, in Aspidisca (e.g., Summers 1935), Euplotes (Voss 1989), or in Strobilidium gyrans sensu Deroux (1974). In species with two or more macronuclear nodules one band per nodule is present (e.g., Summers 1935, Berger 1999, 2006). Usually they migrate from the distal ends of the macronuclear apparatus towards the centre, for example in Euplotes, Diophrys, Strobilidium, or hypotrichs with two nodules (e.g., Griffin 1910, Summers 1935, Deroux 1974, Voss 1989, Berger 1999). In Aspidisca, however, the bands migrate in the opposite direction (Summers 1935), which is very likely an apomorphy for Aspidisca. Cortical granules present. These organelles, which likely belong to the mucocyst type, are present or absent in the hypotrichs. Cortical granules are reported from many other ciliate taxa, for example, Diophrys (Song et al. 2007), colpodids (e.g., Foissner 1993), haptorids (e.g., Foissner 1984, Xu & Foissner 2005), or heterotrichids (e.g., Stein 1867, Foissner et al. 1992). Cortical granules are obviously lacking in oligotrichs (e.g., Foissner et al. 1999, Agatha & Strüder-Kypke 2007) and perhaps also in discocephalids (Lin et al. 2004). Thus, we have to assume that these granules are an old feature, provided that they are homologous. Consequently, cortical granules were already present in the last common ancestor of the hypotrichs. The loss of the granules in various species or genera of hypotrichs evolved convergently. The Stylonychinae are a relatively large group having the loss of cortical granules as apomorphy (Fig. 9a; Berger & Foissner 1997, Berger 1999). The stylonychines are the best founded group of hypotrichs both from the morphological as well as from the molecular biological point of view. Adoral zone of membranelles “short”. In most species of hypotrichs, the length of the adoral zone of membranelles is less than 40%, often between 20% and 35% of body length (for reviews, see Kahl 1932, Berger 1999, 2006, present book). The same is true for the discocephalids (Wicklow 1982b, Lin et al. 2004), indicating that this is a plesiomorphy for the last common ancestor of the hypotrichs. A comparison with the oligo-
PHYLOGENY
31
trichs is difficult because they have a rather different habitus; however, their adoral zone is very prominent because it occupies the anterior cell end. In Phacodinium, the euplotids, and Diophrys the relative size of the adoral zone is much larger than in the hypotrichs; usually it occupies distinctly more than 50% of body length (e.g., Kahl 1932, Curds & Wu 1983, Foissner et al. 1991, Fernández-Galiano & Calvo 1992, Song & Packroff 1993, Song et al. 2007). The last common ancestor of the Stylonychinae again evolved a large adoral zone usually occupying more than 40% of body length (Berger & Foissner 1997, Berger 1999). Adoral zone of membranelles continuous. The adoral zone of most hypotrichs is continuous, that is, it does not show a distinct break in the left anterior corner of the cell where it extends on the dorsal side of the frontal scutum. This basically agrees with the situation in many oligotrichs, discocephalids, Diophrys, euplotids, and Phacodinium (e.g., Kahl 1932, Curds 1975, Wicklow 1982b, Song & Packroff 1993, Agatha 2004, Agatha & Strüder-Kypke 2007), indicating that a continuous row is a plesiomorphy in the ground pattern of the Hypotricha. Two undulating membranes. In the previous monograph I rashly assumed that the presence of two undulating membranes, respectively, the presence of an endoral is an apomorphy of the Hypotricha (Berger 2006, p. 77), simply because I overlooked that Diophrys, a genus usually assigned to the euplotids, has two undulating membranes (e.g., Nobili & Rosati Raffaelli 1971, Hill & Borror 1992, Song & Packroff 1993, Song & Wilbert 1994; Fig. 8a). According to the original description, Cytharoides also has two undulating membranes (Tuffrau 1974), a feature not confirmed by three redescriptions (Agatha et al. 1990, Petz et al. 1995, Song & Wilbert 2000). Hill (1990) described a right and left oral membrane in Uronychia; however, according to Song et al. (2004 and references therein), the oral primordium produces only one very large, almost circular membrane. Other euplotids (Euplotes, Euplotidium, Aspidisca, Uronychia, Certesia, Gastrocirrhus) and Phacodinium almost certainly have only one undulating membrane (e.g., Foissner et al. 1991, Fernández-Galiano & Calvo 1992, Giambelluca et al. 1995, Hu & Song 2003, Song et al. 2004, Lin & Song 2004). Kiitricha, a taxon whose phylogenetic position is not yet clearly recognised, also has two undulating membranes (e.g., Fleury et al. 1986, Song & Wilbert 1997a). Discocephalus and Prodiscocephalus and some related taxa also have a paroral and an endoral (e.g., Wicklow 1982b, Lin et al. 2004). Therefore I prematurely supposed that the discocephalids, which are usually assigned to the euplotids (e.g., Lynn & Small 2002, Adl et al. 2005), belong to the Hypotricha (Berger 2006, p. 30, 77). For a somewhat more detailed discussion and correction of this assumption, see chapter 2.3. In Diophrys the undulating membranes and the general appearance of the oral apparatus and several other features, for example, the formation of the dorsal kineties at two levels, the resorption of parental somatic ciliature, the origin of the oral primordium near the left transverse cirrus, agree very well with the situation in the Hypotricha (e.g., Song & Packroff 1993, Berger 1999, 2006), strongly indicating that Diophrys is more closely related to the group formed by the oligotrichs + hypotrichs than to the euplotids (Fig. 6a). A separation of Diophrys from the euplotids is also supported by a different
32
GENERAL SECTION
PHYLOGENY
33
stop codon usage (Perez-Romero et al. 2001). UAA, UAG, and UGA are the universal stop codons (e.g., Seyffert 2003) which are also used in Entodinium and Spathidium (Perez-Romero et al. 2001). Oxytricha, Stylonychia, Paraurostyla, and Diophrys use only UGA as stop codon whereas UAA and UAG are translated to glutamine (Prescott 1994, Perez-Romero et al. 2001). The same usage is described for Paramecium and Tetrahymena, indicating that this is a plesiomorphy for the hypotrichs and Diophrys. Since plesiomorphies do not support monophyla, but paraphyla, this stop codon feature cannot be used to support the relationship of Diophrys to the group oligotrichs + hypotrichs. By contrast, Euplotes uses, as is usual, UAA and UAG as stop codons, but translates UGA into cysteine (Table 3a), indicating that this is an apomorphy for Euplotes, perhaps even for the euplotids (Borror & Hill 1995). Stop codon usage of further euplotids (e.g., Aspidisca, Uronychia), discocephalids, and oligotrichs are needed to get a better insight into the phylogeny of the spirotrichs. A position of Diophrys closer to the group oligotrichs + hypotrichs than to the euplotids is also shown by some phylogenies based on 18S rRNA gene sequences (e.g., Song et al. 2004, their Fig. 37, not Fig. 38; Li & Song 2006, their Fig. 3, not Fig. 2; Shao et al. 2006) and by the supposed presence of a perilemma (see this plesiomorphy). Interestingly, Uronychia is usually closely related to Diophrys in molecular trees, although it does not have two undulating membranes (see above for some uncertainties). By contrast, Šlapeta et al. (2005) found that Diophrys branches off outside the group euplotids + (oligotrichs + hypotrichs). Agatha (2004) assumed that the second undulating membrane is a common feature for the euplotids, the oligotrichs, and hypotrichs with a convergent loss of one membrane in some euplotids and all oligotrichs. The presence of two undulating membranes in the halterids (Halteria, Meseres) supports most molecular trees which show that the halterids are hypotrichs. For a discussion of the two undulating membranes of Halteria, see Foissner et al. (2004). Undulating membranes long and curved. The undulating membranes of the hypotrichs are of rather different arrangement and length. Berger & Foissner (1997) and Berger (1999) assumed that relatively long and more or less distinctly curved undulat-
b
Fig. 8a, b Diophrys scutum (from Song & Packroff 1993. Protargol impregnation). Infraciliature of ventral and dorsal side and nuclear apparatus. Note the two undulating membranes and the large transverse cirri. Broken lines connect cirri which originate from same frontal-ventral-transverse cirri anlage. Dotted line connects frontal cirri; arrow marks cirrus which is obviously homologous to the buccal cirrus of the 18-cirri hypotrichs. AZM = adoral zone of membranelles, CC = caudal cirri (originate from rightmost dorsal kinety), E = endoral, LMR = left marginal row, P = paroral, TC = transverse cirri, I–VI = frontalventral-transverse cirri anlagen, 1 = dorsal kinety 1 (= leftmost kinety). Fig. 8c, d Marginotricha faurei (from Wicklow 1982b [designated as Psammocephalus faurei in this paper]. Protargol impregnation). Infraciliature of ventral side of interphasic specimen and very late divider. Broken lines in (d) connect cirri originating from the two rightmost frontal-ventral-transverse cirri anlagen, which are certainly homologous to the anlagen V and VI of the Hypotricha because they also form the two pretransverse ventral cirri and two frontoterminal cirri. Note that the median cirral row in the interphasic specimen (c) is a pseudorow (Fig. 1e) because each cirrus originates from a different anlage (d). FT = frontoterminal cirri, LMR = left marginal row, PTVC = pretransverse ventral cirri, RMR = right marginal row, TC = transverse cirri.
34
GENERAL SECTION
Table 3a Genetic code deviations of some ciliates (after Prescott 1994 and Perez-Romero et al. 2001) Species
Entodinium
Universal stop codon use UAA
UAG
UGA
stop
stop
stop
Spathidium
stop
stop
stop
Paramecium
glutamine
glutamine
stop
Tetrahymena
glutamine
glutamine
stop
Oxytricha
glutamine
glutamine
stop
Stylonychia
glutamine
glutamine
stop
Paraurostyla
glutamine
glutamine
stop
Diophrys
glutamine
glutamine
stop
Euplotes
stop
stop
cysteine
ing membranes represent the ancestral state (Oxytricha-pattern) within the oxytrichids. Interestingly, Diophrys shows more or less exactly such a pattern (Fig. 8a), indicating that this is a plesiomorphy in the ground pattern of the Hypotricha. Three frontal cirri. The first occurrence of this feature is difficult to determine because these are the anteriormost cirri of the three leftmost anlagen (I–III), which are already present in euplotids, indicating that it occurred for the first time in the last common ancestor of the group euplotids + (oligotrichs + hypotrichs), that is, immediately after Phacodinium branched off. Anyhow, it is very likely a plesiomorphy for the last common ancestor of the Hypotricha. One buccal cirrus. This cirrus (= cirrus II/2), which is usually located immediately right of the paroral, is already present in euplotids (e.g., Wallengren 1900), Diophrys (e.g., Song & Packroff 1993), and very likely discocephalids (Wicklow 1982b). Thus, the presence of this cirrus is a plesiomorphy for the last common ancestor of the Hypotricha (e.g., Fig. 2a). For the complicated terminology of this cirrus, see Berger (1999). Two frontoterminal cirri. Many hypotrichs have two so-called frontoterminal cirri or migratory cirri originating from the rightmost anlage, which is homologous to anlage VI of the 18-cirri hypotrichs (e.g., Fig. 3a). They migrate anteriorly to near the distal end of the adoral zone and do not participate in anlagen formation. Two frontoterminal cirri are already present in Marginotricha faurei, a discocephalid (Wicklow 1982b, p. 319, Fig. 8c, d), indicating that this feature occurred in an ancestor before the discocephalids branched off, that is, it is a plesiomorphy for the last common ancestor of the Hypotricha. Two pretransverse ventral cirri. These two cirri are widely distributed within the hypotrichs (for review, see Berger 1999, 2006, present book). They can also be found in discocephalids (e.g., Wicklow 1982b, Lin et al. 2004; Fig. 8d), showing that they were already present in a stem-line species before the discocephalids branched off.
PHYLOGENY
35
Consequently, the presence of these two cirri is a plesiomorphy for the last common ancestor of the Hypotricha. Five transverse cirri. Transverse cirri are a typical part of the hypotrich cirral pattern. Five transverse cirri, that is, the rearmost, usually distinctly enlarged cirri formed by the frontal-ventral-transverse cirri anlagen II–VI are already present in the euplotids and Diophrys (e.g., Wallengren 1900, Kahl 1932, Curds & Wu 1983, Voss 1989, Foissner et al. 1991, Song & Packroff 1993, Song et al. 2007). Of course this pattern was frequently modified and transverse cirri have often been lost in various taxa. Discocephalids have more than five large transverse cirri due to an increase of the anlagen number (Wicklow 1982b, Lin et al. 2004); likely this is an apomorphy for this group. Transverse cirri large. In euplotids, Diophrys (Fig. 8a), discocephalids (Fig. 8c), and many hypotrichs the transverse cirri are distinctly larger than the remaining frontal-ventral and marginal cirri, showing that this is a plesiomorphy in the last common ancestor of the Hypotricha. One left and one right marginal row. All hypotrichs have a left and a right marginal row composed of a considerable number of cirri (e.g., Kahl 1932). Such distinct rows are lacking in euplotids (e.g., Curds 1975, Curds & Wu 1983), but present in the discocephalids (Wicklow 1982b, Lin et al. 2004). Thus, the marginal rows are a plesiomorphy for the Hypotricha and not an apomorphy as I supposed in the monograph of the urostyloids, where I assumed that the discocephalids are a subgroup of the hypotrichs because they have two undulating membranes (Berger 2006, p. 78). Euplotids and Diophrys often have two left marginal cirri and caudal cirri formed by the rightmost dorsal kineties (e.g., Voss 1989, Song & Packroff 1993). In Diophrys scutum the rightmost dorsal kinety produces three caudal cirri (Song & Packroff 1993). Perhaps this is a pre-stage of the right marginal row because marginal rows and dorsal kineties are (very likely) homonomous structures (Berger et al. 1985a, p. 309). Caudal cirri present. Caudal cirri are a rather old feature within the spirotrichs, already occurring in the euplotids. However, note that the presence of exactly three caudal cirri (one per bipolar kinety) is obviously a novelty for the Hypotricha (see apomorphies). Oral primordium originates on cell surface. In all Hypotricha the oral primordium of the opisthe originates more or less on the cell surface, that is, epiapokinetally (Foissner 1996). This also applies to the discocephalids (Wicklow 1982b). By contrast, in the euplotids and the oligotrichs the oral primordium develops hypoapokinetally; however, this mode very likely evolved convergently (for review, see Foissner 1996, Agatha 2004, Agatha & Strüder-Kypke 2007). Interestingly, in Diophrys the rear portion of the adoral zone is also formed in a groove (e.g., Song & Packroff 1993). However, the question is whether or not the modes of euplotids and Diophrys are homologous. In the monograph on urostyloids I considered the formation of the oral primordium on the cell surface as apomorphy of the Hypotricha, due to a different interpretation of the data (Berger 2006). Now I follow Foissner (1996), who assumed that the epiapokinetal formation is a plesiomorphy for the last common ancestor of the Hypotricha.
36
GENERAL SECTION
Adoral membranelles mature in an anterior to posterior direction. This mode is present in all hypotrichs (e.g., Berger 1999, 2006, present book) and probably oligotrichs (e.g., Agatha 2003, Agatha et al. 2004), but also in discocephalids (Wicklow 1982b) and Diophrys (Song & Packroff 1993). By contrast, membranelles are formed in the opposite direction in Euplotes (Voss 1989). Unfortunately, I did not find data about Phacodinium to make an outgroup comparison. Anyhow, the present feature is obviously a plesiomorphy for the last common ancestor of the Hypotricha. Frontal-ventral-transverse cirri originate from six (I–VI) anlagen. This is a relatively old feature already present in euplotids and Diophrys (e.g., Wallengren 1900, Curds & Wu 1983, Song & Packroff 1993). That is, the six anlagen forming the 18frontal-ventral-transverse cirri of the Hypotricha are a plesiomorphy. Marginal rows and dorsal kineties divide intrakinetally. Intrakinetal proliferation formation is a rather old plesiomorphy because even present in, for example, the spathidiids (Berger et al. 1983). Only in few hypotrich taxa (e.g., Thigmokeronopsis; Maregastrostyla, Fig. 26n) do at least one of these structures originate de novo, which has to be interpreted as apomorphy (for review, see Berger 2006). If more than one marginal row per side is present then each row divides individually (e.g., Architricha; Gupta et al. 2006) or they originate from a single anlage per filial product (e.g., Pseudourostyla, Ponturostyla [Fig. 138r], Allotricha; Berger 1999, 2006, Song 2001). Marginal rows and dorsal kineties originate independently for proter and opisthe. In almost all hypotrichs marginal rows and dorsal kineties develop more or less independently for proter and opisthe. The dorsal kineties of Diophrys and Uronychia are formed in the same manner (e.g., Song & Packroff 1993, Song et al. 2004), strongly indicating that this feature is a plesiomorphy for the last common ancestor of the Hypotricha. Usable data for the discocephalids are lacking (Wicklow 1982b). Consequently, the formation of the dorsal kineties via primary primordia in Maregastrostyla pulchra (Fig. 26m) has to be interpreted as apomorphy. Parental somatic ciliature completely replaced during cell division. Usually, in the hypotrichs no part of the parental somatic ciliature (frontal-ventral-transverse cirri, marginal rows, dorsal kineties, caudal cirri) is retained in postdividers (for reviews, see Berger 1999, 2006, present book). The same is true for the discocephalids (Wicklow 1982b) and Diophrys (Song & Packroff 1993). By contrast, in the euplotids and the oligotrichs, and in ciliates in general, at least parts of the parental somatic ciliature are present in the post-dividers (e.g., Foissner 1996, Agatha 2004, Agatha & Strüder-Kypke 2007). Thus, the complete replacement of the somatic ciliature is very likely a plesiomorphy in the last common ancestor of the Hypotricha.1 This hypothesis requires the assumption that the preservation of parental ciliature in the oligotrichs is an apomorphy; however, perhaps my interpretation is incorrect. In various hypotrichs parts of the parental ciliature (e.g., marginal rows in Coniculostomum or parts of the dorsal ciliature in Parakahliella; Kamra & Sapra 1990, Berger & Foissner 1989a, Berger et al. 1985a) are retained in the post-dividers. These specific features have to be interpreted as apomorphies of the individual groups. 1
Erroneously, I listed this feature as apomorphy for the Hypotricha in a recent abstract (Berger 2008).
PHYLOGENY
37
Silverline system fine-meshed. The silverline system of the Hypotricha is finemeshed (for reviews, see Berger 1999, 2006). The same is true for oligotrichs (e.g., Foissner et al. 1991, 1999) and Diophrys (e.g., Curds & Wu 1983, Agamaliev 1983, Alekperov & Asadullayeva 1999, p. 224) showing that this pattern is a plesiomorphy for the last common ancestor of the Hypotricha. Three-layered cyst wall. Oxytrichids have a four-layered cyst wall according to the review by Gutiérrez et al. (2003). Non-Dorsomarginalia, that is, hypotrichs branching off outside the Dorsomarginalia have only three layers; the same is true for Diophrys (Walker & Maugel 1980, Gutiérrez et al. 2003). By contrast, true euplotids (e.g., Euplotes) and many taxa outside the spirotrichs, have only two layers, strongly indicating the last common ancestor of the hypotrichs had the plesiomorphic number of three layers. No detailed data are available for the oligotrichs (e.g., Agatha & Strüder-Kypke 2007). Perilemma present. This unit-membrane like layer covers the whole cell of the oligotrichs and hypotrichs (e.g., Laval-Peuto 1975, Bardele 1981, WirnsbergerAescht et al. 1989, Wasik & Mikolajczyk 1992, Modeo et al. 2003, Agatha 2004, Foissner 2005, Foissner & Pichler 2006). It is obviously lacking in the euplotids and other ciliates (Bardele 1981). Bardele (1981) and other workers also reported a lack of the perilemma in Halteria. This is difficult to explain, inasmuch as it is present in Meseres, the closest relative (e.g., Katz et al. 2005, Foissner & Pichler 2006). Only recently, this structure was attested for Halteria by Foissner et al. (2007, p. 311). However, a perilemma is very likely already present in Diophrys because Raffaelli (1970, p. 233) wrote “The pellicle is composed of two membranes: a double outer membrane and a plasmatic membrane without anything between the two. Such a pellicle is different from that described in several other species of Euplotes”. This strongly indicates – together with some other features (e.g., two undulating membranes) – that Diophrys is not a euplotid, but branched off later within the spirotrichs (see chapter 2.3). Unfortunately, no decision can be made about the presence/absence of a perilemma in the discocephalids because relevant data are lacking (Wicklow 1982b). However, if Fig. 6a is correct and if Diophrys indeed has a perilemma then the discocephalids very likely also have such a membrane. Anyhow, the perilemma is a young plesiomorphy in the last common ancestor of the Hypotricha. As discussed in the previous paragraph, the sistergroup relationship of the oligotrichs and hypotrichs is not only indicated by most molecular analyses, but also by the presence of a perilemma. Thus, the taxon comprising the Oligotricha and Hypotricha is named Perilemmaphora1 tax. nov.: Spirotricha with a perilemma. Perhaps this taxon also includes Diophrys and the discocephalids (Fig. 6a); however, further ultrastructural studies on these two taxa are needed. Recently, Foissner et al. (2007) proposed the name Oligotrichia for the group oligotrichs + hypotrichs. By contrast, Lynn & Small 1
Perilemmaphora is a composite of perilemma, the main (sole?) morphological apomorphy, and the Greek noun phor·a (carrier; owner of a feature), meaning ciliates bearing a perilemma. Perilemma itself is a composite of the Greek prefix peri+ (around) and the Greek noun lemm- (cover, sheath). Note that the term perilemma is also used, for example, for the tissue surrounding the nervous system of insects (e.g., Ashhurst 1959).
38
GENERAL SECTION
(2002, p. 394, 465) unified in the taxon Oligotrichia the Halteriida and the Strombidiida. Adl et al. (2005, p. 436) mentioned as representatives of the Oligotrichia Cyrtostrombidium, Laboea, and Strombidium, that is, the Choreotrichia and Stichotrichia (= Hypotricha in my terminology) including the halterids are not included. This shows that the name Oligotrichia comprises a very different number of taxa, depending on the system used. By contrast, the Perilemmaphora comprise those spirotrichs which have a perilemma (Fig. 6a). UGA used as stop codon and UAA and UAG translated to glutamine. Details see plesiomorphy “two undulating membranes” and Table 3a. Telomeric Repeat Sequence TTTTGGGG. Telomeres are specialised DNAprotein complexes at the ends of eukaryotic chromosomes (e.g., Blackburn 1990, Berg et al. 2003, Seyffert 2003). Oxytricha, Stylonychia, and Euplotes have the sequence TTTTGGGG (e.g., Klobutcher et al. 1981). By contrast, the sequence is TTGGGG in Tetrahymena and Glaucoma or TTGGG(T/G) in Paramecium (e.g., Fanti & Pimpinelli 1999). Anyhow, the sequence TTTTGGGG is obviously a plesiomorphy for the last common ancestor of the Hypotricha. Benthic modus vivendi. Almost all hypotrichs are benthic, that is, limnetic and marine species are bottom-dwellers creeping on detritus, stones, or macrophytes. Only few taxa became real planktonts, for example, Hypotrichidium, Pelagotrichidium, Pseudostrombidium, Spiretella (for reviews, see Berger 1999, p. 494 and Foissner et al. 1999, p. 677ff). By contrast, almost all oligotrichs are obligatorily pelagic, that is, the oligotrichs are the planktontic part of the spirotrichs. Agatha & Strüder-Kypke (2007) did not use this feature in their phylogenetic analyses, but included it in the character cell shape; accordingly, they interpreted the globular or obconical shape, which is due to the planktonic life, as apomorphy of the oligotrichs (= Oligotrichea in their paper). I agree with this assumption. By contrast, the euplotids, Diophrys, and the discocephalids are – like the hypotrichs – benthic, showing that this mode of life is a plesiomorphy for the Hypotricha. Whether the last common ancestor of the hypotrichs was marine or limnetic remains uncertain. The soil was likely colonised several times independently within various higher taxa of the hypotrichs, for example, by Keronella within the urostyloids (Berger 2006, p. 1020), by Rigidocortex within the stylonychines (Berger 1999, p. 717), and by Lamtostyla within the amphisiellids.
2.2.3 Features not Considered in the Ground Pattern For the following features the state (apomorphic or plesiomorphic) in the last common ancestor of the Hypotricha is difficult to estimate, for example, because relevant data are lacking either for the hypotrichs or for the out-groups. In some other cases the situation is rather tricky, making a reliable estimation difficult. Thus, they are not yet considered in the ground pattern. Of course there are many other features which will be more or less useful for the estimation of the phylogeny sooner or later. Two macronuclear nodules. All hypotrichs have a macronucleus composed of two or more nodules (for reviews, see Kahl 1932, Berger 1999, 2006). Species which have
PHYLOGENY
39
the two nodules only indistinctly separated are rare (e.g., Orthoamphisiella breviseries Foissner, Agatha & Berger, 2002). Phacodinium, Euplotes, and many oligotrichs have, like many other ciliates, only one macronucleus, indicating that the stem-species of the spirotrichs had a single macronucleus. However, the situation within the euplotids and oligotrichs is rather diverse, that is, species with one or more macronuclear nodules exist (e.g., Curds & Wu 1983, Foissner et al. 1999, Agatha & Strüder-Kypke 2007, Miao et al. 2007). In addition, Diophrys (e.g., Song & Packroff 1993, Song et al. 2007) and discocephalids (e.g., Wicklow 1982b, Lin et al. 2004) have two or more nodules. This indicates that within the spirotrichs two or more nodules evolved convergently. A fragmentation of the macronucleus, likely not a very complex feature, is also known from many non-spirotrich ciliates, for example, the colpodids (e.g., Foissner 1993) or haptorids (e.g., Lynn & Small 2002, Foissner & Xu 2007). Agatha & Strüder-Kypke (2007) assumed that a macronucleus composed of two nodules is a plesiomorphy for the euplotids, the oligotrichs, and hypotrichs, that is, they obviously defined two nodules as apomorphy for the spirotrichs. Accordingly, species with only one macronucleus either lost one nodule or the two nodules fused to a single mass. Since both hypotheses result in a rather high number of homoplasies I preliminarily put the number of macronuclear nodules in the list of features not used in the ground pattern. In the monograph of urostyloids I (likely incorrectly) supposed that two macronuclear nodules are an apomorphy for the Hypotricha (Berger 2006, p. 33). Borror & Wicklow (1983) in their revision on urostyloids supposed that a high number of macronuclear nodules is the “primitive” (= plesiomorphic) condition. However, this hypothesis is very likely incorrect. Resting cyst. Walker & Maugel (1980) proposed a classification system based on the presence/absence of basal bodies, cilia, and cortical microtubules in the resting cyst. They distinguished two types, namely NKR-cysts (non-kinetosome-resorbing) and KR-cysts (kinetosome-resorbing). Later, Matsusaka et al. (1989) found the PKR-type (partially-kinetosome-resorbing). Within the hypotrichs two types occur, namely the PKR-type in taxa which mainly branch off outside the Dorsomarginalia (e.g., Anteholosticha adami, Pseudourostyla levis, Gonostomum affine) and the KRtype in the oxytrichids.1 Unfortunately, no details about resting cysts of oligotrichs have been available so far (Agatha & Strüder-Kypke 2007, p. 52). Only recently, Foissner et al. (2007, p. 308) reported that Rimostrombidium has a PKR or KR cyst. Meseres, a halteriid, also has a KR-cyst (e.g., Foissner 2005, Foissner et al. 2005a, Foissner & Pichler 2006, Foissner et al. 2006), whereas euplotids have NKR cysts (e.g., Rawlinson & Gates 1985, 1986, Martin-Gonzalez et al. 1992). Foissner et al. (2007, p. 308) supposed that the PKR and KR cysts represent the plesiomorphic state within the spirotrichs because they are more common in this group than the NKR cysts. Obviously they made an ingroup comparison, a method usually not suitable to decide between an apomorphic and plesiomorphic state (Ax 1984, p. 131; 1
Grim & Manganaro (1985, p. 357) assigned the cyst of Pseudourostyla cristata (Urostyloidea; Berger 2006) to the NKR-type (see Martin-Gonzalez et al. 1992 for review). However, they studied only early (young) cysts so that this feature must not be overinterpreted.
40
GENERAL SECTION
Wägele 2001, p. 172). Phacodinium, a very near relative of the group euplotids + Perilemmaphora, has – like the euplotids – a NKR cyst (Calvo et al. 1992). When Phacodinium is used as outgroup then one has to postulate that the NKR cyst is the plesiomorphic type within the spirotrichs. Consequently, the PKR/KR type has to be interpreted as novelty in the stemline of the Perilemmaphora. Perhaps most non-spirotrich ciliates have NKR cysts (e.g., Rawlinson & Gates 1986, Walker et al. 1989, Repak & Anderson 1990, Calvo et al. 2003). In colpodids the oral ciliature is resorbed while the somatic kinetosomes are retained (e.g., Ruthmann & Kuck 1985, Martin-Gonzalez et al. 1992, Foissner 1993, Díaz et al. 2000). Of course there are several other features of the resting cysts which are perhaps good phylogenetic markers (Foissner et al. 2007). However, more taxa have to be studied in detail. Recently, Müller (2007) found that Meseres has the same type of emergence from the cyst like oxytrichids. Phago-assistant membrane or buccal seal. This structure covers the buccal cavity of the Hypotricha (for details, see Sui et al. 2003 and Foissner & AL-Rasheid 2006). Unfortunately, the situation outside the Hypotricha is unclear so that it is – at the present state of knowledge – not possible to estimate the state in the last common ancestor of the hypotrichs. I found no comment whether this membrane is the same as the perilemma which covers the whole cell in hypotrichs and oligotrichs. Detailed transmission electron microscopical investigations are needed to clear up the situation. Distal end of adoral zone of membranelles. The distal end of the adoral zone ends at variable position in the Hypotricha, that is, it extends almost not (e.g., Gonostomum; for review, see Berger 1999) to distinctly (e.g., Retroextendia; Berger 2006, p. 732) onto the right body margin. Oligotrichs have a more or less “circular” adoral zone due to the frontal location (e.g., Kahl 1932, Agatha 2004) as apomorphy. In euplotids and Phacodinium the distal end of the adoral zone does not usually extend far posteriorly, whereas in Diophrys scutum the DE-value is 0.44, that is, rather high (Berger 2006, p. 18). By contrast, in the discocephalids the adoral zone almost encircles the characteristic head, possibly an apomorphy of this group. Reorganisation of parental adoral zone of membranelles. This is a highly interesting feature, but difficult to interpret. In the hypotrichs, no (e.g., many [all?] oxytrichids, Holosticha) or some (e.g., Eschaneustyla, Maregastrostyla) parental membranelles are (clearly recognisably) reorganised or the whole parental adoral zone is replaced, for example, in the pseudokeronopsids (for reviews, see Berger 1999, 2006) and Trachelostyla (Shao et al. 2007). In the oligotrichs and the euplotids the parental oral structures are not reorganised (e.g., Agatha & Strüder-Kypke 2007, Voss 1989, Petz & Foissner 1993). By contrast, the parental oral apparatus of Diophrys and the discocephalids is partially redifferentiated (Wicklow 1982b, Song & Packroff 1993). Thus, the situation in the last common ancestor of the Hypotricha is difficult to estimate. I suppose that there no or only the proximal-most membranelles are reorganised. Frontal-ventral-transverse cirri anlagen of proter and opisthe develop independently. This feature is present in Euplotes (e.g., Wise 1965, Voss 1989), the Urostyloidea (for review, see Berger 2006), Trachelostyla (Shao et al. 2007), and many
PHYLOGENY
41
Dorsomarginalia (e.g., Berger 1999). By contrast, in several Dorsomarginalia (e.g., Urosoma; Berger 1999) and some amphisiellids (present book), but also in some euplotids (e.g., Uronychia; Hill 1990, Song et al. 2004), Diophrys (e.g., Song & Packroff 1993), and discocephalids (Wicklow 1982b), at least some anlagen of the proter and the opisthe have a common origin, that is, are formed via so-called primary primordia. Since the oligotrichs have lost the cirri, this group cannot be used as outgroup. Consequently, at present I am not able to estimate the state of this feature in the last common ancestor of the Hypotricha. Frontal-ventral-transverse cirri mature in an anterior to posterior direction. The formation of frontal-ventral-transverse cirri proceeds from anterior to posterior in the Hypotricha (for reviews, see Berger 1999, 2006, present book). By contrast, in euplotids and Diophrys each streak matures in a posterior to anterior direction (e.g., Wicklow 1983a, Voss 1989, Song & Packroff 1993, Song et al. 2004). No detailed information about this trait is available for the discocephalids (Wicklow 1982b) making an outgroup comparison impossible. Thus, we do not yet know when the reversa of the maturation direction occurred. Length of dorsal bristles. Most hypotrichs have dorsal bristles which are less than 5 µm long; very often they have a length of only 2–3 µm. The same is true for most euplotid and Diophrys species (e.g., Curds & Wu 1983, Song & Packroff 1993, Song et al. 2007). However, exceptions exist, for example, Diophrys hystrix where the bristles are about 15 µm long (e.g., Song et al. 2007). Most discocephalids have relatively long bristles, that is, between 5 µm and 10 µm (Wicklow 1982b, Lin et al. 2004). Oligotrichs usually have short bristles (e.g., Foissner et al. 1999, Agatha & Strüder-Kypke 2007). Probably the last common ancestor of the Hypotricha also had short bristles. Kinetodesmal fibres. These fibres are resorbed in late dividers and are therefore lacking in dorsal dikinetids of interphasic specimens of the Hypotricha. By contrast, they are present in euplotids (for review, see Lynn 1991). In the oligotrichs the situation is rather complicated (Agatha 2004): a short kinetodesmal fibre is present in the tintinnid Cyttarocylis brandti (Laval-Peuto 1994, p. 193), but is lacking in another tintinnid (Petalotricha) and Strobilidium velox (for review, see Agatha 2004). Agatha & Strüder-Kypke (2007) did not discuss this feature. For the cladistic analysis, Agatha (2004) assumed a lack of this fibre in morphostatic Oligotrichida. Since she proposed a sister-group relationship of the hypotrichs and euplotids she had to presume a convergent loss of the kinetodesmal fibres in the hypotrichs and oligotrichs (= Oligotrichea in her paper). Because of the uncertain situation in the oligotrichs it is not possibly to estimate whether the lack of the kinetodesmal fibres is an apomorphy or a young plesiomorphy for the last common ancestor of the Hypotricha. Kinetodesmal fibres are permanently present in the discocephalids (Wicklow 1982b), strongly indicating that this group branched off outside the hypotrichs, that is, discocephalids are very likely no hypotrichs. This morphological argumentation is supported by recent molecular data (Miao et al. 2007). Micronuclear DNA polymerase alpha genes not scrambled. Chang et al. (2003) found that the micronuclear DNA polymerase alpha genes of the urostyloids Uro-
42
GENERAL SECTION
styla grandis and Holosticha kessleri (= H. gibba in Berger 2006) are not scrambled. On the other hand, they found that in Uroleptus sp. and Paraurostyla weissei these genes are scrambled with a configuration similar to that in the oxytrichids Stylonychia lemnae and Oxytricha. Thus, they suggested that the DNA polymerase alpha gene became scrambled in a species diverging later than Holosticha (that is, urostyloids), but earlier than Uroleptus. This molecular feature therefore supports the Dorsomarginalia (Fig. 9a; Berger 2006). Obviously the unscrambled configuration of the urostyloids is the plesiomorphic state. I found no data about the state in the oligotrichs or outside the Perilemmaphora (oligotrichs + hypotrichs). Micronuclear actin I gene MDSs (macronuclear destined segments) not scrambled. The micronuclear actin I gene of Urostyla grandis and Engelmanniella mobilis consists of three and four non-scrambled MDSs, respectively (Hogan et al. 2001). Dalby & Prescott (2004) concluded that the non-scrambling is the plesiomorphic state. By contrast, the actin I gene became scrambled both in the Uroleptus and the Oxytrichidae branch, but in two different, completely unrelated patterns (Dalby & Prescott 2004). Thus, the “actin I gene scrambling type Uroleptus” is obviously an apomorphy for the Uroleptus group and the “actin I gene scrambling type Oxytrichidae”, beside the fragmentation of dorsal kinety 3, a further apomorphy of the Oxytrichidae cluster (Berger 2006).
2.3 Comments on the Evolution within the Spirotricha According to molecular data the three major taxa of the spirotrichs (euplotids, oligotrichs, hypotrichs) are related as shown in Fig. 6b. Diophrys and discocephalids are usually considered as euplotids (e.g., Wicklow 1982b, Lynn & Small 2002, Adl et al. 2005, Song et al. 2007). However, these two taxa differ in several features from the euplotids, strongly indicating that this assumption is incorrect. The following features indicate that Diophrys is not a euplotid, but branched off later in the spirotrich tree (Fig. 6a; for a more detailed discussion of the features, see chapter 2.2.2):1 (i) Two undulating membranes: Diophrys has, like the discocephalids and the hypotrichs, two distinct undulating membranes (e.g., Song & Wilbert 1993, Wicklow 1983b, Lin et al. 2004, Berger 1999, 2006, present book). The oligotrichs have lost one membrane in the last common ancestor, which is therefore an apomorphy for this taxon (Agatha & Strüder-Kypke 2007). By contrast, the true euplotids very likely never had a second membrane in their stem-lineage (e.g., Curds & Wu 1983, Wicklow 1983a, Foissner et al. 1991, Lin & Song 2004). (ii) The adoral membranelles mature in an anterior to posterior direction in all hypotrichs (e.g., Berger 1999, 2006, present book) and probably oligotrichs (e.g., Agatha 2003, Agatha et al. 2004), but also in discocephalids (Wicklow 1982b) and Diophrys 1
The halteriids (Halteria, Pelagohalteria, Meseres) are not considered because it is not yet clear whether they are oligotrichs or hypotrichs.
PHYLOGENY
43
(Song & Packroff 1993). By contrast, membranelles are formed in the opposite direction in Euplotes (e.g., Voss 1989). (iii) Euplotes uses other stop codons than Diophrys, Oxytricha, Stylonychia, and Paraurostyla (Prescott 1994, Perez-Romero et al. 2001, Table 3a). Further details, see chapter 2.2.2, plesiomorphy “two undulating membranes”. (iv) The silverline system of Diophrys is, like that of hypotrichs and oligotrichs, fine-meshed (see plesiomorphies above). By contrast, Euplotes, Cytharoides, Aspidisca, and likely other euplotids have a wide-meshed system (e.g., Klein 1936, Foissner et al. 1991, Petz et al. 1995). (v) The somatic parental ciliature is resorbed during cell division in hypotrichs, Diophrys (e.g., Song & Packroff 1993), and the discocephalids (Wicklow 1982b). By contrast, in the euplotids and the oligotrichs, and in ciliates in general, at least parts of the parental somatic ciliature are present in the post-dividers (e.g., Berger et al. 1983, Foissner 1996, Agatha 2004, Agatha & Strüder-Kypke 2007). (vi) Hypotrichs and oligotrichs have a perilemma. Very likely, Diophrys also has such a membrane. By contrast, a perilemma is lacking in the euplotids and other ciliates (details see plesiomorphy above). (vii) In phylogenies based on gene sequences Diophrys is often not in the euplotid cluster, but is more closely related to the group formed by the oligotrichs + hypotrichs (e.g., Song et al. 2004, their Fig. 37; Shao et al. 2006). Chen & Song (2002) also found such a relationship, but did not include the oligotrichs into their analyses. Interestingly, Uronychia is often very close to Diophrys in molecular trees, although they differ distinctly in the morphology. Very likely there are further differences between the euplotids and Diophrys. It is beyond the scope of this book to make a detailed phylogenetic analysis of the spirotrichs; however, the brief review shows that there are several morphological, ontogenetic, ultrastructural, and molecular features which do not support the classification of Diophrys in the euplotids. Chen & Song (2002) came to a similar conclusion based on their gene sequence data, but did not provide background information. Only recently, Foissner et al. (2007) provided a very detailed analysis of the phylogeny of the core spirotrichs using morphological features. The systematic position of the discocephalids is not known in detail. As mentioned above, they are usually classified as subgroup of the euplotids (e.g., Lynn & Small 2002, p. 423; Adl et al. 2005, p. 436). Since these species are rare, relatively little is known about their morphology and cell division (e.g., Wicklow 1982b, Lin et al. 2004). Previously I assigned the discocephalids to the Hypotricha because they have two undulating membranes (Berger 2006, p. 30, 77). However, this foundation was incorrect because based on a plesiomorphy, and not on an apomorphy; I overlooked that already Diophrys has an oral apparatus closely resembling that of the hypotrichs (Fig. 8a). The presence of kinetodesmal fibres in morphostatic specimens (Wicklow 1982b) strongly indicates that the discocephalids branch off outside the hypotrichs, which lack this fibre. The following features indicate that the discocephalids are more closely related to the group oligotrichs + hypotrichs than to the
44
GENERAL SECTION
euplotids (for details, see plesiomorphies discussed above): (i) two frontoterminal cirri; (ii) two pretransverse ventral cirri; (iii) distinct marginal rows; (iv) adoral membranelles mature in posteriad direction; (v) oral primordium originates on cell surface; (vi) parental somatic ciliature completely replaced during cell division. Just recently, Miao et al. (2007; their Fig. 8) also demonstrated that Prodiscocephalus borrori, a discocephalid, is the next relative of the group oligotrichs + hypotrichs. By contrast, in the tree published by Shao et al. (2007), P. borrori clusters in the euplotids (Hypotrichia in their terminology) revealing that the phylogenetic position of the discocephalids is still uncertain. Unfortunately, no data are available about the presence/absence of the perilemma in the discocephalids because the study by Wicklow (1982b) did not deal with this feature. However, if a perilemma is already present in Diophrys (see this feature above; Raffaelli 1970) then such a structure is very likely also present in the discocephalids.
2.4 Comments on the Evolution within the Hypotricha Berger & Foissner (1997) and Berger (1999) hypothesised the 18-frontal-ventraltransverse cirri pattern as apomorphy of the Oxytrichidae. Eigner (1997, p. 553, 570) assumed that this pattern evolved twice or more times independently. However, this is extremely unlikely because it is too complex. In 2004 we published the CEUU-hypothesis which explains the convergent evolution of the midventral cirral pattern in urostyloids and Uroleptus (Foissner et al. 2004) and just recently I hypothesised that the 18-cirri pattern is the major morphological apomorphy of the taxon Hypotricha (Berger 2006, p. 33). In addition, Berger (1999) and Berger (2006) defined the Oxytrichidae mainly via the oxytrichid dorsal kinety fragmentation (kinety 3 fragments) and Berger (2006, p. 38) established the Dorsomarginalia for all Hypotricha with a dorsomarginal kinety as morphological apomorphy. This indicates that the ventral cirral pattern is no longer the sole feature for the major classification of hypotrichs. Neokeronopsis is an impressive example where the ventral cirral pattern feigns an incorrect, namely, urostyloid origin (Wang et al. 2007). However, only if the correct features are used at a certain level will morphological and molecular data come to the same classification. Neokeronopsis has dorsomarginal kineties and a dorsal kinety fragmentation so that it unequivocally belongs to the Oxytrichidae (Berger 2006). Since it has a flexible body and distinct cortical granules it cannot belong to the Stylonychinae (Berger 1999) and the Cyrtohymena-like undulating membranes indicate that it is closely related to flexible Cyrtohymena species (Berger 2006, p. 1191). Berger & Foissner (1997) and Berger (1999) divided the 18-cirri oxytrichids into the Stylonychinae (e.g., rigid body, lack of cortical granules) and Oxytrichinae with the participation of the postoral ventral cirrus V/3 in primordia formation as apomorphy. However, while the stylonychines are confirmed by all molecular analyses
PHYLOGENY
45
(e.g., Hewitt et al. 2003, Foissner et al. 2004, Shao et al. 20071), the oxytrichines reveal as paraphyletic group. Paraphyletic groups are based on plesiomorphies, indicating that the features used to define the oxytrichids and oxytrichines were not appropriate at those levels. In molecular trees, 18-cirri hypotrichs occur throughout the Hypotricha tree (e.g., Schmidt et al. 2007). This agrees with my assumption that this pattern already evolved in the stem-line of the Hypotricha (Berger 2006, Fig. 14a), that is, 18-cirri hypotrichs can occur at all sites of the Hypotricha tree. In addition, all major cirral patterns can be more or less easily derived from this type. In Gonostomum, 18-cirri hypotrichs with anteriorly located postoral ventral cirri (for review, see Berger 1999), the dorsal kineties are formed in the same simple way as in the urostyloids and amphisiellids. Previously, when we classified Gonostomum as oxytrichid, we postulated a loss of both the oxytrichid dorsal kinety fragmentation and dorsomarginal rows (Berger & Foissner 1997, Berger 1999). This classification was mainly based on the assumption that the 18 frontal-ventral-transverse cirri pattern is an apomorphy for the oxytrichids. By contrast, the present hypothesis allows a much more parsimonious explanation of the simple dorsal kinety pattern of Gonostomum. Thus, Gonostomum very probably belongs to the non-dorsomarginalian hypotrichs (Fig. 7a) and not to the oxytrichids as supposed previously (Berger 1999). A rather basal branching off of Gonostomum is also indicated by recent molecular analyses (e.g., Schmidt et al. 2007, Shao et al. 2007). Previously, I classified 18-cirri hypotrichs with dorsomarginal kineties and lacking kinety fragmentation (e.g., Urosomoida, Urosoma) or without dorsomarginal kinety and without fragmentation (e.g., Gonostomum) in the oxytrichids (Berger & Foissner 1997, Berger 1999). We assumed that the absence of these features is due to loss. Now I suppose that the simple dorsal kinety formation pattern present in Gonostomum (only intrakinetal proliferation) is the state already present in the last common ancestor of the Hypotricha. In a stem-line within the Hypotricha a dorsomarginal kinety evolved resulting in the Dorsomarginalia, a (hopefully monophyletic) group comprising a major part of the Hypotricha (Berger 2006). Oxytrichid dorsal kinety fragmentation, the main apomorphy of the Oxytrichidae (Berger 1999, 2006; Fig. 9a), originated (very likely) within one branch of the Dorsomarginalia. As a consequence of this new system, I have to assume that Gonostomum branched off outside the Dorsomarginalia, and Urosomoida and Urosoma outside the Oxytrichidae, but within the Dorsomarginalia. Since the 18-frontal-ventral-transverse cirri pattern is hypothesised as apomorphy of the Hypotricha, the presence of this pattern cannot be used to define a subgroup of the Hypotricha, for example, the Oxytrichidae. Of course, details of this 18-cirri pattern – for example, the special ar1
Note that these authors use an outdated classification because they assigned Pattersoniella and Gastrostyla to the amphisiellids. However, Pattersoniella was already assigned to the stylonychines by Berger in 1999 because of the rigid body and the oxytrichid kinety fragmentation. This classification was later confirmed by molecular data (e.g., Bernhard et al. 2001). Gastrostyla was assigned to the oxytrichids by Berger (1999) mainly because of the oxytrichid dorsal kinety fragmentation; later we recognised that it belongs to the stylonychines (Foissner et al. 2004).
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GENERAL SECTION
Fig. 9a Diagram of supposed phylogenetic relationships within the Hypotricha (original). Paraphyletic groups marked by quotation marks. The question marks indicate the uncertainty in the classification of the “non-xy taxa”. At the present state of knowledge it is impossible to estimate which taxa branched off first within a certain group. Main morphological autapomorphies (squares 1–4): 1 – 18 frontal-ventraltransverse cirri; three dorsal kineties; three caudal cirri; contractile vacuole at left body margin; high support by molecular data (see also 9 in Fig. 6a). 2 – dorsomarginal kineties present. 3 – fragmentation of dorsal kinety 3. 4 – body rigid; adoral zone of membranelles m40% of body length; cortical granules lacking; high support by molecular data. For details see chapter 2.4 and Berger (2006, p. 33). Well-known representatives: Stylonychinae: Coniculostomum, Gastrostyla, Histriculus, Laurentiella, Onychodromus, Pattersoniella, Pleurotricha, Rigidocortex, Steinia, Sterkiella, Stylonychia, Styxophrya, Tetmemena “Non-stylonychine Oxytrichidae”: Apoamphisiella, Australocirrus, Cyrtohymena, Neokeronopsis, Notohymena, Oxytricha, Paraurostyla, Pseudouroleptus, Rubrioxytricha, Territricha “Non-oxytrichid Dorsomarginalia”: Hemiurosoma, Nudiamphisiella, Parakahliella, Uroleptus, Urosoma, Urosomoida, Vermioxytricha “Non-dorsomarginalian Hypotricha”: Amphisiellidae, Gonostomum, Trachelostylidae, Urostyloidea, Wallackia
rangement of the frontoventral cirri in Urosoma-species – are usable markers at certain levels. The Oxytrichidae are now defined via the oxytrichid dorsal kinety fragmentation. Future studies will show whether this proposal allows a better explanation of the phylogeny of the Hypotricha than previous systems. Urostyloids lack dorsomarginal kineties, indicating that they branched off very early in the Hypotricha tree (Berger 2006; Fig. 9a), a hypothesis basically confirmed by gene sequence analysis (e.g., Foissner et al. 2004, Schmidt et al. 2007, Shao et al. 2007). According to the analyses by Schmidt et al. (2007), the urostyloids are not
PHYLOGENY
47
monophyletic which is not impossible from the morphological point of view if we consider that the zigzagging cirral pattern evolved several times independently (e.g., urostyloids, Uroleptus, Neokeronopsis, Pattersoniella). However, some other nonmonophyletic taxa discussed by Schmidt et al. (2007) are simply due to misinterpretations. For example, Pseudokeronopsis qingdaoensis, which does not cluster with three other Pseudokeronopsis species, is a junior synonym of Oxytricha crassa, assigned to Thigmokeronopsis by Berger (2006, p. 873) and just recently transferred to Apokeronopsis by the Song group. Consequently, the molecular date do not contradict the morphological data, but confirm them. In the tree provided by Shao et al. (2007) the urostyloids also form a monophyletic group. The amphisiellids, the major group revised in the present book, is characterised by the so-called amphisiellid median cirral row (Fig. 2a). This row originates from anlagen IV, V, and VI or V and VI, due to an increase of the number of cirri formed per anlage (Fig. 5a). Amphisiellids form their cirri, like the 18-cirri hypotrichs, from six (I–VI) anlagen and lack dorsomarginal kineties and dorsal kinety fragmentation. Consequently, I have to assume that they branch off, like the urostyloids and some other taxa (e.g., Gonostomum), outside the Dorsomarginalia. According to Schmidt et al. (2007), Amphisiella magnigranulosa (= Uroleptoides magnigranulosus in present book) clusters with some urostyloids, a classification difficult to explain from the morphological point of view. However, at least it does not cluster with species having a dorsomarginal row or even a dorsal kinety fragmentation, indicating the groups based on the dorsal ciliature have a relevance. Recently, Gong et al. (2007) found a close relationship of Amphisiella annulata and Hemigastrostyla enigmatica (details see Addenda). Fig. 9a shows the possible relationships within the hypotrichs. Note that the Stylonychinae, Oxytrichidae, Dorsomarginalia, and Hypotricha are not only defined via morphological traits, but also by various molecular features (for review, see Berger 2006). I preliminarily eliminate the paraphyletic group Oxytrichinae (Berger & Foissner 1997, Berger 1999), which is obviously an artificial assemblage of flexible 18-cirri hypotrichs. Since both features (flexible body, 18-cirri pattern) are plesiomorphies within the hypotrichs, they cannot be used to elucidate the phylogeny within the hypotrichs. At the present state of knowledge it is difficult to estimate the “exact” position of many taxa, for example, Uroleptus, the amphisiellids, the urostyloids. For Uroleptus, for example, we can only say that (i) it is very likely not an oxytrichid because it lacks the oxytrichid dorsal kinety fragmentation; (ii) it is very likely not an urostyloid because it has a dorsomarginal row. Thus, I (preliminarily) classify Uroleptus as “non-oxytrichid Dorsomarginalia” (Fig. 9a). Since both amphisiellids and urostyloids lack an oxytrichid dorsal kinety fragmentation as well as a dorsomarginal kinety, they are (preliminarily) assigned to the “non-dorsomarginalian Hypotricha”. Gonostomum-like hypotrichs very likely also belong to this group, a position also supported by certain molecular trees (Schmidt et al. 2007, Shao et al. 2007). Interestingly, Gonostomum, Trachelostyla (branches off as first species in the molecular
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GENERAL SECTION
Hypotricha tree provided by Schmidt et al. 2007), and some amphisiellids (e.g., Lamtostyla granulifera, Fig. 3a) have the postoral ventral cirri right of the proximal portion of the adoral zone (Fig. 4a). Perhaps this is the position already present in the last common ancestor of the Hypotricha (Fig. 7a) because in euplotid-like ancestors almost all cirri were right of the adoral zone due to the large relative length of this structure. Consequently, the “postoral” position of the postoral ventral cirri would be a feature which later evolved when the relative length of the adoral zone became lower.
3
Previous Classifications and Revisions
The “family Amphisiellidae” was established three times, namely by Jankowski (1979), Hemberger (1982), and Small & Lynn (1985). The original classifications of the Amphisiellidae and Trachelostylidae and some later systems are presented in Tables 4–14. I did not change the original presentations, that is, (i) the sequence of the genera; (ii) the spelling of the genus and author(s) names; and (iii) the year of publication are as in the papers. For a brief discussion of the schemes shown in the tables below, see the systematic section. Several genera – although classified in the amphisiellids or trachelostylids by some authors – are not considered in the present monograph, except when they belong to higher taxa already reviewed by Berger (1999, 2006). For a foundation of the exclusions, see “Taxa not considered” (p. 466, 512). There is no modern, detailed guide to the species of the Amphisiellidae. The reviews by Kahl (1932, 1933) contain only the “subgenus Holosticha (Amphisiella)” because most terrestrial species, the major part of the amphisiellids, have only been discovered in the last decades. Consequently, almost all terrestrial species have been described in more or less detail after silver impregnation. For a discussion of the classification used in the present book, see the systematic section, as well as the phylogeny section (chapter 2). Briefly, amphisiellids are, according to Eigner & Foissner (1994), hypotrichs with primarily six frontal-ventraltransverse cirri anlagen and a more or less distinct frontoventral row, the so-called amphisiellid median cirral row, which is composed of cirri of anlagen (from anterior to posterior) VI and V or VI, IV, and V. In addition, dorsal kineties originate only by intrakinetal proliferation. Consequently, all taxa which show dorsomarginal kineties and/or oxytrichid kinety fragmentation have been excluded from the amphisiellids. The trachelostylids are obviously a small group of marine 18-cirri hypotrichs, inter alia, with anteriorly displaced postoral ventral cirri and a more or less distinct head. In addition, the present review contains some genera of very uncertain phylogenetic position, for example, Apourosomoida. For classification of the Amphisiellidae and the other taxa reviewed in the present book, see table of content.
CLASSIFICATION Table 4 Classification of amphisiellid ciliates according to Jankowski (1979) Family Amphisiellidae fam. n. Amphisiella
Table 5 Classification of amphisiellid ciliates according to Hemberger (1982) Family Amphisiellidae n. fam. Amphisiella Gourret & Roeser, 1888 Cladotricha Gajewskaja, 1925 Epiclintes Stein, 1862 Kahliella Corliss, 1960 Paraurostyla Borror, 1972 Pseudouroleptus n. gen. Trachelochaeta Srámek-Husek, 1954 Uroleptoides Wenzel, 1953 Incertae sedis Balladyna Kowalewski, 1882 Banyulsella Dragesco, 1953 Lacazea Dragesco, 1960
Table 6 Classification of amphisiellid ciliates according to Small & Lynn (1985) Family Amphisiellidae n. fam. Amphisiella Eschaneustyla Kahliella Onychodromopsis Onychodromus Paraurostyla
Table 7 Classification of amphisiellid ciliates according to Foissner (1988) Amphisiellidae Jankowski, 1979 Amphisiella Gourret & Roeser, 1888 Amphisiellides nov. gen. Paramphisiella nov. gen. Hemiamphisiella nov. gen
Table 8 Classification of amphisiellid ciliates according to Tuffrau & Fleury (1994) Family Amphisellidae Jankowski, 1979 Amphisiella Gourret et Roeser 1888 Balladyna Kowalewski 1882 Balladynella Stiller 1974 Coniculostomum Njiné 1978 Onychodromopsis Stokes 1887 Paraurostyla Borror 1972 Parurosoma Gelei 1954 Psammomitra Borror 1972
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GENERAL SECTION
Table 8 Continued Territricha Berger et Foissner 1988 Trachelochaeta Sramek-Husek 1954 Trachelostyla Kahl 1932 Wallackia Foissner 1976
Table 9 Classification of amphisiellid ciliates according to Eigner & Foissner (1994) Family Amphisiellidae Jankowski, 1979 Amphisiella Gourret & Roeser, 1888 Amphisiellides Foissner, 1988 Paramphisiella Foissner, 1988 Paragastrostyla Hemberger, 1985 Gastrostyla Engelmann, 1862 Hemiamphisiella Foissner, 1988 Pseudouroleptus Hemberger, 1985
Table 10 Classification of amphisiellid ciliates according to Petz & Foissner (1996) Amphisiellidae Amphisiella Gourret & Roeser, 1888 Amphisiellides Foissner, 1988 Gastrostyla Engelmann, 1862 Hemiamphisiella Foissner, 1988 Paragastrostyla Hemberger, 1985 Paramphisiella Foissner, 1988 Pseudouroleptus Hemberger, 1985
Table 11 Classification of amphisiellid ciliates according to Shi et al. (1999) and Shi (1999) Family Amphisiellidae Hemberger, 1982 (Shi et al. 1999) Family Amphisielidae Jankowski, 1979 (Shi 1999) Circinella Foissner, 1994 Uroleptoides Wenzel, 1953 Paragastrostyla Hemberger, 1985 Orthoamphisiella Eigner & Foissner, 1991 Amphisiella Gourret & Roeser, 1888 Amphisiellides Foissner, 1988 Lamtostyla Buitkamp, 1977 Mucotrichidium Foissner, 1990 a a
The correct authorship is Foissner, Oleksiv & Müller, 1990.
Table 12 Classification of amphisiellid ciliates according to Lynn & Small (2002) Family Amphisiellidae Jankowski, 1979 Psammomitra Borror, 1972 Circinella Foissner, 1994 Pseudouroleptus Hemberger, 1981 Gastrostyla Engelmann, 1862
PARASITISM, ECOLOGY
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Table 12 Continued Hemiamphisiella Foissner, 1988 Paramphisiella Foissner, 1988 Amphisiellides Foissner, 1988 Paragastrostyla Hemberger, 1981 Orthoamphisiella Eigner and Foissner, 1991 Periholosticha Hemberger, 1981 Uroleptoides Wenzel, 1953 Amphisiella Gourret and Roeser, 1888
Table 13 Classification of trachelostylid ciliates according to Small & Lynn (1985) Family Trachelostylidae n. fam. Psammomitra Trachelostyla Urosoma Urosomoides
Table 14 Classification of trachelostylid ciliates according to Lynn & Small (2002) Family Trachelostylidae Small & Lynn, 1985 Terricirra Berger and Foissner, 1989 Hemisincirra Foissner, 1984 Trachelostyla Kahl, 1932 Lamtostyla Buitkamp, 1977 Gonostomum Sterki, 1878
4
Parasitism
There is no report verifying that amphisiellids or other species reviewed in the present book are attacked by, for example, suctorians, the most common parasites in oxytrichids and urostyloids (for review, see Berger 1999, 2006). However, if you see such an infection you can try to identify the suctorian with the reviews by Matthes (1988), Dovgal (2002), and Lynn & Small (2002).
5
Ecology, Occurrence, and Geographic Distribution
Little is known about these topics. In contrast to oxytrichids and urostyloids, which can be found in all major biotopes (sea, freshwater, soil), most amphisiellids live in terrestrial habitats, a distinctly smaller number (the core amphisiellids) are marine, the trachelostylids as are, and only a minute number can be found in brooks, rivers, lakes, or ponds. The other taxa in the present book also occur almost exclusively in soil. The very low number of freshwater species explains why the amphisiellids do not occur in the old literature, for example, in Ehrenberg (1838), Claparède &
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GENERAL SECTION
Lachmann (1858), or Stein (1859) because these authors studied mainly small, stagnant water bodies. For the same reason, no species is included in the list of biological indicators for water quality assessment (e.g., Berger & Foissner 2003); in addition, amphisiellids do not occur in ordinary sewage treatment plants (Curds 1975a, Ganner et al. 2002). The name-bearing group of the amphisiellids, Amphisiella, and the trachelostylids are marine and several species were included in the reviews by Kahl (1932, 1933). The marine species are, like most other hypotrichs, bottom dwellers creeping on, for example, debris, stones, or macrophytes. Species found in the marine interstitial are summarised by Carey (1992) and Patterson et al. (1989). No symbiotic or parasitic species is known, and likely no species is obligatorily anaerobic (Fenchel & Finlay 1995). The discovery of the terrestrial amphisiellids started in the fifties of the past century, inter alia, by Wenzel (1953), Hemberger (1982, 1985), and W. Foissner who provided many detailed studies including ontogenetic ones (see reference section). The species discussed here have basically the same food spectrum as other hypotrichs (for review, see Berger 1999, 2006), that is, they feed on bacteria, cyanobacteria, algae (including diatoms), auto- and heterotrophic flagellates, and other ciliates. However, small metazoans, for example, rotifers, are usually not ingested because most species are too small for such prey. The menu of the individual species is of course often much shorter than the list above, some are likely specialists ingesting, for example, only bacteria. As in other hypotrichs and ciliates in general, relatively little is known about the geographic distribution of amphisiellids and the other species reviewed. The amphisiellids as such are distributed world-wide. Of course we are unable to say in most cases whether a certain group or species is an endemic taxon or a cosmopolitan, because too few reliable records are published. An example for a very confined distribution is possibly Apourosomoida natronophila, a species so far only recorded from a soda lake in Kenya (Fig. 112a, b). Apourosomoida itself is perhaps restricted to saline habitats in Africa, because the type species was recorded only in highly saline soils in Namibia (Foissner et al. 2002); however, a very similar population was found in an Australian salt lake (Ruinen 1938), suggesting that it is restricted to the southern hemisphere. In the descriptions of the individual species all records published so far are mentioned. There is no doubt that in some cases the determinations are incorrect. Thus, records not substantiated by serious morphological data should be used with caution for biogeographical interpretations. As in other groups of hypotrichs, or ciliates and protists in general, many more species than reviewed in the present book exist because only few areas have been studied in detail, for example, the Namib desert (Foissner et al. 2002). During this survey a rather high number of amphisiellids and amphisiellid-like hypotrichs has been discovered. In addition, the number of described trachelostylids and marine amphisiellids also increases constantly thanks to the research in China (e.g., Hu & Song 2002, Gong et a. 2006, Li et al. 2007, Xu & Lei 2007).
COLLECTING, OBSERVING, STAINING
6
53
Collecting, Culturing, Observing, and Staining of Hypotrichous Ciliates
A detailed description of these topics for all ciliates is given by Foissner (1991, 1993) and Foissner et al. (1991, 1999, 2002).
6.1
Collecting and Culturing
Amphisiellids occur mainly in marine and terrestrial (litters, humic and mineral soil horizons) habitats. Only very few species occur in freshwater. The most effective means for collecting and culturing amphisiellids and other ciliates from soils and mosses is the non-flooded petri dish method as described by Foissner (1987; see also Foissner 1993 and Foissner et al. 2002). Here, 10–200 g of fresh or air-dried soil or litter sample are placed in a petri dish (10–20 cm across) and saturated, but not flooded, with distilled water. A ciliate, flagellate, and naked amoeba fauna, often very rich, develops within a few days. Inspection of the cultures on days 2, 4, 6, 10, 14, and 20 usually suffices. Subsequent inspections reveal only few species due to the effects of ciliatostasis (Lüftenegger et al. 1987). Paraholosticha and Keronopsis species usually occur after few hours, the very common Gonostomum affine can be found also in old cultures. Several conditions influence the outcome of the method: (i) air-dried soil often yields more individuals and species than fresh soil, perhaps due to reduced microbiostasis; (ii) the sample should contain ample litter and plant debris and must be spread over the bottom of the petri dish in an at least 1 cm thick layer; (iii) the soil may not be flooded. Water should be added to the sample until 5–20 ml drains off when the petri dish is tilted and the soil is gently pressed with a finger. This run-off contains the protozoa and can be used for further preparations such as silver staining. The methods for culturing hypotrichous ciliates are treated only briefly here as detailed culturing methods – if available – are provided in the species descriptions. Furthermore, the general procedures as described, for instance by Dragesco & Dragesco-Kernéis (1986), Finlay et al. (1988), Foissner et al. (1991, 2002), Galtsoff et al. (1959), Lee et al. (1985), Mayer (1981), and Provasoli et al. (1958) apply also to the hypotrichs. In general, bacteriovorous hypotrichs thrive on various media (e.g., diluted lettuce and/or hay extracts, table waters [e.g., Volvic], tap water) enriched with a little dried yolk, rice grains, or crushed wheat grains to promote bacterial growth. Some predatory species grow well with small ciliates (e.g., species of the Tetrahymena pyriformis complex, Glaucoma scintillans) as food. For marine species, artificial sea water (e.g., the supersoluble seasalt Biosal by Aqualine Buschke, Berg, Germany) can be used.
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6.2
GENERAL SECTION
Observing Living Hypotrichs
Many physical and chemical methods have been described for retarding the movement of ciliates in order to observe structural details (for literature see Foissner 1991). Chemical immobilisation – for example, by nickel sulfate – or physical slowing down by increasing the viscosity of the medium (e.g., methyl cellulose) are rarely helpful. These procedures often change the shape of the cell or cause premortal alterations of various cell structures. The following simple method is therefore preferable: place about 0.5 ml of the raw sample on a slide and pick out (collect) the desired specimens with a micro-pipette under a compound microscope with low magnification (for example, objective 4:1, ocular 10 ×). If specimens are large enough, they can be picked out from a petri dish under a dissecting microscope. Working with micro-pipettes, the diameter of which must be adjusted to the size of the specimens, requires some training. Transfer the collected specimens, which are now in a very small drop of fluid, onto a slide. Apply small dabs of Vaseline (Petroleum jelly) to each of the four corners of a small cover glass (Fig. 10a; the four dabs can be also applied to the slide); it is useful to apply the jelly by an ordinary syringe with a thick needle. Place the cover glass on the droplet containing the ciliates. Press on the vaselined corners with a mounted needle until ciliates are held firmly between slide and cover glass (Fig. 10b–d). As the pressure is increased the ciliates gradually become less mobile and more transparent. Hence, first the location of the main cell organelles (e.g., nuclear and oral apparatus, contractile vacuole) and then details (e.g., cortical granules, micronucleus) can easily be observed under low (100–400 ×) and high (1000 ×; oil immersion objective) magnification. The colour of the cortical granules and/or the cytoplasm must be studied with well-adjusted bright field. The shape of the cells is of course altered by this procedure. Therefore, specimens taken directly from the raw culture with a large-bore (opening about 1 mm) Pasteur pipette must first be investigated under low magnification (100–400 ×), that is, without cover glass. Some species are too fragile to withstand handling with micro-pipette and cover glass trapping without deterioration. Investigation with low magnification also requires some experience, but it guarantees that the outlines of undamaged cells are recorded. Video-microscopy is very useful at this point of investigation, especially for the registration of the swimming behaviour. A compound microscope equipped with Normarski differential interference contrast optics is best for discerning the arrangement of the cirri and dorsal cilia in living hypotrichs. If not available, use bright-field. The nuclear apparatus is wellrecognisable with differential interference contrast or phase-contrast when specimens are strongly squeezed. Species that were not observed in life often cannot be identified after silver impregnation with certainty because important characters (e.g., size, shape, colour of cortical granules, colour of cytoplasm) are not known. As already mentioned above, the correct colour can only be seen with a well-adjusted bright field illumination.
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6.3
55
Staining Procedures
There are many methods for staining ciliates, but only protargol silver impregnation yields (usually) good results in amphisiellid hypotrichs. Thus, familiarity with this method is an absolute prerequisite for the description of amphisiellids. It is thus described in detail. Simple, namely molecular, formulae are given for the chemicals used, since usually only these are found in the catalogues of the suppliers (e.g., Merck). Other silver impregnation methods (dry silver nitrate method, wet silver nitrate method, silver carbonate method), detailed literature, and some general instructions are to be found in the reviews by Foissner (1991, 1993) and Foissner et al. (1991, 1999). Apart from silver impregnation, some other staining techniques are useful for taxonomic work with ciliates, especially the Feulgen nuclear reaction and supravital staining with methyl green-pyronin in order to reveal the nuclear apparatus and, respectively, the extrusomes.
6.3.1 Feulgen Nuclear Reaction Descriptions of this method are to be found, for example, in Dragesco & DragescoKernéis (1986) and Lee et al. (1985). The Feulgen reaction reveals the nuclear apparatus very distinctively, but, because these organelles usually stain well with protargol, it is seldom used for hypotrichs.
6.3.2 Supravital Staining with Methyl Green-Pyronin This simple method was described by Foissner (1979). It is an excellent technique for revealing the mucocysts of most ciliates. Mucocysts are stained deeply and very distinctively blue or red, and can be observed in various stages of explosion because the cells are not killed instantly. The nuclear apparatus is also stained. Procedure (after Foissner 1991) 1. Pick out desired ciliates with a micro-pipette and place the small drop of fluid in the centre of a slide. 2. Add an equally sized drop of methyl green-pyronin and mix the two drops gently by swivelling the slide. Remarks: If ciliates were already mounted under the coverslip, add a drop of the dye at one edge of the coverslip and pass it through the preparation with a piece of filter paper placed at the other end of the coverslip. 3. Place a coverslip with vaselined corners on the preparation.
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Fig. 10a–f Live observation and staining of urostyloid ciliates (from Foissner 1991). a–d: Preparation of slides for observing living ciliates. e: Staining jar for eight and 16 (back to back) slides, respectively. f: Watch-glass for protargol procedure according to Wilbert.
Remarks: Observe immediately. Cells die in the stain within 2 min. Mucocysts stain very quickly and many can be observed at various stages of explosion. To reveal the nuclear apparatus, cells should be fairly strongly squashed (= flattened). The preparation is temporary. After 5–10 min the cytoplasm often becomes heavily stained and obscures other details. Reagents 1 g methyl green-pyronin (Chroma-Gesellschaft, Schmid GmbH and Co., Köngen/N., Germany) add 100 ml distilled water This solution is stable and can be used for years.
6.3.3 Protargol Methods Protargol methods are indispensable for descriptive research on amphisiellids and hypotrichs in general. The first procedures were provided by Kirby (1945), Moskowitz (1950), Dragesco (1962), and Tuffrau (1964, 1967), and many more modifications were subsequently proposed (see Foissner 1991 for references). Here, two variations which produce good results are described. These procedures work well with most ciliate species, but require at least 20 specimens. A single specimen can-
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not usually be handled successfully. Depending on the procedure used, protargol can reveal many cortical and internal structures, such as basal bodies, fibrillary systems, nuclear apparatus. The silverlines (which have no systematic value in the amphisiellids), however, never impregnate. The shape of the cells is usually well preserved in permanent slides, which is an advantage for the investigation, but makes photographic documentation more difficult. However, pictures as clear as those taken from wet silver carbonate impregnations can be obtained with the Wilbert modification (procedure B) if the cells are photographed prior to embedding in the albumenglycerol. Procedure A (after Foissner 1991) The quality of the slides is usually adequate but frequently not as good as with the Wilbert modification. The latter demands more material and experience; inexperienced workers may easily lose all the material. As in all protargol methods, the procedure is rather time-consuming and complicated because subject to many factors. A centrifuge may be used for step 2; staining jars (Fig. 10e) are necessary for steps 6–16. 1. Fix organisms in Bouin’s or Stieve’s fluid for 10–30 min. Remarks: The fixation time has little influence on the quality of the preparation within the limits given. Ratio fixative to sample fluid should be at least 2:1. Pour ciliates into fixative using a wide-necked flask in order to bring organisms in contact with the fixative as quickly as possible. Both fixatives work well but may provide different results with certain organisms. Stieve’s fluid may be supplemented with some drops of 2% osmium tetroxide for better fixation of very fragile hypotrichs. This increases the stability of the cells but usually reduces their impregnability. Alternatively, 70% ethanol can be used for fixation (W. Foissner, pers. comm.). 2. Concentrate by centrifugation and wash organisms three to four times in distilled water. Remarks: There are now two choices: either to continue with step 3, or to transfer the material through 30–50–70% alcohol into 70% alcohol (ethanol), where it remains stable for several years. Transfer preserved material back through the graded alcohol series into distilled water prior to continuing with the next step. Impregnability of preserved material may be slightly different. 3. Clean 8 ordinary slides (or less if material is very scarce) per sample. The slides must be grease-free (clean with alcohol and flame). Remarks: Insufficiently cleaned slides may cause the albumen to detach. Mark slides on back if several samples are prepared together. Alternatively you can use SuperFrost slides which are ready to use. In addition, these slides have a field enabling
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simple labelling with a pencil. Use staining jars with eight sections so that you can work with 16 slides simultaneously by putting them back to back (Fig. 10e). 4. Put 1 drop each of albumen-glycerol and concentrated organisms in the centre of a slide. Mix drops with a mounted needle and spread over the middle third. Remarks: Use about equally sized drops of albumen-glycerol and suspended (in distilled water) organisms to facilitate spreading. The size of the drops should be adjusted so that the middle third of the slide is covered after spreading. Now remove sand, grains, etc. The thickness of the albumen layer should be equal to that of the organisms. Some thicker and thinner slides should, however, also be prepared because the thickness of the albumen layer greatly influences the quality of the preparation. Cells may dry out and/or shrink if the albumen layer is too thin; if it is too thick, it may detach, or the cells may become impossible to study with the oil immersion objective. 5. Allow slides to dry for at least 2 h at room temperature. Remarks: We usually dry slides overnight, that is, for about 12 h. However, slides may be allowed to dry for up 24 h, but no longer if quality is to be maintained. Oven-dried (2 h at 60°C) slides are usually also of poorer quality. 6. Place slides in a staining jar (Fig. 10e) filled with 95% alcohol (ethanol) for 20 to 30 min. Place a staining jar with protargol solution into an oven (60°C). Remarks: Slides should not be transferred through an alcohol series into concentrated alcohol as this causes the albumen layer to detach! Decrease hardening time to 20 min if albumen is already rather old and/or not very sticky. 7. Rehydrate slides through 70% alcohol and two distilled water steps for 5 min each. 8. Place slides in 0.2% potassium permanganate solution. Remove first slide (or pair of slides) after 60 s and the rest at 15 s intervals. Collect slides in a staining jar filled with distilled water. Remarks: Bleaching is by permanganate and oxalic acid (step 9). The procedure described above is necessary because each species has its optimum bleaching time. The sequence in which slides are treated should be recorded because the immersion time in oxalic acid must be proportional to that in the permanganate solution. The albumen layer containing the organisms should swell slightly in the permanganate solution and the surface should become uneven. If it remains smooth, the albumen is too sticky and this could decrease the quality of the impregnation. If the albumen swells strongly, it is possibly too weak (old) and liable to detach. Use fresh KMnO4 solution for each series.
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9. Quickly transfer slides to 2.5% oxalic acid. Remove first slide (or pair of slides) after 160 s, the others at 20 s intervals. Collect slides in a staining jar filled with distilled water. Remarks: Same as for step 8! Albumen layer becomes smooth in oxalic acid. 10. Wash slides three times in distilled water for 3 min each. 11. Place slides in warm (60°C) protargol solution and impregnate for 10–15 min at 60°C. Remarks: Protargol solution can be used only once. 12. Remove staining jar with the slides from the oven and allow to cool for 10 min at room temperature. Remarks: In the meantime organise six staining jars for developing the slides: distilled water – distilled water – fixative (sodium thiosulfate) – distilled water – 70% alcohol – 100% alcohol (ethanol). 13. Remove the first slide from the protargol solution and drop some developer on the albumen layer. Move slide gently to spread developer evenly. As soon as the albumen turns yellowish, pour off the developer, dip slide in the first two distilled water steps for about 2 s each and stop development by submerging the slide in the fixative (sodium thiosulfate), where it can be left for 1–5 min. Remarks: Now control impregnation with the compound microscope. The impregnation intensity is sufficient if the infraciliature is just recognisable. The permanent slide will be too dark if the infraciliature is distinct at this stage of the procedure! The intensity of the impregnation can be controlled by the concentration of the developer and the time of development. 5–10 s usually suffice for the diluted developer! Development time increases with bleaching time. Therefore commence development with those slides which were in the bleaching solutions for 60 and 120 s, respectively. The thinner the albumen layer, the quicker the development. 14. Collect slides in the fixative (sodium thiosulfate) and transfer to distilled water for up to 5 min. Remarks: Do not wash too long; the albumen layer is very fragile and detaches rather easily! 15. Transfer slides to 70% – 100% – 100% alcohol for 5 min each. 16. Clear by two 10 min transfers through xylene. 17. Mount in synthetic neutral mounting medium. Remarks: Do not dry slides between steps 16 and 17! Mounting medium should be rather viscous to avoid air-bubbles being formed when solvent evaporates during drying. If air-bubbles develop in the mounted and hardened slide, re-immerse in
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xylene for some days until the coverslip drops off. Remount using a more viscous medium and remove possible sand grains protruding from the gelatine. Usually, some air-bubbles are found immediately after mounting; these can be pushed to the edge of the coverslip with a finger or mounted needle. The preparation is stable. Reagents a) Bouin’s fluid (prepare immediately before use; components can be stored) 15 parts saturated, aqueous picric acid (C6H3N3O7; preparation: add an excess of picric crystals to, for example, 1 litre of distilled water; shake solution several times within a week; some undissolved crystals should remain; filter before use) 5 parts formalin (HCHO; commercial concentration, about 37%) 1 part glacial acetic acid (= concentrated acetic acid; C2H4O2) b) Stieve’s fluid (slightly modified; prepare immediately before use; components can be stored) 38 ml saturated, aqueous mercuric chloride (dissolve 60 g HgCl2 in 1 litre of boiling distilled water) 10 ml formalin (HCHO; commercial concentration, about 37%) 3 ml glacial acetic acid (= concentrated acetic acid; C2H4O2) c) Albumen-glycerol (2–4 month stability) 15 ml egg albumen 15 ml concentrated (98–100%) glycerol (C3H8O3) Pre-treatment of the egg albumen and preparation of the albumen-glycerol: separate the white carefully from the yolk and embryo of three eggs (free range eggs are preferable to those from battery chickens, whose egg white is less stable and sticky). Shake the white by hand (do not use a mixer!) for some minutes in a narrowmouthed 250 ml Erlenmeyer flask until a stiff white foam is formed. Allow the flask to stand for about 1 min. Then pour the viscous rest of the egg white in a second Erlenmeyer flask and shake again until a stiff foam is formed. Repeat until most of the egg white is either stiff or becomes watery; usually 4–6 Erlenmeyer flasks of foam are obtained. Leave all flasks undisturbed for about 10 min and discard the watery albumen from the last flask. During this time a glycerol-like fluid percolates from the foam. This fluid is collected and used. Add an equal volume of concentrated glycerol and a small thymol crystal (C10H14O) for preservation to the mixture. Mix by shaking gently and pour mixture into a small flask. Leave undisturbed for two weeks. A whitish slime settles at the bottom of the flask. Decant the clear portion, discard slime and thymol crystal. A “good” albumen-glycerol drags a short thread when touched with a needle. The albumen is too thin (not sticky enough) or too old if this thread is not formed. Fresh albumen which is too thin may be concentrated by leaving it open for some weeks so that water can evaporate. If the albumen is too sticky, which may cause only one side of the organisms to impregnate well, it is di-
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luted with distilled water or old, less sticky albumen to the appropriate consistency. The preparation of the albumen-glycerol must be undertaken with great care because much depends on its quality. Unfortunately, all commercial products which have been tried detach during impregnation. A somewhat simpler method to produce the albumen-glycerol is described by Adam & Czihak (1964, p. 274): the white of one or two fresh chicken egg(s) and the same amount of concentrated glycerol are well stirred to a homogenous, thick fluid (a magnetic stirrer can be used). Then filter through cotton wool. Add a small thymol crystal to the filtrate. The albumenglycerol can be used right away. d) 0.2% potassium permanganate solution (stable for about 1 d) 0.2 g potassium permanganate (KMnO4) are dissolved in 100 ml distilled water e) 2.5% oxalic acid solution (stable for about 1 d) 2.5 g oxalic acid (C2H2O4·2H2O) are dissolved in 100 ml distilled water f) 0.4–0.8% protargol solution (stable for about 1 d) 100 ml distilled water add 0.4–0.8 g protargol Remarks: Use light-brown “protargol for microscopy” (for example, Merck’s “Albumosesilber für die Mikroskopie” or “Proteinate d’Argent”, Roques, Paris, France). Some dark-brown, cheaper products do not work! Sprinkle powder on the surface of the water in the staining jar and allow to dissolve without stirring. g) Developer (mix in sequence indicated; sodium sulphite must be dissolved before hydroquinone is added) 95 ml distilled water 5 g sodium sulphite (Na2SO3) 1 g hydroquinone (C6H6O2) Remarks: This recipe yields the stock solution which is stable for some weeks and should be used undiluted for certain ciliates (step 13). Usually, however, it must be diluted with tap water in a ratio of 1:20 to 1:50 to avoid too rapid development and one-sided impregnation of the organisms. Freshly prepared developer is usually inadequate (the albumen turns greenish instead of brownish). The developer should thus be prepared from equal parts of fresh and old (brown) stock solutions. Take great care with the developer as its quality contributes highly to that of the slides. If the developer has lost its activity (which is not always indicated by a brown colour!) the silver is not or only insufficiently reduced and the organisms stain too faintly. A fresh developer should therefore be prepared for each “impregnation week”, and some old developer kept. Fresh developer can be artificially aged by adding some sodium carbonate (Na2CO3). However, better results are obtained with air-aged solutions, that is, by a developer which has been kept uncovered for some days in a wide-mouthed bottle. It first turns yellowish, then light brown (most effective), and later dark brown and viscous (at this stage the developer has lost most of its activity,
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but is still suitable for artificial ageing of fresh developer = 1:1 mixture mentioned above). During the last years, we obtained very good slides with the low-speed developer used by Fryd-Versavel (pers. comm. to W. Foissner). It is composed of 7 g boric acid, 1.5 g hydroquinone, 10 g sodium sulphite, and 75 ml acetone, all solved, one by one, in 420 ml distilled water. This developer is stable for some weeks and should be used only once. Pour developer into a staining jar and immerse slides, one by one, controlling impregnation intensity after 30–60 s. Usually, developing is finished within 1–5 min (if not, double protargol concentration because slides should not be too long in the developer, as the albumen may detach). The further procedure is as described above (steps 14–17). In many cases commercial paper developers (for example, Ilford Multigrade) yield very good results. h) Fixative for impregnation (stable for several years) 25 g sodium thiosulfate (Na2S2O3·5H2O) are dissolved in 1000 ml distilled water Procedure B (after Wilbert 1975 and Foissner 1991) This modification produces excellent results but demands much experience. Manipulate large cells with micropipettes in a watch-glass, whereas the centrifuge is used for steps 1–4, 7, 8 if cells are smaller than about 150 µm. The watch-glass method is used when there are only few specimens of larger cells; thus an attempt is worthwhile even if only 20 cells are available. The organisms are very soft after development and fixation, and are thus easily compressed between slide and coverslip, which greatly facilitates photographic documentation. 1. Fix organisms as described in protargol procedure A. 2. Wash and, if so desired, preserve organisms as described in protargol procedure A. Remarks: Wash cells either in the centrifuge (small species) or in a watch-glass (Fig. 10f). To change fluids allow cells to settle on bottom of watch-glass and remove supernatant with a micro-pipette under the dissecting microscope; concentrate cells in the centre of the watch-glass by gentle swirling. 3. Leave organisms in a small amount of distilled water and add, drop by drop, diluted sodium hypochlorite (NaClO) and bleach for about 1–3 min under the dissecting microscope (for small specimens, various concentrations of NaClO can be applied in centrifuge tubes, keeping the reaction time constant, for example, 1 min). Remarks: This is the critical step in this modification. If bleaching is too strong or too weak all is lost: cells either dissolve or do not impregnate well. Systematic investigations showed that not the bleaching time but the amount of active chloride in the sodium hypochlorite and the pre-treatment of the cells (fixation method, fresh or
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preserved material) are decisive for the quality of the preparation. Different species need different concentrations. Unfortunately, the concentration of active chloride in the commercial products varies (10–13%) and is dependent on the age of the fluid. It is thus impossible to provide more than only a few guidelines: 100 ml distilled water + 0.2–0.4 ml NaClO (if product is fresh and cells were not stored in alcohol) or 100 ml distilled water + 0.5–1.6 ml NaClO (if product is older and cells were stored in alcohol). The transparency of the cells under the dissecting microscope may serve as a further indicator: fixed, unbleached cells appear dark and opaque, whereas accurately bleached cells are almost colourless and rather transparent (depends, however, also on size and thickness of the cell). Thus, increase the concentration of sodium hypochlorite stepwise if cells appear too dark with the recommended concentrations. We routinely start with 3 different hypochlorite concentrations if enough material is available. 4. Wash organisms at least 3 times with distilled water and finally once in the protargol solution. Remarks: Wash thoroughly, especially when fluids are changed with micro-pipettes, because even the slightest traces of the sodium hypochlorite disturb impregnation. 5. Transfer to 1% protargol solution and impregnate for 10–20 min at 60°C. Remarks: This and the next step should be carried out in a watch-glass even for material which is otherwise manipulated with the centrifuge. The impregnation time depends on the kind of material and the degree of bleaching. Check the progress of impregnation every 3–4 min under the compound microscope by picking out a few cells with the micro-pipette under the dissecting microscope; add these to one drop of developer. Dilute developer and/or interrupt development of adding a little fixative (sodium thiosulfate) if impregnation is strong enough. 6. Remove most of the protargol solution with a micro-pipette and add some drops of developer to the remainder containing the organisms. Remarks: Fresh, undiluted developer is usually used (but see step 5). Control development in compound or dissecting microscope. As soon as the infraciliature becomes faintly visible, development must be stopped by adding a few drops of sodium thiosulfate. Judging the right moment is a question of experience; the permanent slide will be too dark if the infraciliature is very distinct at this stage of the procedure! Generally: if bleaching was correct, specimens cannot be over-impregnated. 7. Stabilise the impregnation by two, approximately 5-minute transfers through sodium thiosulfate. Remarks: The developer need not be removed before fixation. For small species this and the next step can be carried out in a centrifuge. Larger species must be manipulated with micro-pipettes because cells become very fragile and would be damaged in a centrifuge. Cells are very soft at this stage and can thus be easily compressed and photographed. Transfer some of the more darkly impregnated specimens with a
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very small amount of the fixative onto a clean slide using a micro-pipette and cover with a coverslip. Organisms are usually flattened by the weight of the coverslip; excess fluid my be removed from the edge of the coverslip with a piece of filter paper. Frequently, even better micrographs are obtained if specimens are flattened before fixed with sodium thiosulphate; that is, together with some developer. 8. Wash very thoroughly in distilled water (three times with the centrifuge; 7–10 times in watch-glass with micro-pipettes). Finally remove as much of the water as possible. Remarks: Even the slightest traces of the fixative destroy the impregnation within a few days or weeks. 9. Smear a moderately thick layer of albumen-glycerol on a clean slide with a finger. Drop impregnated, washed cells on the albumised slide with a large-bore pipette (opening about 1 mm) and dry preparation for at least 2 h. Remarks: The cells are too fragile to be spread with a needle. With much care it is possible to orientate cells using a mounted eyelash. Commercial albumen-glycerol can be used. 10. Harden albumen by two 10-minute transfers through concentrated alcohol (ethanol). Remarks: This and the next step are best carried out in staining jars. The albumen layer turns milky and opaque. 11. Clear by two 5-minute transfers through xylene. Remarks: The albumen layer turns transparent. 12. Mount in synthetic neutral mounting medium. Remarks: Do not dry slides between steps 11 and 12! Mounting medium should be rather viscous to avoid air-bubbles being formed when solvent evaporates during drying. If air-bubbles develop in the mounted and hardened slide, re-immerse in xylene for some days until the coverslip drops off. Remount using a more viscous medium and remove possible sand grains protruding from the albumen. Usually, some air-bubbles are found immediately after mounting; these can be pushed to the edge of the coverslip with a finger or mounted needle. The preparation is stable. Reagents If not stated otherwise, the same reagents as in the protargol procedure A are to be used (see above).
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6.4
65
Preparation for Scanning Electron Microscopy
Hypotrichs cannot usually be identified solely by scanning electron microscopy because only a limited number of characters is revealed. However, this method is useful in that it allows a three-dimensional view of the object, as well as for documenting details which are difficult to reveal with other methods. For a detailed instruction of preparation for scanning electron microscopy, see Foissner (1991, 1993), Foissner et al. (1991, 1999), and other textbooks.
7
Species Concept and Nomenclature
7.1 Species Concept The species concept used here is the same as described by Foissner et al. (2002). Briefly, we usually apply the “morphospecies” concept as basically defined by Nixon & Wheeler (1990): “A species is the smallest aggregation of populations (sexual) or lineages (asexual) diagnosable by a unique combination of character states in comparable individuals (semaphoronts).” That is, I consider two populations as belonging to two different species if they differ from each other in at least one “important” morphological feature (e.g., number of macronuclear nodules). Of course, there is no strict consensus about the importance of various features and, unfortunately, for many species several features (e.g., presence/absence of cortical granules or caudal cirri; number and arrangement of dorsal kineties; length of dorsal bristles; exact arrangement of cirri) are not known, making revisions rather difficult. Often it is a matter of taste whether or not two species are synonymised. To overcome these difficulties I have kept the descriptions and the ecological data of synonyms separate, especially when the descriptions did not fit in all important details. The presence or absence of a certain cirral group or part of the dorsal ciliature (e.g., postperistomial cirrus, transverse cirri, caudal cirri) is generally considered as diagnostic character for supraspecific taxa. However, features of the cirral pattern are certainly not the sole source to elucidate the phylogenetic relationships. The consistence of the cell (flexible vs. rigid), the presence/absence of cortical granules, and the relative length (i.e., a quantitative feature!) of the adoral zone have been successfully used to characterise the Stylonychinae (Berger & Foissner 1997, Berger 1999), and in the urostyloids the individually dividing macronuclear nodules are the apomorphy of the Pseudokeronopsinae (Berger 2006). Moreover, molecular markers will significantly increase our knowledge on the phylogeny of hypotrichs. Relatively many features have evolved convergently, perhaps even the rigidity of the cortex (stylonychines, Berger 1999; Rigidothrix, Foissner & Stoeck 2006) or the zigzagging cirral pattern of the urostyloids (Foissner et al. 2004, Berger 2006) which occurs, inter alia, in Uroleptus (e.g., Foissner et al. 1991, Eigner 2001) and several oxytrichids as, for example, Neokeronopsis (Berger 1999, 2006).
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For a discussion of the advantages and disadvantages of various species concepts, see textbooks and papers on evolution (e.g., Ax 1984, 1995, Sudhaus & Rehfeld 1992, Winston 1999, Wägele 2001, Knoop & Müller 2006, Lecointre & Le Guyader 2006, Weisse 2006) and references cited by Foissner et al. (2002, p. 35).
7.2 Nomenclature In the case of nomenclatural problems, the ICZN (1964, 1985, 1999) have been consulted, depending on the date when the paper was published. For explanation of nomenclatural terms (e.g., nomen nudum, holotype), see the glossary of the ICZN (1999) or various textbooks (e.g., Lincoln et al. 1985, CBE 1996, Winston 1999). I tried to explain the meaning and origin of the scientific names using, inter alia, the ICZN (1985, 1999), Werner (1972), Hentschel & Wagner (1996), Cole (2000), and Latin/German dictionaries. Likely less than 50% of the original descriptions contain an etymology section. The gender of ciliate genus-group names can be found, inter alia, in the valuable catalogue by Aescht (2001). I did not consult a Latin/Greek linguist; thus, deficiencies cannot be excluded. Note that in the reviews by Kahl (1932, 1933) only the type genus of the Amphisiellidae and some trachelostylids are included. In addition, Kahl classified Amphisiella as subgenus of Holosticha, a urostyloid genus (Berger 2003, 2006). Most other species reviewed in the present monograph are not revised in Kahl’s papers because they have been discovered in the last five decades during the systematic investigation of soil from throughout the world (for reviews, see Foissner 1987, 1998 and Foissner et al. 2002) and the detailed research on marine habitats, mainly from China (e.g., Hu et al. 2003). For authorship and date of hypotrichs not reviewed in the present volume, see Berger (1999, 2001, 2006) and the permanently updated “Online Catalogue of Ciliate Names. 1. Hypotrichs and Euplotids” at http://www.protozoology.com. A comprehensive bibliography comprising 6062 references about hypotrichs and euplotids is provided by Berger (2006a). As in the first two volumes of the revision of hypotrichs (Berger 1999, 2006), higher taxa are not provided with categories (e.g., family, order)1, simply because categories do not contain information and cannot be defined objectively (for details see, e.g., Ax 1995, Westheide & Rieger 1996, Wägele 2001). For example, the taxon Hypotricha was established as order by Stein (1859). Since then it also attained the categories suborder, subclass, and even class (for review, see Berger 2001). How1
According to Foissner et al. (2007, p. 312), Berger (2006) has written that “higher taxa do not exist in nature”. In addition, I should have suggested a new higher category for the group composed of hypotrichs and oligotrichs. Both statements by Foissner et al. (2007) are incorrect. Apparently they refer to the legend of Fig. 13a in Berger (2006, p. 32) where I wrote “I do not use categories (e.g., family, order), simply because they do not exist in nature” (see also Berger 2006, p. 67). Obviously Foissner et al. (2007) did not distinguish between the terms taxon and category (for explanation of the difference, see, for example, Ax 1999 and Wägele 2001).
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67
ever, to avoid inflation of names I use those which are available. Therefore the “defined” endings (ICZN 1999, Article 29.2; e.g., -idae, -inae) have no meaning in this book.
7.3 Summary of New Taxa and Nomenclatural Acts Within this book the nomenclatural acts listed below have been undertaken. Several distinct (monophyletic?) groups within the amphisiellids (see groups I to IV in table of contents) and within some genera, for example, in Lamtostyla can be distinguished. However, further results, especially molecular data, should be awaited before naming them. New combinations: Anteholosticha heterocirrata (Hemberger, 1985) comb. nov. (p. 640; basionym: Perisincirra heterocirrata); Anteholosticha verrucosa (Foissner & Schade in Foissner, 2000) comb. nov., stat. nov. (p. 647; basionym: Hemisincirra gellerti verrucosa); Apourosomoida natronophila (Dietz, 1965) comb. nov. (p. 530; basionym: Uroleptus natronophilus); Bistichella buitkampi (Foissner, 1982) comb. nov. (p. 535; basionym: Paraurostyla buitkampi); Bistichella humicola (Gellért, 1956) comb. nov. (p. 556; basionym: Uroleptus humicola); Bistichella namibiensis (Foissner, Agatha & Berger, 2002) comb. nov. (p. 538; basionym: Amphisiella namibiensis); Bistichella procera (Berger & Foissner, 1987) comb. nov. (p. 547; basionym: Pseudouroleptus procerus); Bistichella terrestris (Hemberger, 1985) comb. nov. (p. 554; basionym: Pseudouroleptus terrestris); Caudiamphisiella antarctica (Wilbert & Song, 2005) comb. nov. (p. 129; basionym: Amphisiella antarctica); Hemiamphisiella terricola qingdaoensis (Song & Wilbert, 1989) comb. nov., stat. nov. (p. 310; basionym: Uroleptoides qingdaoensis); Hemisincirra buitkampi (Jankowski, 1979) comb. nov. (p. 393; basionym: Perisincirra buitkampi); Kleinstyla bavariensis (Foissner, Agatha & Berger, 2002) Foissner, Agatha & Berger comb. nov. (p. 140; basionym: Gastrostyla (Kleinstyla) bavariensis); Kleinstyla dorsicirrata (Foissner, 1982) Foissner, Agatha & Berger comb. nov. (p. 140; basionym: Gastrostyla dorsicirrata); Lamtostyla elegans (Foissner, Agatha & Berger, 2002) comb. nov. (p. 192; basionym: Amphisiella elegans); Lamtostyla procera (Foissner, Agatha & Berger, 2002) comb. nov. (p. 183; basionym: Amphisiella procera); Lamtostyla quadrinucleata (Berger & Foissner, 1989) comb. nov. (p. 190; basionym: Amphisiella quadrinucleata); Lamtostyla vitiphila (Foissner, 1987) comb. nov. (p. 187; basionym: Uroleptoides vitiphila); Lamtostylides edaphoni (Berger & Foissner, 1987) comb. nov. (p. 324; basionym: Lamtostyla edaphoni); Lamtostylides halophilus (Foissner, Agatha & Berger, 2002) comb. nov. (p. 336; basionym: Lamtostyla halophila); Lamtostylides hyalinus (Berger, Foissner & Adam, 1984) comb. nov. (p. 347; basionym: Tachysoma hyalina); Lamtostylides kirkeniensis (Berger & Foissner, 1988) comb. nov. (p. 333; basionym: Lamtostyla kirkeniensis); Lamtostylides pori (Wilbert & Kahan, 1986) comb. nov. (p. 344; basionym: Perisincirra pori); Maregastrostyla pulchra (Pereyaslawzewa, 1886) comb. nov. (p. 137; basionym: Stilonichia pulchra);
68
GENERAL SECTION
Nudiamphisiella illuvialis (Eigner & Foissner, 1994) comb. nov. (p. 569; basionym: Amphisiellides illuvialis); Spetastyla mystacea (Stein, 1859) Foissner, Agatha & Berger comb. nov. (p. 140; basionym: Oxytricha mystacea); Spetastyla mystacea mystacea (Stein, 1859) Foissner, Agatha & Berger comb. nov. (p. 140; basionym: Oxytricha mystacea); Spetastyla mystacea minima (Hemberger, 1985) Foissner, Agatha & Berger comb. nov. (p. 140; basionym: Gastrostyla minima); Trachelostyla rostrata (Lepsi, 1962) comb. nov. (p. 498; basionym: Trachelostyla rostrata); Uroleptoides binucleatus multicirratus (Foissner, Agatha & Berger, 2002) comb. nov. (p. 267; basionym: Amphisiella binucleata multicirrata); Uroleptoides longiseries (Foissner, Agatha & Berger, 2002) comb. nov. (p. 249; basionym: Amphisiella longiseries); Uroleptoides magnigranulosus (Foissner, 1988) comb. nov. (p. 273; Amphisiella magnigranulosa); Uroleptoides multinucleatus (Foissner, Agatha & Berger, 2002) comb. nov. (p. 255; basionym: Amphisiella multinucleata); Uroleptoides polycirratus (Berger & Foissner, 1989) comb. nov. (p. 285; basionym: Amphisiella polycirrata); Uroleptoides terricola (Gellért, 1956) comb. nov. (p. 279; basionym: Amphisiella terricola). New genera: Bistichella gen. nov. (p. 532; type species: Paraurostyla buitkampi Foissner, 1982); Caudiamphisiella gen. nov. (p. 129; type species: Amphisiella antarctica Wilbert & Song, 2005); Lamtostylides gen. nov. (p. 322; type species: Lamtostyla edaphoni Berger & Foissner, 1987); Maregastrostyla gen. nov. (p. 136; type species: Stilonichia pulchra Pereyaslawzewa, 1886). New higher taxon: Perilemmaphora tax. nov. (p. 37). New ranks: Anteholosticha verrucosa (Foissner & Schade in Foissner, 2000) comb. nov., stat. nov. (now species rank; p. 647); Hemiamphisiella terricola terricola Foissner, 1988 stat. nov. (now subspecies rank; p. 296); Hemiamphisiella terricola qingdaoensis (Song & Wilbert, 1989) comb. nov., stat. nov. (now subspecies rank; p. 310); Kleinstyla Foissner, Agatha & Berger, 2002 (now genus rank; p. 140); Spetastyla Foissner, Agatha & Berger, 2002 (now genus rank; p. 140). Corrected name: Hemiurosoma polynucleatum (p. 634). New synonyms: Metastrongylidium Xu & Lei, 2007 is synonymous with Spiroamphisiella Li, Song & Hu, 2007 (p. 148); Metastrongylidium distichum Xu & Lei, 2007 in synonymous with Spiroamphisiella hembergeri Li, Song & Hu, 2007 (p. 150); Maregastrostyla Berger, 2008 (p. 136) is synonymous with Protogastrostyla Gong, Kim, Kim, Min, Roberts, Warren & Choi, 2007 (p. 690).
SPECIES CONCEPT, NOMENCLATURE
69
7.4 Deposition of Slides If mentioned in the individual papers, the site where the slide(s) (holotype; paratype; neotype; voucher) has (have) been deposited, is given in the corresponding entry of the list of synonyms. For a detailed list of type specimens deposited in the collection “diverse invertebrates (except insects)” of the Biology Centre Linz (Upper Austria), see Aescht (2003; also available at www.biologiezentrum.at). Slides used for original observations are also deposited in this collection.
B Systematic Section This treatise is about the Amphisiellidae, the Trachelostylidae, and several genera of uncertain position. However, the cirral pattern of the latter is somewhat reminiscent of amphisiellids and therefore they are included in this monograph. In addition, supplements to the urostyloids (Berger 2006) and oxytrichids (Berger 1999) are provided. With the key below and the subsequent keys you can identify all species reviewed in this book. Unfortunately, there are no “simple” features showing that a certain species must be included in this revision. However, if one of the following combinations of characteristics applies to your specimen/population you have a good chance of finding it in this volume. y
Body flexible; three enlarged frontal cirri; one more or less distinct frontoventral row (e.g., Fig. 2a), or two widely separated frontoventral rows (e.g., Fig. 113e), or 18-cirri hypotrichs which do not have cirri in the postoral area (e.g., Lamtostyla longa- and Lamtostyla granulifera-group, Trachelostylidae; Fig. 3a, 4a)
y
Marine, cephalised 18-cirri hypotrichs (Trachelostylidae; Fig. 4a)
y
Slender or vermiform soil species without distinct frontoventral row (e.g., Hemisincirra, Erimophrya, Hemiurosoma, Vermioxytricha, Apourosomoida)
Representatives of all genera in this book are summarised on the following plates (Fig. 11a–d, 12a–g, 13a–f, 14a–d, 15a–h). A key to all hypotrich genera and higher taxa will be provided in the last volume of the series.
Key to the Taxa Treated in Present Book Before you start with the key you should familiarise yourself with the main morphological terms (see general section). Note that in the key the neutral term frontoventral row is used, even for amphisiellids where this row is usually termed amphisiellid median cirral row. Some key features (e.g., caudal cirri present/absent, transverse cirri present/absent [must not be confused with caudal cirri or marginal cirri!]) are rather difficult to recognise, especially in species with a slender body or posteriorly narrowed body. Thus, protargol preparations, or at least very detailed live observations using interference contrast, are needed. Note that some genera are included twice because they cannot be assigned unambiguously to a single character state! 1 Marine or inland saltwater . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 - Terrestrial or limnetic. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
71
72
SYSTEMATIC SECTION
Fig. 11a–d Ventral infraciliature of representative of marine amphisiellids (sources of illustrations see individual descriptions). a: Amphisiella annulata; caudal cirri lacking. b: Caudiamphisiella antarctica; caudal cirri present. c: Maregastrostyla pulchra; caudal cirri present. d: Spiroamphisiella hembergeri; caudal cirri present.
2 3 4 5
(29) Distinct frontoventral row or postoral row present (e.g., Fig. 11a, c, 14c). . 3 Postoral area without frontoventral cirri (Fig. 14a, b) . . Trachelostylidae (p. 471) Body slenderly spindle-shaped (Fig. 13f). . . . . . . . . . . . . . . Cossothigma (p. 382) Body not as above. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Body distinctly twisted about main axis (Fig. 11d). . . . Spiroamphisiella (p. 148) Body not distinctly twisted. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Transverse cirri distinctly displaced anteriad so that they do not project beyond rear body end (Fig. 11c). . . . . . . . . . . . . . . . . . . . . . . . . Maregastrostyla (p. 136)
Fig. 12a–e Ventral infraciliature of terrestrial amphisiellids, which have six frontal-ventral-transverse cirri anlagen (sources of illustrations see individual descriptions). a: Lamtostyla lamottei; caudal cirri lacking. b: Lamtostyla granulifera; caudal cirri lacking; three dorsal kineties. c: Lamtostyla longa; caudal cirri lacking; five dorsal kineties. d: Uroleptoides binucleatus; caudal cirri lacking. e: Hemiamphisiella terricola; caudal cirri present. Fig. 12f, g Ventral infraciliature of terrestrial amphisiellids which lack frontal-ventral-transverse cirri anlage IV (sources of illustrations see individual descriptions). f: Lamtostylides edaphoni; caudal cirri lacking. g: Paramphisiella acuta; caudal cirri present.
d
KEY TO TAXA
73
74
SYSTEMATIC SECTION
Fig. 13a–f Infraciliature of ventral side of genera classified as incertae sedis in the amphisiellids (sources of illustrations see individual descriptions). a: Afroamphisiella multinucleata; caudal cirri lacking. b: Hemisincirra buitkampi; caudal cirri lacking. c: Terricirra viridis; caudal cirri lacking. d: Mucotrichidium hospes; caudal cirri present. e: Tetrastyla oblonga; presence/absence of caudal cirri not known. f: Cossothigma dubium; presence/absence of caudal cirri not known.
KEY TO TAXA
75
Fig. 14a, b Infraciliature of ventral side of trachelostylids (sources of illustrations see individual descriptions). a: Trachelostyla pediculiformis; body not twisted; caudal cirri present. b: Spirotrachelostyla spiralis; body twisted; caudal cirri present. Fig. 14c, d: Two genera of unknown position in the Hypotricha (sources of illustrations see individual descriptions). c: Apourosomoida halophila; caudal cirri present. d: Bistichella buitkampi; caudal cirri lacking.
-
Transverse cirri not distinctly displaced anteriad, that is, protrude beyond rear body end (e.g., Fig. 11a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5a 5a Postoral cirral row present (Fig. 14c). . . . . . . . . . . . . . . . Apourosomoida (p. 514) - Frontoventral row present (Fig. 11a, b). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6 Caudal cirri present. . . . . . . . . . . . . . . . . . . . . . . . . . . . Caudiamphisiella (p. 129) - Caudal cirri absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amphisiella (p. 84) 7 (1) Limnetic1. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 - Terrestrial. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
1
If your “limnetic” specimen/population does not belong to Mucotrichidium hospes or Tetrastyla oblonga (see next couplet), select lead “terrestrial” because due to various events, for example, flooding, primarily terrestrial species can also occur in freshwater.
76
SYSTEMATIC SECTION
KEY TO TAXA
77
8 Body broadly spindle-shaped; cirral rows twisted; lives in spawn of evertebrates (Fig. 13d). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mucotrichidium (p. 441) - Body broad elliptical to roughly rectangular; cirral rows not twisted; reliable recorded only from New Zealand so far (Fig. 14e). . . . . . . . . . . Tetrastyla (p. 463) 9 (7) Cortical granules dark green or blue green; food vacuoles with parallelarranged bacteria; undulating membranes form acute angle (Fig. 14c, 93a, f). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Terricirra (p. 447) - No such combination of features. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 10 Two distinctly separated frontoventral rows; more than one buccal cirrus; postperistomial cirrus absent (Fig. 14d). . . . . . . . . . . . . . . . . . . . . Bistichella (p. 532) - No such combination of features. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 11 Two distinctly separated frontoventral rows; one buccal cirrus; postperistomial cirrus present (Fig. 15g). . . . . . . . . . . . . . . . . . . . . . . . . Pseudouroleptus1 (p. 658) - No such combination of features. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 12 One frontoventral row present (e.g., Fig. 12e, 15e). . . . . . . . . . . . . . . . . . . . . . 13 - No distinct frontoventral row present (e.g., Fig. 13b, 15c, d). . . . . . . . . . . . . . . 25 13 Frontoventral row composed of irregularly arranged cirri (Fig. 15e). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha (p. 636) - Frontoventral row composed of regularly arranged cirri (Fig. 12d, g, 13a, 15a, f). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 14 Frontoventral row ends at 50% of body length or behind (e.g., Fig. 12d, e). . . 15 - Frontoventral row ends at less than 50% of body length (Fig. 15b–e). . . . . . . . 19 15 Postperistomial cirrus present (Fig. 12e). . . . . . . . . . . . Hemiamphisiella (p. 288) - Postperistomial cirrus lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 16 More than one cirrus left of anterior portion of frontoventral row2 (Fig. 12d, 15f). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 - One cirrus left of anterior portion of frontoventral row (Fig. 12g, 13a). . . . . . . 18 17 Transverse cirri present; caudal cirri likely lacking (Fig. 12d). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Uroleptoides (p. 224) - No such combination of features. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17a
b Fig. 15a–d Incertae sedis in the non-oxytrichid Dorsomarginalia (sources of illustrations see individual descriptions). a: Nudiamphisiella interrupta; caudal cirri present. b: Erimophrya glatzeli; caudal cirri present. c: Vermioxytricha arenicola; caudal cirri absent. d: Hemiurosoma terricola; caudal cirri present. Fig. 15e Supplement to the Urostyloidea (source of illustration see individual description). e: Anteholosticha heterocirrata; caudal cirri lacking; cirri of frontoventral row arranged in zigzag pattern. Fig. 15f–h Supplement to the Oxytrichidae; dorsal kinety fragmentation present (sources of illustrations see individual descriptions). f: Amphisiellides atypicus; caudal cirri present. g: Pseudouroleptus caudatus; caudal cirri present. h: Ponturostyla enigmatica; caudal cirri absent. 1
See also Hemiamphisiella. Note that Uroleptoides multinucleatus rarely has only one cirrus left of the anterior portion of the amphisiellid median cirral row (Fig. 48f).
2
78
SYSTEMATIC SECTION
17a Transverse cirri present; caudal cirri present (Fig. 15f). . Amphisiellides (p. 651) - Transverse cirri lacking (do not confuse transverse cirri with a short longitudinal row in posterior body portion); caudal cirri present (Fig. 15a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nudiamphisiella (p. 560) 18 (16) Caudal cirri present (Fig. 12g). . . . . . . . . . . . . . . . . . Paramphisiella (p. 351) - Caudal cirri absent (Fig. 13a). . . . . . . . . . . . . . . . . . . . . . Afroamphisiella (p. 371) 19 (14) One to several postperistomial cirri present, that is, frontoventral row with distinct longitudinal break (Fig. 14c, 15b). . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 - Frontoventral row without longitudinal break (Fig. 12f, 13a, b, 15a). . . . . . . . . 21 20 1–3 postoral cirri; usually non-saline soil (Fig. 15b). . . . . . . Erimophrya (p. 577) - Usually five (3–6) postoral cirri; highly saline soil (Fig. 14c). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apourosomoida (p. 514) 21 (19) Transverse cirri present (e.g., Fig. 12f). . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 - Transverse cirri lacking (e.g., Fig. 13a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 22 Only one cirrus left of anterior portion of frontoventral row; two macronuclear nodules (Fig. 12f). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lamtostylides (p. 322) - No such combination of features. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 23 Two or four macronuclear nodules. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lamtostyla (p. 161) or Hemisincirra namibiensis (p. 418) - More than four macronuclear nodules. . . . . . . . . . . . . . . . . Hemisincirra (p. 387) 24 (21) Caudal cirri present (Fig. 15a). . . . . . . . . . . . . . . . . Nudiamphisiella (p. 560) - Caudal cirri absent (Fig. 13a). . . . . . . . . . . . . . . . . . . . . . Afroamphisiella (p. 371) 25 (12) Transverse cirri present (Fig. 15d). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 - Transverse cirri absent (Fig. 15c). . . . . . . . . . . . . . . . . . . Vermioxytricha (p. 596) 26 Frontoventral cirri arranged in Urosoma-pattern (Fig. 15d, 131i, h). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemiurosoma (p. 614) - Frontoventral cirri not arranged in Urosoma-pattern. . . . . . . . . . . . . . . . . . . . . 27 27 One cirrus left of anterior portion of frontoventral row; two macronuclear nodules (e.g., Fig. 66b). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lamtostylides (p. 322) - No such combination of features. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 28 Two macronuclear nodules (Fig. 12b, c). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lamtostyla granulifera-group (p. 205) and Lamtostyla longa-group (p. 218) - More than two macronuclear nodules (Fig. 13b). . . . . . . . . Hemisincirra (p. 387) 29 (1) About 6–8 marginal rows per side (Fig. 15h). . . . . . . . . Ponturostyla (p. 672) - Usually 1 marginal row per side (Spiroamphisiella with 1 long and 1 short right marginal row). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Amphisiellidae Jankowski, 1979 1979 1982 1985 1988 1994 1994 1996 1999 1999 2002 2005
Amphisiellidae fam. n. – Jankowski, Trudy zool. Inst., 86: 78 (original description; Table 4). Amphisiellidae n. fam.1 – Hemberger, Dissertation, p. 15 (original description; Table 5). Amphisiellidae n. fam.2 – Small & Lynn, Phylum Ciliophora, p. 457 (original description; Table 6). Amphisiellidae Jankowski, 1979 – Foissner, Stapfia, 17: 112 (description of three new genera). Amphisellidae Jankowski, 1979 – Tuffrau & Fleury, Traite de Zoologie, 2: 141 (incorrect subsequent spelling; Table 8). Amphisiellidae Jankowski, 1979 3 – Eigner & Foissner, J. Euk. Microbiol., 41: 254 (improved diagnosis; Table 9). Amphisiellidae – Petz & Foissner, Acta Protozool., 35: 276 (characterisation and key to amphisiellid genera; Table 10). Amphisiellidae Hemberger, 1982 – Shi, Acta Zootax. sin., 24: 252 (brief revision of hypotrichs in Chinese; Table 11). Amphisiellidae Hemberger, 1982 – Shi, Song & Shi, Progress in Protozoology, p. 99 (brief revision of hypotrichs in Chinese; Table 11). Amphisiellidae Jankowski, 1979 4 – Lynn & Small, Phylum Ciliophora, p. 450 (guide to ciliate genera; Table 12). Amphisiellidae Jankowski, 1979 – Berger, Int. Congr. Protozool., 12: 106 (brief note on amphisiellids).
Nomenclature: The taxon Amphisiellidae has been established three times as new family (see list of synonyms and remarks below). Now, the Amphisiellidae Jankowski, 1979 are generally accepted (e.g., Lynn & Small 2002). Characterisation: Hypotricha with a more or less long frontoventral row (= amphisiellid median cirral row) composed of an anterior portion formed by the anteriorly migrating cirri (= frontoterminal cirri) of anlage VI, a middle portion formed by the anteriormost cirrus/cirri of anlage IV, and a posterior portion formed by anlage V.5
1
Hemberger (1982) provided the following diagnosis: Hypotrichida mit mindestens je 1 Marginalreihe und mindestens 1 Ventralreihe; Cirren der Ventralreihe(n) nicht in der für die Familie Urostylidae Bütschli typischen Zick-Zack-Anordnung; ausdifferenzierte Frontalcirren vorhanden oder fehlend; alle Cirren bzw. Cirrenreihen entwickeln sich aus longitudinalen Anlagen. 2 Small & Lynn (1985) provided the following diagnosis: At least 1 or more frontoventral cirral files extends well past mid-ventrum. 3 Eigner & Foissner (1994) provided the following improved diagnosis: Euhypotrichina with an amphisiellid median cirral row of which the anterior segment is formed by cirri of the rightmost ventral anlage and the posteriormost segment by cirri of the second ventral anlage from right. A middle segment my be formed by neighboring anlagen. 4 Lynn & Small (2002) provided the following characterisation: Single ventral cirral file, except Pseudouroleptus; anterior segment of this file formed by cirri from rightmost ventral anlage and posterior segment from second ventral anlage from right. 5 Note that this characterisation reflects the situation in the last common ancestor of the Amphisiellidae. It does not exclude taxa with more or less distinct deviations (e.g., amphisiellid median cirral row composed of two portions only) because these features have very likely evolved only in subgroups of the amphisiellids). For plesiomorphies, see remarks.
79
80
SYSTEMATIC SECTION
The ground pattern of the Amphisiellidae: In the present chapter the supposed ground pattern of the Amphisiellidae is discussed. Briefly, the ground pattern of a monophylum (evolutionary unit) is the combination of features of the stem-species from which the monophylum evolved, that is, it is a summary of apomorphies and more or less young plesiomorphies (Ax 1995); old plesiomorphies, for example, the presence of cilia in the amphisiellids, are usually not included in the ground pattern. Apomorphy of the Amphisiellidae: Very likely the amphisiellids have only one morphological apomorphy which was recognised and discussed in detail by Eigner & Foissner (1994) for the first time. Amphisiellid median cirral row. This row is a mixed row, that is, it is composed of two or more (three in present case) true rows (see chapter 1.7 for terminology; Fig. 1f, 2a, 5a). Obviously, it evolved by an increase of the number of (i) frontoterminal cirri originating, as is usual, from anlage VI and (ii) postoral ventral cirri produced by anlage V. The “frontoterminal” cirri form the anterior portion, the “postoral ventral” cirri form the posterior portion of the amphisiellid median cirral row. In 18-cirri hypotrichs, cirrus IV/3 is usually more or less in line with the two frontoterminal cirri and the two postoral ventral cirri V/3 and V/4, indicating that the amphisiellid median cirral row was originally composed of three portions (Fig. 16b in Berger 2006). Such a state is still present, for example, in Hemiamphisiella (Fig. 58f). However, in most species the amphisiellid median cirral row is formed only by anlagen V and VI, that is, the anteriormost cirrus/cirri of anlage IV are no longer part of the row, but have been displaced leftwards to form, together with cirrus III/2, the conspicuous group “cirri left of the anterior portion of the amphisiellid median cirral”, a feature used to define several amphisiellid genera (Foissner 1988). The plesiomorphies of the Amphisiellidae are basically the same as for the Hypotricha, please refer to chapter 2.2 of the general section where the ground pattern of the Hypotricha is discussed in detail (see also next paragraph). Remarks: The characterisation above contains only the supposed apomorphy of the last common ancestor of the amphisiellids. The other features, for example, three frontal cirri, buccal cirrus and transverse and caudal cirri present, one left and one right marginal row, dorsal ciliature composed of bipolar kineties only, are plesiomorphies taken over from the ground pattern of the Hypotricha (chapter 2.2). The exact origin of the amphisiellids within the Hypotricha tree is still uncertain. However, the following hypothesis can be put forward: The Amphisiellidae branched off outside the Dorsomarginalia. Amphisiella, the name-bearing type of the group and some other “typical” amphisiellids (e.g., Lamtostyla) have a simple dorsal kinety pattern, that is, kineties originate by intrakinetal proliferation only (Fig. 9a). This pattern – which is also present, for example, in the urostyloids (Berger 2006) and Gonostomum (Berger 1999) – is the same as supposed for the last common ancestor of the Hypotricha and lacks dorsomarginal kineties, which are characteristic for the Dorsomarginalia Berger, 2006, a group comprising a major part of hypotrichs. Previously we supposed that this simple pattern, termed Gonostomum pattern by Berger & Foissner (1997) and Berger (1999), has evolved
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via the complex Oxytricha pattern (dorsomarginal kineties and kinety fragmentation present) by loss of (i) kinety fragmentation and (ii) dorsomarginal kineties. However, this explanation is not parsimonious and was mainly initiated by the assumption that the 18-frontal-ventral-transverse cirri pattern present in many oxytrichids (including Gonostomum), is an apomorphy of the Oxytrichidae. By contrast, I now suppose that the 18-cirri pattern is an apomorphy of the Hypotricha (Berger 2006, p. 33; details see chapter 2 of general section, Fig. 6a, 9a). The present hypothesis is supported by molecular data showing that Gonostomum, which basically has 18frontal-ventral-transverse cirri (for review, see Berger 1999), branches off very basally within the hypotrichs (Fig. 2 in Schmidt et al. 2007). The type of the amphisiellids, Amphisiella with the marine Amphisiella capitata as type species, was discovered by Pereyaslawzewa (1886). Kahl (1932), the first reviser, accepted Amphisiella and added some new species, but classified it as subgenus of Holosticha Wrześniowski, 1877, a typical urostyloid (for review, see Berger 2006, p. 88). Holosticha sensu Kahl was very inhomogeneous because basically nothing was known about the ontogenetic processes; all Oxytrichidae – that is, all Hypotricha according to my terminology – with “1–3 closed ventral cirral rows” have been included in Holosticha. Later, Amphisiella was again raised to genus level, but the classification in the holostichids, respectively, urostyloids was retained (e.g., Fauré-Fremiet 1961, Borror 1972, Stiller 1974a, Corliss 1979). Jankowski (1979) was the first who recognised Amphisiella as representative of a higher taxon and therefore established the Amphisiellidae. Hemberger (1982)1 came to the same conclusion and again established the Amphisiellidae because he did not know Jankowski’s paper, which was published in a little known and not widely distributed Russian journal just three years previously. Again three years later, Small & Lynn (1985) introduced the name Amphisiellidae for the third time as new family. Now, the Amphisiellidae are generally assigned to Jankowski (e.g., Eigner & Foissner 1994, Lynn & Small 2002). The classifications of the amphisiellids proposed by various authors are rather different (Tables 4–14). It is not possible to compare all these systems with the present classification (see contents). For a foundation of the exclusion of taxa from the amphisiellids, see the “Taxa not considered” chapter. Generally, the amphisiellids are a relatively small group compared to the urostyloids or oxytrichids. In addition, major taxa (e.g., Lamtostyla, Uroleptoides) are not very well defined because the type species are not studied in detail. To decide whether or not a species belongs to the amphisiellids, its ontogenesis has to be known because the main morphological apomorphy (amphisiellid median cirral row) is basically an ontogenetic feature: the row must be formed by cirri produced by anlage VI (anterior portion) and anlage V (posterior portion); in some species the row is tripartite because the middle portion is formed by anlage IV. Consequently, spe1
Hemberger (1982, p. 16) wrote that Borror (1972) assigned Amphisiella to the Amphisiellidae. However, Borror (1972) classified Amphisiella in the Urostylidae. Obviously, Hemberger (1982) equated the Urostylidae of Borror (1972) with his amphisiellids.
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SYSTEMATIC SECTION
cies which have a frontoventral row formed only by a single anlage, for example, Circinella or Orthoamphisiella, do not belong to the amphisiellids at the present state of knowledge. According to Eigner (1999, p. 46), Amphisiella marioni, a junior synonym of the type species of Amphisiella, belongs to the Oxytrichidae, that is, he synonymised the amphisiellids with the oxytrichids or he considered the amphisiellids as subgroup of the oxytrichids. However, as explained above, the amphisiellids and oxytrichids have, inter alia, a very different dorsal kinety pattern, indicating that Eigner’s (1999) proposal is incorrect. Eigner & Foissner (1994) made the most detailed study of amphisiellids; they significantly increased our knowledge about the cell division and introduced the term amphisiellid median cirral. They tried to estimate the phylogeny of the amphisiellids, but failed, inter alia, because they included taxa with rather different dorsal ciliature, for example, Gastrostyla, which has, indeed, a frontoventral row originating in the same way as the amphisiellid median cirral (Hemberger 1982, Foissner et al. 2002). Eigner & Foissner (1994) ignored the complex dorsal infraciliature of Gastrostyla (dorsomarginal rows and kinety fragmentation present vs. lacking in amphisiellids), which clearly shows that Gastrostyla is a true oxytrichid (for review, see Berger 1999, p. 789). Molecular data even show that Gastrostyla steinii Engelmann, 1862 – type of Gastrostyla – is a Stylonychinae (e.g., Foissner et al. 2004, Schmidt et al. 2007, Shao et al. 2007), that is, an oxytrichid with, inter alia, rigid body and lacking cortical granules (Berger & Foissner 1997, Berger 1999). Consequently, we have to assume that an “amphisiellid median cirral row” evolved at least twice independently. However, this is not the only complex cirral pattern which evolved convergently in the very long history of hypotrichs, which perhaps started at least about 400 million years ago (Wright & Lynn 1997). For example, the impressive midventral pattern characterising the urostyloids (no dorsomarginal kineties and no oxytrichid kinety fragmentation) has evolved convergently in Uroleptus (dorsomarginal kineties present, kinety fragmentation lacking; Foissner et al. 2004) and some oxytrichids, for example, Neokeronopsis (for review, see Berger 2006). Obviously, the evolutionary pressure is much greater on the ventral ciliature, which is involved in movement and very likely also in food uptake, than on the dorsal ciliature, which has possibly “no function” and is therefore more conservative. In the present book the Amphisiellidae are divided into three groups, excluding the genera classified as incertae sedis. Group I comprises all marine taxa which uniformly have a very prominent amphisiellid median cirral row (p. 84). These are the core amphisiellids. Group II includes all terrestrial genera which form their frontalventral-transverse ciliature, like group I genera, from the ordinary six anlagen (p. 161). This group also comprises species which have the plesiomorphic number of 18-frontal-ventral-transverse cirri (Fig. 12b, c). Group III comprises two terrestrial genera which have lost anlage IV and therefore produce only one cirrus left of the anterior portion of the amphisiellid median cirral row (p. 322). Although the amphisiellid median cirral row is a rather impressive feature I am not quite certain that
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it evolved only once and therefore unifies the amphisiellids as monophylum; perhaps it evolved independently in a marine and a terrestrial branch. Molecular data will perhaps show which assumption is correct. Unfortunately, I failed, like Eigner & Foissner (1994), to create a meaningful tree showing the relationships of the amphisiellid genera, most of which are more or less homogenous in the present review. A relatively high number of genera is classified as incertae sedis in the amphisiellids (see contents), inter alia, some difficult taxa like Hemisincirra. This uncertain classification is mainly due to the lack of a distinct “amphisiellid median cirral row” and the lack of ontogenetic data showing the formation of the cirral row. Amphisiellides is excluded from the amphisiellids and preliminarily classified in the oxytrichids because the type species very likely forms one kinety via dorsal kinety fragmentation; in addition, the frontoventral row originates from a single anlage, preventing the classification in the amphisiellids (Fig. 135a–e). By contrast, Nudiamphisiella is excluded from the Amphisiellidae because a dorsomarginal kinety is present (Fig. 119i, 120o, r–t). Since I do not know to which higher taxon it belongs, I preliminarily classify it as incertae sedis in the non-oxytrichid Dorsomarginalia, together with some deviating “oxytrichids” discovered by Foissner et al. (2002). I suppose that these genera are not oxytrichids because they lack dorsal kinety fragmentation. Since their cirral pattern shows some resemblance to amphisiellids, they are included in the present book. However, these genera likely do not form a monophyletic group (further details, see p. 560). Just recently, Paiva & Silva-Neto (2007) published an important paper about Strongylidium Sterki, 1878. They redescribed S. pseudocrassum Wang & Nie, 1935 and found a great resemblance to Hemiamphisiella and Pseudouroleptus, inter alia, because all these taxa have a postperistomial cirrus (= cirrus IV/2). However, there are some distinct differences in ontogenetic features so that synonymy can be excluded. Strongylidium pseudocrassum and Hemiamphisiella are obviously closely related because both lack a dorsal kinety fragmentation (vs. present in Pseudouroleptus; Fig. 136c) and the frontoventral row is formed by the anlagen IV (forms middle portion of row), V (rear portion), and VI (anterior portion). Unfortunately, the type species of Strongylidium is not described in detail so that the phylogenetic position of this genus remains uncertain. Probably, relevant molecular biological data will give us a better insight into the systematics of this little-known group. Anyhow, the strongylidids, which are perhaps closely related to the amphisiellids, will be treated in a later volume of the monographic series.
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SYSTEMATIC SECTION
Group I: Marine Amphisiellids This rather small group comprises the core amphisiellids, marine hypotrichs with a prominent amphisiellid median cirral row formed from two or three anlagen. Besides Amphisiella, the name-bearing group of the amphisiellids, three monotypic genera are included, namely (i) Caudiamphisiella, a new genus establish for an Amphisiella-like species with caudal cirri; (ii) Maregastrostyla, also a new genus for Gastrostyla pulchra, which lacks dorsal kinety fragmentation and dorsomarginal rows, features present in G. steinii, type of Gastrostyla; and (iii) Spiroamphisiella, which has a strongly twisted body. Although the cirral pattern of these taxa is rather homogenous, I am not quite certain that they form a monophyletic group. If yes, a subgroup comprising these marine amphisiellids should be established. However, molecular data should be awaited to show whether or not they cluster together.
Amphisiella Gourret & Roeser, 1888 1888 Amphisiella, nov. gen. – Gourret & Roeser, Archs Biol., 8: 180 (original description; no formal diagnosis provided). Type species (by monotypy): Amphisiella marioni Gourret & Roeser, 1888. 1932 Amphisiella Gourret und Roeser, 1888 – Kahl, Tierwelt Dtl., 25: 589 (revision of hypotrichs; see nomenclature). 1933 Amphisiella Gourret & Roeser, 1888 – Kahl, Tierwelt N.- u. Ostsee, 23: 112 (guide to marine ciliates; see nomenclature). 1961 Amphisiella Gourret & Roeser – Corliss, Ciliated protozoa, p. 169 (revision). 1972 Amphisiella Gourret & Roeser, 18871 – Borror, J. Protozool., 19: 9 (revision of hypotrichs). 1974 Amphisiella Gourret & Roeser – Stiller, Fauna Hung., 115: 94 (revision). 1979 Amphisiella Gourret et Roeser, 1888 – Jankowski, Trudy zool. Inst., Leningr., 86: 50 (generic catalogue of hypotrichs). 1979 Amphisiella Gourret & Roeser, 1888 – Tuffrau, Trans. Am. microsc. Soc., 98: 526 (revision of hypotrichs). 1979 Amphisiella Gourret & Roeser, 1888 – Corliss, Ciliated protozoa, p. 309 (revision). 1982 Amphisiella Gourret & Roeser, 18882 – Hemberger, Dissertation, p. 22 (revision of non-euplotid hypotrichs). 1983 Amphisiella Gourret & Roeser, 1888 – Curds, Gates & Roberts, British and other freshwater ciliated protozoa, p. 398 (guide to ciliate genera). 1985 Amphisiella – Small & Lynn, Phylum Ciliophora, p. 457 (guide to ciliate genera). 1987 Amphisiella Gourret et Roeser, 1888 – Tuffrau, Annls Sci. nat. (Zool.), 8: 115 (revision of hypotrichs). 1988 Amphisiella Gourret & Roeser, 18883 – Foissner, Stapfia, 17: 112 (improved diagnosis and discussion). 1
Borror (1972) provided the following diagnosis: Cirri in 3 ventral rows plus frontal and transverse cirri. Usually 2 macronuclei. 2 Hemberger (1982) provided the following diagnosis: Je 1 rechte und linke Marginalreihe; 1 Ventralreihe; deutlich differenzierte Frontal- und Transversalcirren; Morphogenesebeginn am linken Transversalcirrus; Frontalcirrenentwicklung aus 5 longitudinalen Anlagen; Anlagenentwicklung der Ventralreihe in Verlauf der bestehenden. 3 Foissner (1988) provided the following improved diagnosis: Amphisiellidae mit mehr als 1 Cirrus links der Ventralreihe im Frontalfeld. Transversalcirren vorhanden.
Amphisiella
85
1992 Amphisiella Gourret and Roeser, 1888 – Carey, Marine interstitial ciliates, p. 178 (guide). 1992 Amphisiella Gourret and Roeser, 18881 – Voß, Europ. J. Protistol., 28: 413 (improved diagnosis). 1994 Amphisiella Gourret & Roeser, 18882 – Eigner & Foissner, J. Euk. Microbiol., 41: 255 (improved diagnosis). 1996 Amphisiella Gourret and Roeser, 18883 – Petz & Foissner, Acta Protozool., 35: 277 (improved diagnosis). 1999 Amphisiella Gourret & Roeser, 1888 – Shi, Acta Zootax. sinica, 24: 255 (generic revision of hypotrichs). 1999 Amphisiella Gourret & Roeser, 1888 – Shi, Song & Shi, Progress in Protozoology, p. 102 (generic revision of hypotrichs). 2001 Amphisiella Gourret & Roeser 1888 – Aescht, Denisia, 1: 21 (catalogue of generic names of ciliates). 2001 Amphisiella Gourret and Roeser, 1888 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 7 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Amphisiella Gourret and Roeser, 1888 – Lynn & Small, Phylum Ciliophora, p. 454 (guide to ciliate genera). 2006 Amphisiella Gourret & Roeser, 1888 – Berger, Monographiae biol., 85: 1208 (brief note about the systematic position).
Nomenclature: Amphisiella is the diminutive of the genus-group name Amphisia Sterki, 1878 (Gourret & Roeser 1888), a junior synonym of Holosticha Wrześniowski, 1877 (for derivation of Amphisia, see Berger 2006, p. 89). Feminine gender because ending with the Latin suffix -ella (ICZN 1999, Article 30.1.3). Nominotypical genus of the Amphisiellidae. Kahl (1932, p. 571, 589; 1933) classified Amphisiella as subgenus of Holosticha, that is, the correct spelling in Kahl’s papers is Holosticha (Amphisiella) Gourret & Roeser, 1888. Incorrect subsequent spellings: Amphisiela (Tirjaková 1988, p. 501); Amphysiella Gourret & Roeser (Stiller 1974, p. 46). Characterisation (A = supposed apomorphy): Amphisiellidae with continuous adoral zone of membranelles. Undulating membranes straight and parallel. Three enlarged frontal cirri. Buccal cirrus present. Two or more cirri left of anterior portion of amphisiellid median cirral row, which originates from anlage V (posterior portion) and VI (anterior portion). Postperistomial cirrus lacking. Usually two pretransverse ventral cirri. Five or more prominent transverse cirri. One left and one right marginal row. More than three dorsal kineties (A) each originating intrakinetally. Caudal cirri lacking. Saltwater. 1 Voß (1992) provided the following improved diagnosis: Amphisiellidae with 1 right and left marginal row, 1 ventral row and more than 1 fronto-ventral cirrus left of the ventral row. Caudal cirri missing, at least 2 transverse cirri. During morphogenesis the ventral row is composed of two anlagen, frontal cirri and fronto-ventral cirri are produced in 4 anlagen. 2 Eigner & Foissner (1994) provided the following improved diagnosis: The amphisiellid median cirral row (ACR) originates from two rightmost anlagen. More than one cirrus left of ACR. Transverse cirri obliquely arranged, originate from more than one anlage. Caudal cirri lacking. 3 Petz & Foissner (1996) provided the following improved diagnosis: The oral primordium originates in close contact with the amphisiellid median cirral row (ACR). The ACR commences anlagen formation within-row and originates from two rightmost anlagen. All dorsal kineties develop intrakinetally. More than one cirrus left of ACR. Transverse cirri obliquely arranged, originate from more than one anlage. Caudal cirri lacking.
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SYSTEMATIC SECTION
Table 15 Morphometric data on Amphisiella annulata (an1, from Berger 2004; an2, an3, year 1996 and year 2000 population from Hu et al. 2004) Characteristics a Body, length
Body, width
Body length:width, ratio Adoral zone of membranelles, length
Adoral zone, relative length (%) Anterior body end to distal end of adoral zone, distance Anterior body end to paroral, distance Paroral, length Anterior body end to buccal cirrus, distance Anterior body end to cirrus III/2, distance Anterior body end to anlagen IV cirri b, distance Anterior body end to amphisiellid median cirral row, distance Anterior body end to left marginal row, distance Posterior body end to left marginal row, distance Anterior body end to right marginal row, distance Posterior body end to right marginal row, distance Rearmost transverse cirrus to rear body end, distance Anterior macronuclear nodule, length
Species
mean
M
SD
SE
CV
Max
n
an1 an2 an3 an1 an2 an3 an1 an1 an2 an3 an1 an1
96.4 167.3 152.8 37.4 59.3 42.1 2.6 40.6 65.7 55.3 42.3 11.3
97.0 – – 36.0 – – 2.5 40.0 – – 42.9 11.5
12.3 24.7 21.9 7.7 10.6 9.7 0.4 5.1 8.6 10.0 4.0 3.5
2.3 6.4 4.5 1.4 2.7 2.1 0.1 0.9 2.2 2.0 0.7 0.7
12.8 68.0 121.0 14.8 120.0 203.0 14.3 120.0 200.0 20.5 23.0 55.0 17.9 45.0 80.0 23.1 28.0 64.0 15.9 2.0 3.6 12.5 26.0 50.0 13.1 48.0 78.0 18.1 42.0 78.0 9.4 33.3 49.5 31.0 4.0 19.0
30 15 24 29 15 22 29 29 15 25 29 28
an1 an1 an1
11.3 17.9 9.4
12.0 18.0 9.0
3.1 2.1 3.0
0.6 0.4 0.6
27.3 11.9 31.9
5.0 12.0 4.0
17.0 22.0 18.0
28 27 29
an1
12.7
12.0
3.7
0.7
29.0
7.0
22.0
28
an1
15.9
16.0
3.6
0.7
22.5
9.0
25.0
30
an1
12.4
12.0
3.6
0.7
28.9
4.0
21.0
28
an1
20.3
21.0
4.4
0.8
21.7
10.0
30.0
30
an1
7.0
7.0
2.0
0.4
28.8
4.0
12.0
28
an1
24.2
24.0
4.6
0.9
18.9
16.0
36.0
30
an1
6.2
6.0
1.6
0.3
25.7
4.0
11.0
27
an1
4.5
4.0
1.6
0.3
35.4
2.0
8.0
30
19.2 24.3 34.2 6.9 12.2 17.8 5.2 18.1 7.7 3.4 1.7 2.0 2.0 2.0 2.6
18.0 – – 7.0 – – 4.5 18.0 8.0 3.0 1.6 2.0 – – 2.5
3.6 8.5 5.2 1.2 1.6 4.9 3.0 3.1 1.3 0.8 0.3 0.0 0.0 0.0 1.0
0.7 2.2 1.0 0.2 0.4 1.0 0.5 0.6 0.2 0.1 0.1 0.0 0.0 0.0 0.2
18.6 34.8 15.0 17.0 12.9 0.3 57.8 17.2 16.8 22.7 17.8 0.0 0.0 0.0 39.2
14.0 16.0 27.0 4.0 9.0 12.0 1.0 13.0 5.0 2.0 1.5 2.0 2.0 2.0 1.0
29.0 35.0 44.0 9.0 15.0 28.0 12.0 27.0 10.0 5.0 2.5 2.0 2.0 2.0 5.0
30 15 25 30 15 25 30 30 30 30 30 30 15 25 30
an1 an2 c an3 c Anterior macronuclear nodule, width an1 an2 c an3 c Macronuclear nodules, distance in between an1 Posterior macronuclear nodule, length an1 Posterior macronuclear nodule, width an1 Anteriormost micronucleus, length an1 Anteriormost micronucleus, width an1 Macronuclear nodules, number an1 an2 an3 Micronuclei near anterior an1 macronuclear nodule, number
Min
Amphisiella
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Table 15 Continued Characteristics a Micronuclei near posterior macronuclear nodule, number Micronuclei, total number Adoral membranelles, number
Frontal cirri, number
Buccal cirri, number
Cirri behind right frontal cirrus, number Anlagen IV cirri b, number
Amphisiellid median cirral row, number of cirri Pretransverse ventral cirri, number
Transverse cirri, number
Left marginal cirri, number
Right marginal cirri, number
Dorsal kineties, number
Species
mean
M
SD
SE
CV
Min
Max
n
an1
3.0
2.5
1.3
0.2
42.9
1.0
6.0
30
an1 an3 an1 an2 an3 an1 an2 an3 an1 an2 an3 an1 an2 an3 an1 an2 an3 an1 an2 an3 an1 an2 an3 an1 an2 an3 an1 an2 an3 an1 an2 an3 an1 an2 an3
5.6 3.5 47.2 52.2 44.7 3.0 3.0 3.0 1.0 1.0 1.0 1.0 1.0 1.0 3.1 3.0 3.0 44.5 49.9 42.5 2.0 2.0 2.0 6.0 5.9 6.0 34.6 40.9 36.5 34.1 42.4 34.9 6.4 8.2 7.7
6.0 – 48.0 – – 3.0 – – 1.0 – – 1.0 – – 3.0 – – 45.0 – – 2.0 – – 6.0 – – 34.0 – – 34.0 – – 6.0 – –
1.3 1.0 5.9 5.7 6.6 0.2 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.3 0.0 0.0 5.7 13.0 6.0 0.0 0.0 0.0 0.2 0.5 0.0 3.5 3.5 7.2 3.2 4.3 6.7 0.6 0.5 0.6
0.2 0.2 1.1 1.5 1.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1.2 5.8 1.3 0.0 0.0 0.0 0.0 0.1 0.0 0.6 0.9 1.6 0.6 1.1 1.5 0.1 0.1 0.1
23.5 28.2 12.4 10.9 14.7 6.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 8.3 0.0 0.0 12.9 26.2 14.1 0.0 0.0 0.0 3.1 7.7 0.0 10.0 8.5 19.8 9.3 10.1 19.2 9.2 6.5 7.6
3.0 2.0 31.0 41.0 33.0 3.0 3.0 3.0 1.0 1.0 1.0 1.0 1.0 1.0 3.0 3.0 3.0 25.0 45.0 33.0 2.0 2.0 2.0 5.0 5.0 6.0 25.0 36.0 25.0 28.0 36.0 22.0 6.0 8.0 7.0
8.0 6.0 57.0 62.0 57.0 4.0 3.0 3.0 1.0 1.0 1.0 1.0 1.0 1.0 4.0 3.0 3.0 54.0 61.0 56.0 2.0 2.0 2.0 6.0 7.0 6.0 41.0 46.0 53.0 41.0 49.0 48.0 8.0 10.0 8.0
30 21 28 15 24 30 15 25 30 15 25 30 15 25 30 15 25 24 15 20 28 15 25 29 15 20 29 15 21 27 15 21 23 15 19
a All measurements in µm. CV = coefficient of variation in %, M = median, Max = maximum value, mean = arithmetic mean, Min = minimum value, n = number of individuals investigated, SD = standard deviation, SE = standard error of arithmetic mean. Data based on protargol-impregnated specimens. b
This is the short cirral row left of the anterior portion of the amphisiellid median cirral row.
c
Macronuclear nodule (anterior or posterior) not specified.
Additional characters: Body flexible, broad to elongate elliptical in outline; contractile vacuole indistinct or lacking; dorsal bristles less than 5 µm long. Remarks: The systematics of Amphisiella is complicated. For a long time this group was a melting pot for species from all habitats having a more or less long and
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longitudinally arranged frontoventral row. Twenty years ago, Foissner (1988) established three genera (Amphisiellides, Paramphisiella, Hemiamphisiella) to include largely Amphisiella-like species, mainly from terrestrial habitats. So far, more than 30 species/subspecies have been originally assigned or transferred to Amphisiella (Berger 2001; see also permanently updated version at www.protozoology.com). However, the present review shows that only the type species of Amphisiella and the remaining saltwater species form a rather homogenous group due to the distinctly increased number of dorsal kineties (Table 15). In addition, they have taken over the five prominent transverse cirri (in some species also the pretransverse ventral cirri) from the ground pattern of the hypotrichs. Consequently, only five species are assigned to Amphisiella in the present book (see below). By contrast, the other species assigned to Amphisiella so far are terrestrial, usually have the plesiomorphic number of three dorsal kineties (rarely four kineties are present), usually lack the pretransverse ventral cirri, and the transverse cirri are normally less prominent than in the saltwater species. Further, many terrestrial species have, likely as an adaptation to the limited amount of space in the soil pores, a slender body. In recent years, many terrestrial amphisiellids have been assigned either to Amphisiella or Lamtostyla, which differ, according to Petz & Foissner (1996), only in the beginning of the oral primordium formation. However, since the ontogenesis of L. lamottei, type of Lamtostyla, is not known, the feature origin of oral primordium cannot be used for the separation at the present state of knowledge. Synonymy of Uroleptoides and Amphisiella was first suggested by Borror (1972, p. 9) and later accepted by Jankowski (1979) and Foissner & Blatterer (1990). However, this synonymy is unlikely because of some distinct differences in the cirral pattern (see U. kihni, type of Uroleptoides). Amphisiella was classified in various higher taxa. The assignment to the Oxytrichidae by Kahl (1932) is meaningless because he classified all taxa, except for the euplotids, in the oxytrichids. Fauré-Fremiet (1961, p. 3517), Stiller (1974a, p. 130), Corliss (1977, p. 137; 1979), Tuffrau (1979, 1987), Curds et al. (1983), and Carey (1992) have classified Amphisiella in the Holostichidae, a subgroup of the urostyloids (for review, see Berger 2006). This assignment is certainly incorrect because amphisiellids have the ordinary, plesiomorphic number of six frontal-ventraltransverse cirri anlagen (I–VI), and the cirri of the two rightmost anlagen form a more or less continuous row. By contrast, the urostyloids have increased the number of anlagen which form the zigzagging midventral pairs, the main morphological apomorphy of the urostyloids. Borror (1972) also assigned Amphisiella to the Urostylidae, but later transferred it to the Oxytrichidae (Borror 1979, p. 546), a classification also suggested by Eigner (1997, p. 555; 1999, p. 46) and overlooked by Berger (1999, p. 893) in his “Taxa not considered in the Oxytrichidae” chapter. Foissner & Foissner (1988, p. 81) assigned Amphisiella to the Kahliellidae Tuffrau, 1979, without, however, giving an explanation. Shi (1993) classified Amphisiella and some
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other genera in the Rootletphorida, which is characterised, inter alia, by “leftward transverse rootlet fiber in the marginal cirral base”. Jankowski (1979), Hemberger (1982), and Small & Lynn (1985) considered Amphisiella as type of a separate family (see Amphisiellidae section for details and problems on nomenclature). This classification is accepted by Eigner & Foissner (1994), Tuffrau & Fleury (1994, p. 141), Petz & Foissner (1996), Shi (1999), Shi et al. (1999), Lynn & Small (2002), and myself (further details see remarks at the Amphisiellidae). Species included in Amphisiella (alphabetically arranged basionyms are given): (1) Amphisiella ovalis Fernandez-Leborans & Novillo, 1992; (2) Amphisiella turanica Alekperov & Asadullayeva, 1999; (3) Holosticha annulata Kahl, 1928b; (4) Holosticha (Amphisiella) milnei Kahl, 1932; (5) Oxytricha capitata Pereyaslawzewa, 1886. Species misplaced in Amphisiella: For a long time Amphisiella was a melting pot for species with a frontoventral row, a variable number of cirri left of the anterior portion of this row, and with or without transverse and caudal cirri. In the present review, Amphisiella is confined to saltwater species with distinct transverse cirri and without caudal cirri (details see Amphisiellidae and remarks of genus section). Thus, the following species originally assigned to Amphisiella are very likely misplaced in this genus (if you do not find a certain species in the list below, check the systematic index): Amphisiella acuta Foissner, 1982. Remarks: Now Paramphisiella acuta (p. 352). Amphisiella antarctica Wilbert & Song, 2005. Remarks: Now Caudiamphisiella antarctica (p. 129). Amphisiella arenicola Fernandez-Leborans & Novillo, 1992. Remarks: Classified as species indeterminata (p. 127). Amphisiella australis Blatterer & Foissner, 1988. Remarks: Now Lamtostyla australis (p. 169). Amphisiella binucleata multicirrata Foissner, Agatha & Berger, 2002. Remarks: Now Uroleptoides binucleatus multicirratus (p. 267). Amphisiella dorsicirrata Foissner. Remarks: A nomen nudum mentioned in a faunal list by Foissner (1981, p. 18); details see Berger (1999, p. 803). Amphisiella elegans Foissner, Agatha & Berger, 2002. Remarks: Now Lamtostyla elegans (p. 192). Amphisiella faurei Dragesco, 1963. Remarks: Now Marginotricha faurei (Dragesco, 1963) Lin, Song & Warren, 2004 (p. 144). Detailed redescription, see Wicklow (1982b, p. 319) as Psammocephalus faurei (Dragesco, 1963) Wicklow, 1982. Will be treated in a later volume of the monograph of hypotrichs. Amphisiella lithophora Fauré-Fremiet, 1954. Remarks: Now Marginotricha lithophora (Fauré-Fremiet, 1954) Lin, Song & Warren, 2004 (p. 144). Detailed redescription see Wicklow (1982b, p. 320) as Psammocephalus lithophora (FauréFremiet, 1954) Wicklow, 1982b. Will be treated in a later volume of the monograph of hypotrichs.
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Amphisiella longiseries Foissner, Agatha & Berger, 2002. Remarks: Now Uroleptoides longiseries (p. 249). Amphisiella magnigranulosa Foissner, 1988. Remarks: Now Uroleptoides magnigranulosus (p. 273). Amphisiella multinucleata Foissner, Agatha & Berger, 2002. Remarks: Now Uroleptoides multinucleatus (present book, p. 255). Amphisiella namibiensis Foissner, Agatha & Berger, 2002. Remarks: Now Bistichella namibiensis (p. 538). Amphisiella oscensis Fernandez-Leborans, 1984. Remarks: Incertea sedis in Caudiamphisiella (p. 133) Amphisiella polycirrata Berger & Foissner, 1989. Remarks: Now Uroleptoides polycirratus (p. 285). Amphisiella procera Foissner, Agatha & Berger, 2002. Remarks: Now Lamtostyla procera (p. 183). Amphisiella quadrinucleata Berger & Foissner, 1989. Remarks: Now Lamtostyla quadrinucleata (p. 190). Amphisiella raptans Buitkamp & Wilbert, 1974. Remarks: Now Uroleptoides raptans (p. 252). Amphisiella terricola Gellért, 1956a. Remarks: Now Uroleptoides terricola (p. 279). Holosticha thiophaga Kahl, 1928. Remarks: This species was assigned to the subgenus Holosticha (Amphisiella) by Kahl (1932). Now classified as junior synonym of Holosticha diademata (Rees, 1884) Kahl, 1932 (for review, see Berger 2006, p. 115).
Key to Amphisiella species The identification of the five marine species is not very difficult. It should be possible to determine them after detailed live observations. If caudal cirri are present, see Caudiamphisiella; if the body is strongly twisted, see Spiroamphisiella; and if the transverse cirri are distinctly displaced anteriad, see Maregastrostyla. 1 More than 2 macronuclear nodules (Fig. 20a, 21a). . . . . . . . . . . . . . . . . . . . . . . . 4 - Two macronuclear nodules. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2 Ring-shaped structures present; anterior body portion not cephalised (Fig. 17b, g, n, 19a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 - Ring-shaped structures lacking; anterior body portion more or less distinctly cephalised (Fig. 16a–c). . . . . . . . . . . . . . . . . . . . . . . . . . Amphisiella capitata (p. 91) 3 Several ring-shaped structures usually present (Fig. 17a, b, g). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amphisiella annulata (p. 100) - One ring-shaped structure each in anterior and posterior body portion (posterior ring sometimes lacking) (Fig. 19a). . . . . . . . . . . . . . . Amphisiella milnei (p. 118)
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4 (1) Four macronuclear nodules (Fig. 20a). . . . . . . . Amphisiella turanica (p. 123) - 32–45 macronuclear nodules (Fig. 21a). . . . . . . . . . . . Amphisiella ovalis (p. 125)
Amphisiella capitata (Pereyaslawzewa, 1886) Borror, 1972 (Fig. 16a–e, g–w, Addenda) 1886 Oxytricha capitata n. sp. – Pereyaslawzewa, Zap. novoross. Obshch. Estat., 10: 91, Fig. 15 (Fig. 16a; original description in Russian [I have no translation]; no formal diagnosis provided and no type material available). 1888 Amphisiella marioni, nov. spec. – Gourret & Roeser, Archs Biol., 8: 180, Planche XIV, Fig. 9 (Fig. 16b; original description of synonym; no formal diagnosis provided and no type material available). 1923 Amphisia marioni, Gourret u. Roeser 18881 – Mansfeld, Arch. Protistenk., 46: 128, Fig. 14a–c (Fig. 16c–e; detailed redescription and combination with Amphisia). 1929 Amphisia (Oxytricha) capitata – Hamburger & Buddenbrock, Nord. Plankt., 7: 91 (guide to marine plankton; combination with Amphisia). 1932 Amphisiella (Oxytricha) capitata (Perejaslawzewa, 1885) – Kahl, Tierwelt Dtl., 25: 589, Fig. 10624 (Fig. 16g; revision; combination with Holosticha; see nomenclature for correct name; incorrect year). 1932 Amphisiella marioni Gourret und Roeser, 1888 – Kahl, Tierwelt Dtl., 25: 589, Fig. 1111 (Fig. 16i; revision; combination with Holosticha, see nomenclature for correct name). 1932 Amphisiella marioni Mansfeld, 1926 – Kahl, Tierwelt Dtl., 25: 590, Fig. 10623 (Fig. 16k; incorrect author and year; see nomenclature and remarks). 1933 Amphisiella capitata (Perejaslawzewa 1885) – Kahl, Tierwelt N.- u. Ostsee, 23: 112, Fig. 17.33 (Fig. 16h; guide to marine ciliates; see nomenclature for correct name; incorrect year). 1933 Amphisiella marioni (Mansfeld 1926) – Kahl, Tierwelt N.- u. Ostsee, 23: 112, Fig. 17.34 (Fig. 16j; guide to marine ciliates; see nomenclature for correct name). 1933 Amphisiella marioni (Gourret & Roeser 1888) – Kahl, Tierwelt N.- u. Ostsee, 23: 112, Fig. 17.35 (Fig. 16l; guide to marine ciliates; incorrect author and year; see nomenclature and remarks). 1972 Amphisiella capitata (Perejaslawzewa, 1886) Kahl, 1932 – Borror, J. Protozool., 19: 9, Fig. 12 (Fig. 16w; combination with Amphisiella, that is, incorrect combining author, see nomenclature; revision of hypotrichs). 1972 Amphisiella marioni Gourret & Roeser, 1887 – Borror, J. Protozool., 19: 9 (revision of hypotrichs; incorrect year). 1979 Amphisiella marioni – Borror, J. Protozool., 26: 548, Fig. 5 (Fig. 16u, v; illustration of infraciliature). 1982 Amphisiella marioni Gourret and Roeser, 1887 – Wicklow, Protistologica, 18: 321, Fig. 45a–h (Fig. 16m–t; redescription after protargol impregnation and description of cell division; incorrect year). 1982 Amphisiella capitata (Perejaslawzewa, 1886) Kahl, 1932 – Hemberger, Dissertation, p. 22 (revision of hypotrichs). 1985 Amphisiella marioni – Small & Lynn, Phylum Ciliophora, p. 457, Fig. 28A (Fig. 16u; guide to ciliate genera).
1
Mansfeld (1923) provided the following diagnosis: Körper kontraktil und langgestreckt, 150–200 µ lang und 40–50 µ breit. Das erste Viertel durch beiderseitige Einschnürung seitlich abgesetzt. Rückenseite gewölbt und ohne Wimpern und Borsten. Bauchseite abgeflacht. Peristom im ersten Drittel der Bauchseite reicht nicht bis an den Vorderrand. Der linke Rand trägt eine undulierende Membran, der rechte Membranellen, von denen 12 dicht am Vorderrand deutlich hervortreten. Zwei Randcirren- und einen Bauchcirrenreihe. 7 Aftercirren.
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Fig. 16a–e, g–l Amphisiella capitata (a, from Pereyaslawzewa 1886; b, from Gourret & Roeser 1888; c–e, from Mansfeld 1923; g, h, after Pereyaslawzewa 1886 from Kahl 1932, 1933; i, j, after Gourret & Roeser 1888 from Kahl 1932, 1933; k, l, after Mansfeld 1923 from Kahl 1932, 1933. a–c, g–l, from life; d, e, neutral red staining). a–c, g–l: Ventral views showing, inter alia, cirral pattern and macronuclear nodules (c). Long arrow in (c) marks right marginal row, short arrow denotes amphisiellid median cirral row. Individ-
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1994 Amphisiella marioni Gourret & Roeser, 1888 – Eigner & Foissner, J. Euk. Microbiol., 41: 255 (review of the amphisiellids). 1997 Amphisiella marioni Gourret & Roeser, 1888 – Eigner, J. Euk. Microbiol., 44: 561, Fig. 19 (Fig. 16m; analysis of cell division). 1999 Oxytricha capitata Pereyaslawzewa, 1886 – Berger, Monographiae biol., 78: 243 (brief note). 2001 Oxytricha capitata Pereyaslawzewa, 1886 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 53 (see nomenclature).
Nomenclature: The species-group name capitát·us, -a, -um (Latin adjective [m, f, n]; having a head [caput]; Hentschel & Wagner 1996) obviously refers to the headlike anterior body portion. The species-group name marioni is very likely derived from a personal name. Kahl (1932, 1933) classified Amphisiella as subgenus of Holosticha. Thus, the correct names in his reviews are Holosticha (Amphisiella) capitata (Pereyaslawzewa, 1886) Kahl, 1932 and Holosticha (Amphisiella) marioni (Gourret & Roeser, 1888) Kahl, 1932. Borror (1972) and Carey (1992) obviously overlooked that Kahl (1932) had transferred Oxytricha capitata to Holosticha, and not to Amphisiella (see list of synonyms). For the sake of simplicity I suggest to fix Borror (1972) as combining author (both for A. capitata and A. marioni), although he did not formally transfer it to Amphisiella. In the “Catalogue of ciliate names” I overlooked this combination (Berger 2001, p. 53). Kahl (1932, 1933) considered Mansfeld’s (1923) population as distinct species (see list of synonyms and remarks). The correct heading in his papers would have been “Amphisiella marioni sensu Mansfeld (1923)”. The incorrect authorship “Mansfeld 1926” introduced by Kahl (1932) was also used by Dragesco (1953), Vacelet (1961b), and Agamaliev (1967, 1970). Amphisiella capitata was incorrectly assigned to Kahl by Dzhurtubayev (1978, p. 65). Remarks: None of the two species (Oxytricha capitata, Amphisiella marioni) included has been redescribed in detail with modern methods. The available data caused Hemberger (1982) to synonymise them, a decision preliminary accepted in the present review. However, new data should be awaited before a final judgement is passed on this problem because there are some more or less distinct differences in the original descriptions (see below). Just recently, Li et al. (2007) described and neotypified A. marioni (see Addenda). Pereyaslawzewa (1886) discovered the present species in the Black Sea and described it in a little known Russian journal. Almost simultaneously, Gourret & Roeser (1888) found the same, or a very similar, species in the Mediterranean Sea. I suggest that the French workers did not have the recent original description of Oxytricha capitata and therefore also described it as new species. Interestingly, in both
b
ual sizes not indicated. d, e: Ventral and dorsal view after staining showing the cirral pattern and, very likely, dorsal bristle rows, although Mansfeld wrote that no bristles are present. Page 91. Fig. 16f Amphisiella capitata (from Carey 1992). Insufficient redescription because amphisiellid median cirral row obviously lacking. Page 128.
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cases the nuclear apparatus was not described, so that Kahl (1932) supposed that both species have many macronuclear nodules which are difficult to see in life. Kahl recognised the similarity of O. capitata and A. marioni, but distinguished them by the number of transverse cirri (5 vs. 7). However, the illustration by Pereyaslawzewa shows that she did not clearly separate the transverse cirri from the posterior portion of the left marginal row, so this feature should not be over-interpreted. Moreover, one cannot exclude that Gourret & Roeser (1888) summarised (two?) pretransverse ventral cirri and the (five?) transverse cirri, whereas Pereyaslawzewa (1886) obviously overlooked the difficult-to-recognise pretransverse ventral cirri. The body shape, the remaining cirral pattern, and the habitat agree very well. Noteworthy differences concern the contractile vacuole (left of proximal portion of adoral zone in O. capitata vs. lacking or, if present at all, then behind mid-body in A. marioni) and the distal end of the adoral zone (extends far posteriorly [Fig. 16a] vs. not very far [Fig. 16b]). Mansfeld (1923) redescribed Gourret & Roeser’s species and transferred it to Amphisia, a junior synonym of the urostyloid Holosticha (Berger 2006, p. 88). Some years later, Hamburger & Buddenbrock (1929) transferred Oxytricha capitata to Amphisia too. However, the present species lacks zigzagging midventral pairs, so that a relationship to the urostyloids is very unlikely. Mansfeld (1923) described, like Gourret & Roeser (1888), seven transverse cirri. Moreover, he found that the present species has only two macronuclear nodules (Fig. 16c). For this and other reasons, Kahl (1932, 1933) doubted Mansfeld’s identification, that is, Kahl basically considered Mansfeld population as distinct species (see nomenclature). In this context it should be noted that Kahl’s (1932, 1933) redrawings of O. capitata and A. marioni are not exact, and in some respects almost misleading; for example, according to Pereyaslawzewa (1886) the right marginal row is displaced to near the amphisiellid median cirral row (Fig. 16a), whereas it is near the right body margin in Kahl’s redrawing (Fig. 16g). In this feature, O. capitata and A. marioni are also very similar (Fig. 16a, b). Wicklow (1982b) redescribed Amphisiella marioni after protargol impregnation, and largely confirmed the cirral pattern illustrated by Pereyaslawzewa (1886), Gourret & Roeser (1888), and Mansfeld (1923), and the two macronuclear nodules observed by Mansfeld. However, Wicklow’s population differs from the previous descriptions more or less distinctly in some other features, namely, in the body length (75–125 µm in life? vs. 150–200 µm according to Mansfeld), the characteristic cephalisation (lacking vs. present in all earlier descriptions), and the course of the right marginal row (straight vs. twisted). Possibly the last two features have been lost in Wicklow’s specimens due to the preparation procedure, but one can also not exclude that he observed a different species. Hemberger (1982) synonymised Oxytricha capitata, Amphisiella marioni, A. annulata, and A. milnei because they are similar in body shape and the cirral pattern. Beside O. capitata, I consider A. annulata and A. milnei as distinct species because they have, inter alia, distinct ring-shaped structures in the cytoplasm (see key).
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Because of all these problems (no detailed redescription, synonymy of O. capitata and A. marioni not quite certain) a detailed redescription from life and after protargol impregnation of such a population (better of more populations to show the constancy of the features) is recommended. Probably, only a neotypification of A. capitata or of A. capitata and A. marioni – when reliable differences can be demonstrated – will close the discussion. Small & Lynn (1985) incorrectly wrote that Fig. 16u is from Borror & Evans (1979). The illustration by Carey (1992; Fig. 16f) is very misleading; his contribution is therefore classified as insufficient redescription. Morphology: The description below is based mainly on the life data provided by Mansfeld (1923) and the cirral pattern as described by Wicklow (1982b), unless otherwise indicated. Please note that the synonymy of O. capitata and A. marioni is not definite. See Addenda for a redescription of the synonym A. marioni by Lin et al. (2007). Body size neither mentioned by Pereyaslawzewa (1886) nor by Gourret & Roeser (1888); according to Mansfeld (1923) 150–200 × 40–50 µm in life, according to Wicklow (1982) body length is 75–125 µm (method [from life? protargol impregnation?] not indicated). Body length:width ratio of extended specimens up to 8:1, usually, however, around 4–5:1. Body elongate, anterior quarter of cell usually set off from body proper, that is, cell distinctly cephalised; both cell ends rounded, left cell margin usually slightly more convex than right (Fig. 16c). Body distinctly flattened dorsoventrally, that is, ventral side slightly concave, dorsal side vaulted; anterior portion more distinctly flattened and therefore more translucent than main body portion. Body very flexible and contractile for about 50% of length. Two macronuclear nodules (Wicklow 1982b) left of cell midline behind proximal end of adoral zone (Mansfeld 1923, Fig. 16c). Contractile vacuole lacking (Mansfeld 1923); according to Gourret & Roeser (1888) also usually lacking; if present, however, then in the posterior dorsal portion. By contrast, Pereyaslawzewa (1886) illustrated a vacuole at anterior end of left marginal row (Fig. 16a; see also redrawings by Kahl 1932, 1933; Fig. 16g, h). Pellicle delicate, especially in anterior cell portion so that cephalised oral region frequently bursts under cover glass pressure. Cytoplasm with small, irregular granules; in main axis of oral region and in right body portion large, dark globules. Presence/absence of cortical granules not described for A. marioni and likely also not for O. capitata by Pereyaslawzewa (1886). Usually, Amphisiella capitata moves continuously forward, showing great flexibility and contractility when nestling against sediment particles; sometimes it moves backwards for about one body length. Occasionally it adheres to the substrate with the transverse cirri simultaneously twisting about main body axis. Swims under slow rotation about main body axis. Length of adoral zone rather varying, namely, 29% of body length (Fig. 16b), 33% (Fig. 16m), 44% (Fig. 16a), 47% (Fig. 16c). Zone shaped like a question mark (Mansfeld 1923; Fig. 16c), distal end extends far posteriorly to about 18% of body length (Fig. 16c; DE-value = 0.38); by contrast, in the synonym A. marioni the distal
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end is about at 11% of body length (Fig. 16b); in spite of this the DE-value is also rather high, namely 0.33. However, values must not be over-interpreted because based on live observations only. Distalmost 12 (Mansfeld 1923) to 13 (Wicklow 1982b) membranelles large and usually distinctly spread; proximal portion (= ventral) composed of about 22 membranelles (Wicklow 1982b). Buccal field narrow. Cilia of paroral shorten from anterior to posterior; both undulating membranes rather long and almost straight (Fig. 16c, m). Cytopharynx inconspicuous in life. Exact cirral pattern not known because identification by Wicklow (1982b) not quite certain. However, the available data indicate that the arrangement in A. capitata (Fig. 16a–c) is very similar to that of A. annulata, a species already described by modern methods (Fig. 17g, n). In total, seven isolated cirri on the frontal region, that is, three inconspicuously enlarged frontal cirri, one buccal cirrus near anterior end of paroral, one cirrus (= cirrus III/2) behind right frontal cirrus, and two cirri right of cirrus III/2 (Fig. 16m). Amphisiellid median cirral row composed of two portions (Fig. 16m); bipartition, however, very likely not clearly recognisable in life (Fig. 16a–c). Anterior portion composed of 4–7 cirri, rear portion of about 22 cirri (Wicklow 1982b). According to original descriptions and redescription by Mansfeld (1923), amphisiellid median cirral row extends sigmoidally to very near transverse cirri (Fig. 16a–c), according to Wicklow (1982b) it runs almost longitudinally and ends at about 76% of body length in specimen illustrated (Fig. 16m; see remarks). Seven pretransverse ventral and transverse cirri (Fig. 16b, c, m). Pereyaslawzewa (1886; Fig. 16a) likely did not clearly distinguish them from the left marginal cirri or overlooked the two inconspicuous pretransverse ventral cirri and therefore counted only five cirri. Pretransverse ventral cirri (= accessory transverse cirri according to Wicklow’s terminology) rather small, and – as is usual – ahead of the two rightmost transverse cirri. Transverse cirri slightly enlarged and almost terminal so that they project distinctly (slightly more than 50% according to Mansfeld 1923) beyond rear body end (Fig. 16c, m); sometimes only four transverse cirri present (Wicklow 1982b). Gourret & Roeser (1888) mentioned seven transverse cirri; likely they did not distinguish between pretransverse ventral cirri and transverse cirri. Right marginal row commences distinctly behind anterior end of cell, extends, like amphisiellid median cirral row, sigmoidally to near rear cell end, that is, central and posterior portion of row distinctly displaced inwards so that it runs roughly in parallel with amphisiellid median cirral row (Fig. 16a–c); right row composed of 34 cirri in specimen illustrated by Wicklow (1982b; Fig. 16m). Left row begins left of proximal portion of adoral zone, ends subterminally; composed of 29 cirri in specimen illustrated (Fig. 16m); rear end not distinctly separated from transverse cirri, which are, however, distinctly longer; left marginal cirri become only slightly longer posteriad (Mansfeld 1923). Marginal cirri and cirri of amphisiellid median cirral row beat continuously; frontal cirri move irregularly, whereas adoral membranelles show a permanent, wavelike movement (Mansfeld 1923). Dorsal cilia likely of ordinary length (3–4 µm) because neither mentioned nor illustrated in all papers; arranged in six kineties; caudal cirri definitely lacking (Wick-
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Fig. 16m–p Amphisiella capitata (from Wicklow 1982b. Protargol impregnation). m: Infraciliature of ventral side of non-dividing specimen, size not indicated. Pretransverse ventral cirri encircled. n–p: Infraciliature of ventral side of a very early, an early, and a middle divider. Sizes not indicated. Arrow in (n) marks the front anlagen patch. Detailed explanation see text. BC = buccal cirrus, FT = frontoterminal cirri forming anterior portion of amphisiellid median cirral row, OP = oral primordium, I–VI = frontal-ventraltransverse cirri anlagen. Page 91.
low 1982b). Mansfeld (1923) mentioned that he did not recognise dorsal bristles; however, he observed five (three central, two marginal) rows of dots, possibly cortical granules along dorsal kineties. Cell division (Fig. 16n–t): Ontogenetic data of the synonym A. marioni have been provided by Borror & Wicklow (1982) and Wicklow (1982a, b, 1983). Cell division begins with the proliferation of basal bodies left of each of the 10–12 rearmost cirri of the amphisiellid median cirral row. The basal bodies form a series of roundish primordia that enlarge into an extensive field (Fig. 16n, o). Six frontal-ventral-transverse primordia (I–VI) originate from the oral primordium in total (Fig. 16p). Meanwhile, the parental undulating membranes dedifferentiate and five anlagen obviously form from the buccal cirrus, the cirrus behind the right frontal cirrus, the rear cirrus of the cirral pair right of cirrus III/2, the anterior cirri of the posterior portion and the posterior cirri of the anterior portion of the amphisiellid median cirral row (Fig. 16o, p). In total, six primordia occur both in proter and opisthe. The
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Fig. 16q–w Amphisiella capitata (q–t, from Wicklow 1982b; u, v, from Borror 1979; w, after Kahl 1932 from Borror 1972. q–v, protargol impregnation; w, from life). q–t: Ventral side of middle to very late dividers, sizes not indicated. Broken lines in (t) connect cirri which originate from the same anlage. Pretransverse ventral cirri, respectively, transverse cirri of opisthe connected by dotted line. Arrow in (t) marks a cirrus whose origin is not comprehensible. u, v: Ventral side of a non-divider (103 µm) and anlagen area of opisthe. w: Non-divider. I–VI = cirri anlagen. Page 91.
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frontal cirri, the anterior cirrus of the cirral pair right of cirrus III/2, the anteriormost two cirri of the amphisiellid median cirral row, and the posteriormost cirri of this row are not modified to primordia (Fig. 16q). The anlagen I–IV form the following number of cirri: 1 (left frontal cirrus); 3 (middle frontal cirrus, buccal cirrus, leftmost transverse cirrus); 3 (right frontal cirrus, cirrus III/2, second transverse cirrus from left); 3 (cirral pair right of cirrus III/2, middle transverse cirrus). Anlage V produces the posterior portion of the amphisiellid median cirral row, the left pretransverse ventral cirrus, and the second transverse cirrus from right). Anlage VI produces, as is usual, (i) the frontoterminal cirri (five in specimen shown in Fig. 16t) which, on the other hand, form the anterior portion of the amphisiellid median cirral row; (ii) the right pretransverse ventral cirrus; and (iii) the rightmost transverse cirrus (Fig. 16r–t). The very late divider shown in Fig. 16t has a supernumerary cirrus (arrow) whose origin is incomprehensible. Possibly it is a special feature of this specimen or a parental, not yet resorbed cirrus. The formation of the marginal rows and dorsal kineties proceeds in ordinary manner; each two primordia occur within the parental cirral and bristles rows (Wicklow 1982b), that is, dorsomarginal rows and a dorsal kinety fragmentation are lacking. No data are available about the division of the nuclear apparatus. Very likely it proceeds as in A. annulata (see there), that is, the nodules fuse to a single mass during division and later divide again. Occurrence and ecology: Very likely confined to marine habitats. Type locality of A. capitata is the Black Sea near Sevastopol, Ukraine (Pereyaslawzewa 1886, Kahl 1932). Type locality of the junior synonym A. marioni is the harbour of Bastia (Corsica), Mediterranean Sea (Gourret & Roeser 1888). Mansfeld (1923) isolated the present species from a seawater tank (filled with material from the North Sea and the Mediterranean Sea) of the Aquarium of the city of Berlin, Germany. Borror (1979) found A. marioni in a sample (salinity 38‰) from the coast of the Yucatan peninsula, Mexico. Wicklow (1982b) found it in the same area; however, it is not known whether or not Borror (1979) and Wicklow (1982b) worked with the same population. Records of A. capitata not substantiated by morphological data: Mediterranean Sea near Marseille, France (Vacelet 1960, p. 54; 1961a, p. 15; 1961b, p. 4); at 21–27°C and about 13–18‰ salinity at Bulgarian coast of Black Sea (Detcheva 1977, p. 4; 1980, p. 35; 1982, p. 249; 1983, p. 72); sandy sediment of Odessa Bay and other sites of Black Sea (Dzhurtubayev 1978, p. 65; Kovaleva 1966, p. 1603); in detritus and sand (grain size 0.1–0.7 mm) of the Western Caspian Sea (Agamaliev 1971, p. 383; Agamaliyev 1974, p. 21); sand from an estuary near the Narragansett Marine Laboratory, University of Rhode Island, USA (Lackey 1961, p. 276). I somewhat doubt the record of A. capitata from saline soils of the Hortobágy National Park, Hungary reported by Szabó (1999, p. 249).
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Records of the synonym A. marioni not substantiated by morphological data: Mediterranean Sea near Banyuls-sur-mer and Marseille, France (Dragesco 1953, p. 630; Vacelet 1961b, p. 4); sediment of Loch Eil on the west coast of Scotland, Atlantic Ocean (Wyatt & Pearson 1982, p. 301); coastal areas of the Sea of Cantabria (Spain), Bay of Biscay, Atlantic Ocean (Fernandez-Leborans & Novillo 1993, p. 216); freshwater flats of “Fährmannssand” in the Elbe estuary (Pfannkuche et al. 1975, p. 482); Darss-Zingster lagoon near the city of Rostock, Baltic Sea (Schiewer 1994, p. 78; identified by E.M. Scharf); at 25°C and about 14‰ salinity at Bulgarian coast of Black Sea (Detcheva 1982, p. 249; 1983, p. 72); sediment off west coast of Caspian Sea (Agamaliev 1967, p. 369; 1970, p. 1279). Wicklow (1982b) cultured the populations in 38‰ seawater at 16°C and with the diatom Phaeodactylum as food. According to Wyatt & Pearson (1982), the junior synonym A. marioni is a grazer feeding upon detritus and bacteria. Mansfeld (1923) observed only sludge (sediment) particles in the newly forming food vacuoles.
Amphisiella annulata (Kahl, 1932) Borror, 1972 (Fig. 17a–z, 18a–q, Table 15) 1928 Holosticha annulata – Kahl, Arch. Hydrobiol., 19: 212, Fig. 44f (Fig. 17a; original description; no formal diagnosis provided and no type material available). 1932 Amphisiella (Holosticha) annulata Kahl, 1928 – Kahl, Tierwelt Dtl., 25: 590, Fig. 1121 (Fig. 17b; redescription and revision; see nomenclature for correct name). 1933 Amphisiella annulata Kahl 1928 – Kahl, Tierwelt N.- u. Ostsee, 23: 112, Fig. 17.21 (Fig. 17c; guide to marine ciliates; see nomenclature for correct name). 1963 Holosticha annulata Kahl, 1928 – Borror, Arch. Protistenk., 106: 511, Fig. 118 (Fig. 17d; redescription). 1972 Amphisiella annulata (Kahl, 1928) Kahl, 1932 – Borror, J. Protozool., 19: 9 (combination with Amphisiella, see nomenclature; revision of hypotrichs). 1985 Amphisiella annulata (Kahl, 1928) – Aladro Lubel, An. Inst. Biol. Univ. Méx., 55: 25, Lámina 12, Fig. 4 (Fig. 17e; illustrated record). 1990 Amphisiella annulata (Kahl, 1928) – Aladro Lubel, Martínez Murillo & Mayén Estrada, Manual de ciliados, p. 125, Figure on p. 125 (Fig. 17f; review). 1992 Amphisiella annulata (Kahl, 1928) Kahl, 1930–5 – Carey, Marine interstitial ciliates, p. 179, Fig. 701 (guide; illustration is a redrawing of Fig. 17a). 1999 Amphisiella annulata (Kahl, 1928) – Alekperov & Asadullayeva, Turkish J. Zool., 23: 219, Fig. 8 (Fig. 17w; description of a Caspian Sea population). 2001 Amphisiella annulata (Kahl, 1928) Borror, 1972 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 33 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2003 Amphisiella annulata Kahl, 1928 – Hu, Gong & Song, Pathogenic Protozoa, p. 162, Fig. 5-9A–C (Fig. 17x–z; description of Chinese population; see nomenclature). 2004 Amphisiella annulata (Kahl, 1932) Borror, 19721 – Berger, Acta Protozool., 43: 2, Fig. 1–23 (Fig. 17g–v; detailed redescription; five neotype slides [accession numbers 2003/146–150] have been deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria). 1
Berger (2004) provided the following improved diagnosis solely based on the neotype population: Body size about 130 × 33 µm in life. Body outline elongate elliptical to oval. Two macronuclear nodules. Cortical granules colourless, arranged mainly along dorsal kineties. Amphisiellid median cirral row extends sigmoidally from near right frontal cirrus to near transverse cirri, consists of about 44 narrowly spaced
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Fig. 17a–f Amphisiella annulata (a, from Kahl 1928b; b, from Kahl 1932; c, from Kahl 1933; d, from Borror 1963; e, from Aladro Lubel 1985; f, from Aladro Lubel et al. 1990. a–f, from life). Ventral views showing, inter alia, basic cirral pattern, nuclear apparatus, and ring-shaped structures (a = 120–150 µm, b, c = 150–200 µm, d = 160 µm, e, f = 125 µm). Note that the cirral pattern illustrated by Kahl (1932; Fig. 17b) matches the pattern of the neotype almost perfectly (Fig. 17n). Borror illustrated a large vacuole in the posterior body portion near the right body margin (d); it must not be misinterpreted as contractile vacuole because Borror did not mention such an organelle; interestingly enough, in (e, f) a somewhat smaller vacuole (inclusion?) is shown in a very similar position. Page 100.
2004 Amphisiella annulata (Kahl, 1928) Borror, 1972 – Hu, Warren & Suzuki, Acta Protozool., 43: 363, Fig. 4A–J, 5A–F, 8A–J, Tables 3, 4 (Fig. 18a–q; redescription of two allochronic Yellow Sea populations; site where voucher slides deposited not mentioned).
Nomenclature: No derivation of the name is given in the original description. According to Berger (2004), the species-group name annulát·us, -a, -um (Latin adjective [m, f, n]; ringed, having a small ring [ánnulus]; Hentschel & Wagner 1996) obviously alludes to the ring-shaped structures in the cytoplasm. Kahl (1932, 1933) classified Amphisiella as subgenus of Holosticha. Thus, the correct name in his reviews is Holosticha (Amphisiella) annulata Kahl, 1928. This was obviously overlooked by Borror (1972) and Carey (1992), who assumed that Kahl (1932) had transferred it from the genus Holosticha to the genus Amphisiella (see list of synocirri which are conspicuously wide (4–5 µm!) in middle portion of row. On average 47 adoral membranelles and each 34 cirri in left and right marginal row. More or less invariably 1 buccal cirrus, 1 cirrus behind right frontal cirrus, 3 cirri left of anterior portion of median cirral row, 2 pretransverse ventral cirri, 6 transverse cirri, and 6–7 dorsal kineties. Oral primordium originates from several anlagen pits. Fourth transverse cirrus from left is formed from additional anlage which produces no other cirri.
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Fig. 17g–m Amphisiella annulata (neotype population from Berger 2004. g–l, from life; m, protargol impregnation). g: Ventral view of a representative specimen, 150 µm. Note ring-shaped structures (“hollow” globules?). h, i: Body outline of posteriorly widened specimens in ventral and dorsal view showing, inter alia, marginal cirral rows, amphisiellid median cirral row, the frontal scutum, and a dorsal furrow. j, k: Ring-shaped structures (“hollow” globules?) about 4–8 µm across (fine structure not completely discerned). l: Two size classes of cortical granules are present: (i) 0.8–1.0 µm-sized, colourless globules, which form patches between dorsal bristles; (ii) tiny (about 0.3–0.5 µm across) colourless globules scattered throughout cortex. m: Infraciliature of oral region of neotype specimen (complete cirral pattern, see Fig. 17n). Arrow marks cirrus III/2. Broken lines connect cirri which originate from same anlage (only
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nyms). For the sake of simplicity I suggest fixing Borror (1972) as combining author, although he did not formally transfer it to Amphisiella. Hu et al. (2003) erroneously assumed that Kahl (1928b) had established this species in the genus Amphisiella. Incorrect subsequent spelling: Amphisiella anulata (Fernandez-Leborans & Novillo 1993, p. 216). Remarks: The classification of the present species in the subgenus/genus Amphisiella by Kahl (1932), respectively, Borror (1972) seems correct because the cirral pattern matches that of A. capitata, type of the genus, very well. Further, the cell division in these two species proceeds largely identically. The original description of A. annulata is very brief, but contains most diagnostic features (Kahl 1928b). Accordingly, the present species has five frontal and transverse cirri each (Fig. 17a). As concerns the frontal cirri, Kahl (1928b) very likely recognised only the three enlarged frontal cirri, the buccal cirrus, and cirrus III/2. The short row composed of three cirri left of the anterior portion of the median cirral row is less distinct than the other five “frontal cirri” and thus easily overlooked or misinterpreted as anterior end of the median cirral row. The interpretation of the five transverse cirri is somewhat more difficult because this number was not mentioned by Kahl (1932) and Alekperov & Asadullayeva (1999) and occurred only very rarely in the Adriatic neotype population (Table 15). The following possibilities exist: (i) Kahl (1928b) observed and illustrated a rare specimen, which indeed had only five transverse cirri; (ii) Kahl (1928b) did not count correctly. On page 211, Kahl (1928b) wrote that he would like to investigate the species again, indicating that his observations are not very detailed and precise; (iii) the populations studied by Kahl (1928b) and Kahl (1932) are not conspecific. I prefer possibilities (i) and (ii) because it is unlikely that Kahl (1932) did not recognise his own species, inasmuch as it has a conspicuous feature, namely the large, ring-shaped structures. To avoid this permanent uncertainty I designated a neotype (Berger 2004). Kahl (1932, p. 582, 583) wrote that he had confused A. annulata with his Caudiholosticha setifera (Kahl, 1932) Berger, 2003 (basionym: Holosticha setifera) earlier, likely because both species have large, ring-shaped structures (for review see Berger 2006, p. 281). However, Caudiholosticha setifera, which occurred – like A. annulata – in saltwater habitats near Bad Oldesloe (Germany), has a distinct midventral pattern, a single micronucleus between the two macronuclear nodules, and distinct caudal cirri and therefore differs clearly from A. annulata. Holosticha obliqua Kahl, 1928 sensu Kahl (1932, his Fig. 10626) also has only one micronucleus between the two macronuclear nodules and five transverse cirri (vs. several micronuclei and six transverse cirri in A. annulata). Possibly, this illustration contains fea-
b
shown for anlagen I–IV). ACR = amphisiellid median cirral row, AZM = distal end of adoral zone of membranelles, BC = buccal cirrus, CG = cortical granules, DB = dorsal bristle, FC = right frontal cirrus, FS = frontal scutum, LMR = left marginal row, P = paroral, RMR = right marginal row, IV = cirri formed by anlage IV, 5, 6 = rightmost dorsal kineties. Page 100.
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Fig. 17n–p Amphisiella annulata (neotype population from Berger 2004. Protargol impregnation). n, o: Infraciliature of ventral and dorsal side and nuclear apparatus of neotype specimen, 102 µm (detail of anterior portion, see Fig. 17m). Note that the middle portion of the median cirral row is composed of rather wide, closely spaced cirri (large arrow in n). Small arrow in (n) denotes the “additional” transverse cirrus IVa/1 (see Fig. 17s, v). Pretransverse ventral cirri encircled. p: Infraciliature of left side, 137 µm. Note strong vaulting of dorsal side, that is, specimens almost not flattened dorsoventrally. LMR = left marginal row, MA = macronuclear nodule, MI = micronucleus, TC = leftmost transverse cirrus, 1–6 = dorsal kineties. Page 100.
tures of C. setifera and A. annulata, as supposed by Kahl (1932) himself. Holosticha obliqua is classified as species indeterminata by Berger (2006, p. 183). According to Kahl (1932), the frontal cirri of A. annulata are arranged “as in Oxytricha (3 + 2 + 3)”. This is only correct for the first two parts (3 + 2), that is, the three enlarged frontal cirri (cirri I/1, II/3, III/3), the buccal cirrus (cirrus II/2), and cirrus III/2, which is behind the right frontal cirrus. The last three cirri (... + 3) are formed by the anlage IV in A. annulata, whereas they are produced from two different anlagen in the 18-cirri hypotrichs/oxytrichids, namely the anterior two cirri (cirri
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Fig. 17q, r Amphisiella annulata (neotype population; from Berger 2004. Protargol impregnation. Parental structures white, new black). q: Infraciliature of ventral side and nuclear apparatus of early divider, 121 µm. Oblique arrow marks an anlagen pit. Transverse arrow marks discontinuity in amphisiellid median cirral row (site where the two portions of the row join). Pretransverse ventral cirri circled. r: Infraciliature of ventral side and nuclear apparatus of an early to middle reorganiser. Arrow marks an anlagen pit. Very likely this is a reorganiser because in dividers of such a stage the anlagen for both the porter and the opisthe would be recognisable. Note that the additional anlage IVa is not yet clearly recognisable. Area ahead of transverse cirri not recognisable because covered by debris. MA = macronuclear nodule, MI = micronucleus, I–VI = frontal-ventral-transverse cirri anlagen, 1 = dorsal kinety 1 (= leftmost kinety). Page 100.
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Fig. 17s–u Amphisiella annulata (neotype population from Berger 2004. Protargol impregnation. Parental structures white, new black). s, t: Infraciliature of ventral and dorsal side and nuclear apparatus of late divider, 113 µm. Arrows mark anlagen IVa, which form the additional transverse cirrus (= cirrus IVa/1) marked by an arrow in Fig. 17n. Note that no dorsomarginal kineties are formed in the amphisiellids. u: Nuclear apparatus of a late reorganiser shown in Fig. 17v, 75 µm. LMR = left marginal row, MA = macronucleus, MI = dividing and non-dividing micronucleus, RMR = right marginal row, I, IV, VI = frontal-ventral-transverse cirri anlagen, 1, 7 = dorsal kineties. Page 100.
VI/3, VI/4 = frontoterminal cirri) from anlage VI and the rearmost cirrus from anlage IV (cirrus IV/3; for review on oxytrichids, see Berger 1999). The redescriptions by Borror (1963, Fig. 17d) and Aladro Lubel (1985, Fig. 17e) are less detailed than Kahl’s (1932, Fig. 17b) data. Especially the cirral patterns do not correspond very well with those described by Kahl (1932) and Berger (2004; Fig. 17n). The following differences are the most conspicuous ones: (i) the median
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cirral row is distinctly shortened against unshortened; (ii) only five transverse cirri present (Fig. 17d–f) against six (Fig. 17b, n). Admittedly, five matches the original description by Kahl (1928b) exactly, but, as discussed above, this is obviously not the ordinary number for A. annulata. (iii) Borror and Aladro Lubel did not mention the conspicuous ring-shaped structures, although some globules, which can be also interpreted as rings, are illustrated (Fig. 17d–f). They write about digestive vacuoles and cytoplasmic granules (Aladro Lubel 1985) and many endoplasmic spherical granules 4 µm in diameter (Borror 1963). The non-mention of these large, conspicuous ring-shaped structures indicates that they were not present in the American populations. However, this is not a proof for a misidentification because Kahl (1932) also mentioned that the rings, which could be lithosomes (for review, see Hausmann & Hülsmann 1996), can be absent. It is noticeable that the illustrations provided by Borror (1963; Fig. 17d) on the one hand and by Aladro Lubel (1985, Fig. 17e) and Aladro Lubel et al. (1990, Fig. 17f) on the other are very similar, especially as concerns the anterior shortage of the median cirral row, the five strong transverse cirri, and the six paired cirri ahead of the median cirral row. The Caspian Sea population (Fig. 17w) differs from the Adriatic neotype population mainly by the lack of the ring-shaped structures (vs. present), in the higher number of adoral membranelles (65–70 vs. 31–57), the lack of pretransverse ventral cirri (vs. two such cirri present), and the lower number of dorsal kineties (four vs. usually six). The data by Berger (2004) match the redescription and illustrations by Kahl (1932, 1933) almost perfectly so that the identification is beyond reasonable doubt. Very likely all identifications listed in the synonymy above and the occurrence and ecology section were done after Kahl (1932), and not after the original description (Kahl 1928b). Consequently, the 1932-review was de facto the authoritative redescription. There are only few differences between Kahl’s (1932) data and my observations, which, however, can be explained: (i) body length is 150–200 µm against 100–160 µm. My sole live measurement, which I made some days before the fixation of the material, was 160 × 40 µm, which fits into the range provided by Kahl. Obviously the Adriatic specimens became slightly smaller in cultures as indicated by the morphometric analysis (Table 15). Further, the specimens of Kahl’s (1928b) population were also only 120–150 µm long. (ii) Kahl did not mention the cortical granules, which were present in the population from the Adriatic Sea. However, the granules are colourless and rather small (0.3–1.0 µm) and thus easily overlooked especially with bright field illumination. Kahl (1932) wrote “ ..., usually grey granulated and in between with numerous ring-shaped reserve bodies”; this indicates that the term grey granulated refers to the cytoplasm and not to a cortical granulation. Indeed, the specimens of my population were also steel-grey; however, I did not know whether this impression was mainly due to the cytoplasm or the colourless cortical granulation. (iii) The Adriatic population has two pretransverse ventral cirri, which were neither mentioned nor illustrated by Kahl (1932, 1933). However, they are rather small and at least the left cirrus is almost not set off from the rear end of the
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Fig. 17v Amphisiella annulata (neotype population from Berger 2004. Protargol impregnation. Parental structures white, new black). Infraciliature of ventral side of a late reorganiser, 75 µm (nuclear apparatus shown in Fig. 17u). Cirri originating from same anlage are connected by broken line. Arrow marks additional transverse cirrus originating from anlage IVa, which is formed between anlagen IV and V (see Fig. 17s). AZM = adoral zone of membranelles, LMR = left marginal row, RMR = right marginal row, I–VI = frontal-ventral-transverse cirri anlagen, 1 = dorsal kinety 1 (= leftmost kinety). Page 100.
median cirral row, indicating that Kahl had not distinguished this small cirral group from the median cirral row. But more important than these few differences is the exact agreement in the remaining cirral pattern, which is not very difficult to recognise in life. Kahl (1932) emphasised the short, wide, and narrowly spaced cirri in the marginal and median rows, a feature which is indeed very conspicuous (Fig. 17b, n). Because of the taxonomic problems (e.g., synonymy of A. capitata and A. annulata suggested by Hemberger 1982, see below), Berger (2004, p. 13) neotypified the present species. Hu et al. (2004) described two allochronic populations from a mollusc-culturing water off the coast of Qingdao, China. The Chinese populations agree with the neotype population rather well so that conspecificity is beyond reasonable doubt. The lack of ringshaped structures in the Chinese populations must not be over-interpreted because Kahl (1932) also mentioned that these structures are sometimes lacking. The paper by Hu et al. (2003) is in Chinese and thus only the illustrations are provided (Fig. 17x–y). According to Hemberger (1982), Amphisiella marioni, A. annulata, and A. milnei are junior synonyms of A. capitata. Indeed, synonymy of A. capitata and A. marioni is very likely and therefore also assumed in the present review. However, Amphisiella capitata, which is redescribed by Wicklow (1982b), has (i) only two cirri in the short row left of the anterior portion of the median cirral row (Fig. 17m; vs. three in A. annulata, Fig. 17b, m); (ii) a shorter and distinctly interrupted median cirral row (terminates distinctly ahead of transverse cirri vs. very near transverse cirri and usually continuous), which is composed of about 26–29 ordinary sized and spaced cirri (vs. 25–54, on average 44 rather wide cirri, which are narrowly
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spaced especially in the middle portion); (iii) 4–5 transverse cirri (vs. usually six). Amphisiella capitata and A. annulata agree, inter alia, in the marine habitat, the number of dorsal kineties (six, respectively, 6.4 on average), the arrangement of the undulating membranes (straight and in parallel), and the presence of invariably two pretransverse ventral cirri (termed accessory transverse cirri by Wicklow 1982b) ahead of the two rightmost transverse cirri. Further, morphogenesis obviously proceeds very similarly (see above) so that a close relationship of these two species is very likely. Amphisiella milnei (Fig. 19a–f) has, inter alia, yellowish cortical granules (vs. colourless in A. annulata), five transverse cirri (vs. six), possibly a somewhat different cirral pattern in the frontal area (cp. Fig. 19a with Fig. 17b), and one ring-shaped structure each in the anterior and posterior body portion (rear one sometimes lacking vs. usually several rings scattered throughout cytoplasm). Further, the cirri of the marginal rows and the median row are of ordinary size and arrangement in A. milnei, whereas they are rather wide and narrowly spaced in A. annulata. Amphisiella turanica from the Caspian Sea is somewhat larger (170–210 µm long in life) than A. annulata, has 70–85 adoral membranelles, four dorsal kineties, and, most importantly, four macronuclear nodules (Fig. 20a, b; Alekperov & Asadullayeva 1999). In life, Amphisiella annulata is rather conspicuous mainly because of the wide and narrowly spaced cirri of the median cirral row. This feature and the conspicuous ring-shaped structures enable a very simple identification of this salt water species. Since the cell is rather large and resistant to cover glass pressure, the cirral pattern can be studied in great detail even in life (own observations), and as impressively demonstrated by Kahl more than 70 years ago (Fig. 17b)! Morphology (Fig. 17a–p, w–z, 18a–j, Table 15): the improved diagnosis by Berger (2004) is solely based on data from the Adriatic neotype population. However, it covers Kahl’s and other relevant data rather well. The ring-shaped structures (lithosomes?), although very conspicuous, have been omitted from the diagnosis because they can be absent (Kahl 1932; see below). The neotype population is described first. Data from populations studied by Kahl (1928b, 1932, 1933) and the other workers (Borror 1963, Aladro Lubel 1985, Aladro Lubel et al. 1990, Alekperov & Asadullayeva 1999, Hu et al. 2004) are kept separate. Description of neotype population (Fig. 17g–p, Table 15): size in life about 100–160 × 30–40 µm (I made only one live measurement, namely 160 × 40 µm; the range is derived from the morphometric data shown in Table 15 assuming a shrinkage of up to 30% due to the preparation procedure; Berger et al. 1983); body length:width ratio of live specimens ranging from 3–4:1; prepared specimens only 68–121 µm long, length:width ratio on average 2.6:1 (Table 15). Body outline elongate elliptical (Fig. 17g) to slightly oval (Fig. 17h, i). Body very flexible and often slightly twisted about main body axis, not distinctly contractile, rather resistant to cover glass pressure; ventral side flat, dorsal side often distinctly vaulted so that many specimens are arranged with dorsal or lateral surface above in protargol prepa-
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rations (Fig. 17p). Invariably two macronuclear nodules slightly left of midline; individual nodules ellipsoidal, in life up to about 28 × 12 µm, with many chromatin aggregates of ordinary size; nodules usually connected by fine strand; length:width ratio of anterior nodule ranging from 1.8–4.8:1 (average 2.1:1), posterior nodule 1.4–3.4:1 (average 2.4:1; Table 15). Micronuclei ellipsoidal, arranged close to macronuclear nodules. No contractile vacuole recognisable. Two size-classes of colourless cortical granules: (i) larger globules about 0.8 to Fig. 17w–z Amphisiella annulata (w, from Alekperov & Asa1.0 µm across, form distinct dullayeva 1999; x–z, from Hu et al. 2003. w, wet silver nitrate patches mainly between indiimpregnation; x, from life; y, z, protargol impregnation). w, y, z: vidual bristles of a dorsal kiInfraciliature of ventral and dorsal side and nuclear apparatus, w = 110–145 µm, y, z = 152 µm. x: Ventral view, 141 µm. Page nety; (ii) smaller granules 100. about 0.3–0.5 µm across, more or less densely distributed in whole cortex (Fig. 17g, l); stainability with methyl-green pyronin not checked; sometimes cortical granules impregnate with protargol. Cells appear steel-grey; cytoplasm colourless, contains many greasily shining globules 2–5 µm across and several (up to about 20) ring-shaped structures (“hollow” spheres; lithosomes?) about 4–8 µm in diameter (Fig. 17g, j, k); rings not clearly recognisable in protargol preparations. Movement rapidly gliding, showing great flexibility. Adoral zone occupies 42% of body length, composed of 47 membranelles on average, individual membranelles of ordinary fine structure (Fig. 17g, h, m, n, p). Distal end of adoral zone extends, on average, to 12% of body length on right body margin, proximal portion usually widened slightly spoon-like. Buccal field inconspicuous, that is, narrow to ordinary width. Buccal lip also inconspicuous because does not cover proximal portion of adoral zone distinctly. Undulating membranes more or less straight and in parallel; fine structure of membranes not clearly recognised. Cytopharynx inconspicuous in life and protargol preparations. Cirral pattern and number of cirri of usual variability (Fig. 17g, m, n, p, Table 15). Three distinctly enlarged frontal cirri in ordinary arrangement, that is, in oblique pseudorow along anterior body margin with right cirrus (= cirrus III/3) close
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Fig. 18a–g Amphisiella annulata (from Hu et al. 2004. a–d, from life; e–g, protargol impregnation). a: Ventral view, size not indicated. b: Left lateral view. c: Ventral view of contracted specimen, 105 µm. d: Colourless cortical granules (0.5 µm across) around dorsal bristles and ejected granules (2–3 µm long). e: Infraciliature of ventral side and nuclear apparatus of a specimen from the year 2000 population, size not indicated. f, g: Infraciliature of anterior and posterior portion of the 1996 population. Arrow in (f) marks cirrus III/2; broken lines connect left frontal cirrus with undulating membranes and middle frontal cirrus and buccal cirrus. Cirral row formed by anlage IV encircled by dotted line. Arrows in (g) mark pretransverse ventral cirri. ACR = amphisiellid median cirral row, FC = right frontal cirrus, P = paroral, RMR = right marginal row. Page 100.
behind distal end of adoral zone. Invariably one slightly enlarged buccal cirrus (= cirrus II/2) closely ahead of anterior end of paroral, and one cirrus (= cirrus III/2; Fig. 17m) left behind right frontal cirrus. A short row composed of three, occasionally of four cirri left of anterior portion of amphisiellid median cirral row; very rarely a second row present. Amphisiellid median cirral row commences right of right frontal cirrus, extends slightly to distinctly sigmoidally close to transverse cirri, composed of 44 cirri on average (Fig. 17g, m, n, Table 15); median cirral row usually without distinct break although composed of two portions (see morphogenesis);
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cirri narrowly spaced, especially behind buccal vertex, where they are also rather wide, that is, up to 4–5 µm! Invariably two pretransverse ventral cirri, one ahead of right transverse cirrus, the other roughly in line with amphisiellid median cirral row. Usually six, rarely (1 out of 29 specimens) only five transverse cirri arranged in Jshaped, slightly subterminal pseudorow; cirri about 20 µm long in life and thus distinctly (right) to almost not (left) projecting beyond rear body end. Marginal cirri and cirri of amphisiellid median row only 8–10 µm long in life; marginal cirri relatively narrowly spaced. Right marginal row begins on average at 25% of body length, often more or less distinctly sigmoidal, terminates near right transverse cirrus; usually two ciliated basal body pairs ahead of right marginal row. Left marginal row commences distinctly ahead of level of buccal vertex, ends subterminally. Dorsal cilia 2–3 µm long; about two thirds of specimens with six and one third with seven, roughly bipolar kineties (Fig. 17g, m–p); rarely (1 out of 23 specimens analysed morphometrically) eight kineties present. Caudal cirri lacking. Besides the very brief original description by Kahl (1928b, Fig. 17a), several redescriptions are available, namely by Kahl (1932, 1933; Fig. 17b, c), Borror (1963, Fig. 17d), Aladro Lubel (1985; including a review by Aladro Lubel et al. 1990; Fig. 17e, f), Alekperov & Asadullayeva (1999; Fig. 17w), and Hu et al. (2003, Fig. 17x–z; 2004, Fig. 18a–j). To complete the picture of A. annulata, I provide the original data of these populations separately; see also corresponding illustrations for some features, for example, body outline, cirral pattern. The review by Carey (1992) is not considered further because it contains only a redrawing of Fig. 17a and lacks original data. Population described by Kahl (1928b; Fig. 17a): body length 120–150 µm in life; five frontal cirri and five long transverse cirri; single ventral row; scattered ringshaped “reserve bodies”; lively, metabolic, slightly contractile; two elongated macronuclear nodules. Population described by Kahl (1932, 1933; Fig. 17b, c): body length 150 to 200 µm in life; body slenderly ellipsoidal, soft, flexible, metabolic, slightly contractile, usually greyish granulated, with a variable number (zero to numerous, but never two, that is, never one each in anterior and posterior body portion as in A. milnei) of ring-shaped (probably “hollow-spherical”) reserve bodies; ectoplasm colourless; two elongated macronuclear nodules, each with a single (?) micronucleus; 6–7 transverse cirri (specimen illustrated has six cirri), which project slightly beyond rear body end; frontal cirri likely as in Oxytricha (3 + 2 + 3; homologisation not quite correct, see remarks); cirri of marginal rows and ventral row very short, wide, and narrowly spaced; for cirral pattern, see the excellent illustration (Fig. 17b); posterior portion of adoral zone of membranelles longitudinally arranged. Population described by Borror (1963; Fig. 17d): body size 160 × 30–36 µm in life; body evenly rounded at ends, slightly flat ventrally. Two oval macronuclear nodules, each 18 µm long; cytoplasm with many spherical granules 4 µm across. Adoral zone of membranelles 55 µm long, composed of 60–65 membranelles. Nine frontal cirri, the anterior three larger; ventral row composed of about 40 narrowly
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spaced cirri; five transverse cirri 24 µm long, extending just beyond rear body end; about 50 left and 60 right marginal cirri, very closely set; six dorsal kineties. Population described by Aladro Lubel (1985; Fig. 17e): body size 123 × 21 µm; body outline elongated; two oval-shaped macronuclear nodules, each with a spherical micronucleus; contractile vacuole in posterior body third (side not mentioned and vacuole likely not illustrated); several food vacuoles and cytoplasmic granules; adoral zone about 1/3 of body length; cirri narrowly spaced in all rows; nine frontal cirri, three anteriormost larger; five transverse cirri, which slightly protrude beyond rear body end. Population described by Alekperov & Asadullayeva (1999; Fig. 17w): body length 130–180 µm in life, 110–145 µm in silver preparations. Body elongated, strongly flattened dorsoventrally. Two macronuclear nodules, two micronuclei. Contractile vacuole near proximal end of adoral zone. Cytoplasm transparent, without inclusions. Adoral zone composed of 65–70 membranelles. Two frontal cirri near anterior end, five further frontal cirri at level of anterior portion of undulating membranes. Amphisiellid median cirral row composed of 48 cirri, extends from rear frontal cirri to near the six transverse cirri. 55 right and 40 left marginal cirri. Four dorsal kineties (three bipolar, one shortened; number of kineties possibly not correctly estimated due to inappropriate preparation method). Caudal cirri absent. Populations described by Hu et al. (2004): body size 100–210 × 24–75 µm in life; body length:width ratio about 3–5:1. Body outline usually elongate elliptical with both ends rounded, left margin conspicuously convex in mid-portion, right distinctly sigmoidal. Body dorsoventrally flattened, ventral side flat, dorsal distinctly vaulted in mid-portion (Fig. 18a–c); flexible and slightly contractile. Macronuclear nodules in left body portion behind proximal end of adoral zone. Micronuclei about 4 µm across; usually one or two near macronuclear nodules. Pellicle soft. Cortical granules colourless, 0.5 µm across, arranged around bases of dorsal bristles and in lines along dorsal kineties; ejected granules 2–3 µm long (Fig. 18d; some minor differences in the cortical granulation between the neotype population and the Yellow Sea populations should not be over-interpreted). Body colourless to greyish at low magnification, cytoplasm transparent with several refractive granular inclusions 2–5 µm long. Contractile vacuole not observed, probably absent. Some specimens with 1–2 food vacuoles in mid-body. Movement without peculiarities, that is, crawls on substrate or rotates around main body axis when swimming. Adoral zone occupies 25–33% of body length, distal end extends far posteriorly (DE-value 0.49 for specimen shown in Fig. 18f). Cilia of distal adoral membranelles and frontal cirri about 15 µm long. Transverse cirri about 16–20 µm long, remaining cirri about 10–12 µm long. Cirral pattern as shown in Fig. 18a, e–h, j, that is, basically as in the neotype population. Thus, only additional or deviating observations are provided. Paroral composed of zigzagging basal bodies arranged in parallel to single-rowed endoral; paroral commences more anteriorly than endoral. Basis of each marginal cirrus (and amphisiellid median row cirrus?) composed of two rows of basal bodies.
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Fig. 18h–k Amphisiella annulata (from Hu et al. 2004. Protargol impregnation). h–j: Infraciliature of ventral side and dorsal side and nuclear apparatus of non-dividing specimens, h, i = 196 µm, j = 150 µm. Broken lines connect cirri, which originate from the anlagen I–IV. k: Infraciliature of an early to middle divider, size not indicated. Arrows mark anlagen I of proter and opisthe. FC = right frontal cirrus. Page 100.
Dorsal bristles about 5 µm long and therefore easily recognisable in life; some dorsal kineties shortened at both ends (Fig. 18d, i). Cell division and reorganisation: Only few dividers and reorganisers have been found by Berger (2004). In spite of this, some highly interesting morphogenetic features could be elucidated (Fig. 17q–v). Some early and middle stages of division are lacking so that the origin of the frontal-ventral-transverse cirri anlagen remains basically unknown. The data provided by Hu et al. (2004; Fig. 18k–q) confirm my observations. Cell division commences with the formation of some anlagen pits left of the third quarter of the median cirral row (Fig. 17q). The pits are roundish and up to 5 µm
Fig. 18l–q Amphisiella annulata (from Hu et al. 2004. Protargol impregnation). l, m: Infraciliature of ventral and dorsal side and nuclear apparatus of a middle divider, 145 µm. Arrow in (l) denotes the anlage IVa which produces the additional transverse cirrus (see Fig. 17s). n–q: Infraciliature of ventral and dorsal side and nuclear apparatus of a middle and a late reorganiser, n, o = 193 µm, p, q = size not indicated. Page 100.
d
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deep. Very likely the cirri of the median cirral row beside the pits are not altered at this stage. Ahead of the five pits of the divider shown is a large field of basal bodies, which likely originated by the fusion of some further pits. Fig. 17r shows an early to middle reorganiser in which the anlagen I–VI are clearly recognisable. The additional anlage IVa, which produces the fourth transverse cirrus from left, is not yet clearly recognisable. A middle to late divider shows the differentiation of the frontal-ventraltransverse cirri, which originate from six anlagen (I–VI; Fig. 17s). A curiosity is the formation of a small basal body field between anlagen IV and V (Fig. 17s, arrows). It only forms a transverse cirrus, namely the fourth from left. Consequently, the cirral anlagen produce the following structures: anlage I forms the undulating membranes (paroral, endoral) and the left frontal cirrus (= cirrus I/1); anlage II forms the leftmost transverse cirrus (= cirrus II/1), the buccal cirrus (= cirrus II/2), and the middle frontal cirrus (= cirrus II/3); anlage III produces the second transverse cirrus from left (= cirrus III/1), the cirrus left behind right frontal cirrus (= cirrus III/2), and the right frontal cirrus (= cirrus III/3); anlage IV produces the third transverse cirrus from left (= cirrus IV/1) and the three cirri left of the anterior portion of the amphisiellid median cirral row; anlage IVa (Fig. 17s, arrow; see below for explanation) forms only the fourth transverse cirrus from left (= cirrus IVa/1; Fig. 17n, arrow; Fig. 17v, arrow); anlage V produces the fifth transverse cirrus from left (= cirrus V/1), the left pretransverse ventral cirrus (= cirrus V/2), and the posterior portion (around 31 cirri) of the amphisiellid median cirral row; anlage VI forms the sixth transverse cirrus from left (= cirrus VI/1), the right pretransverse ventral cirrus (= cirrus VI/2), and the anterior portion (about 13 cirri) of the amphisiellid median cirral row. The parental adoral zone of membranelles obviously remains more or less unchanged, whereas the parental undulating membranes are reorganised (Fig. 17s). A late reorganiser shows that the amphisiellid median cirral row of A. annulata is formed by cirri of the two rightmost anlagen (Fig. 17v). Anlage V produces the posterior portion, which commences slightly ahead of the level of the buccal vertex in interphasic specimens. The front portion is formed by the anteriorly migrating cirri (except the right pretransverse ventral cirrus and the right transverse cirrus) of anlage VI. In interphasic specimens the two portions form a continuous row (Fig. 17n); a discontinuity is only very rarely recognisable (Fig. 17q, transverse arrow). The development of the marginal rows and dorsal kineties does not show any peculiarities. Two primordia each occur within the parental cirral and bristle rows. No caudal cirri are formed (Fig. 17s, t). The division of the nuclear apparatus proceeds in ordinary manner, that is, the two macronuclear nodules fuse to a single mass, which subsequently divides. The micronuclei behave like those of other hypotrichs (Fig. 17q, t, u). The morphogenetic pattern of A. annulata largely agrees with that of A. capitata (Fig. 16m–t; Wicklow 1982b). Of course there are also several agreements with many other hypotrichs, for example, that the left frontal cirrus originates from the undulating membrane anlage or that the buccal cirrus is the middle cirrus of anlage
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II. However, these features are symplesiomorphies and are therefore not considered further in the discussion below. A very interesting morphogenetic feature of A. annulata is the formation of roundish anlagen pits left of the middle and posterior portion of the median cirral row (Fig. 17q). Wicklow (1982b) illustrated an early divider of A. capitata, which shows several roundish patches of basal bodies, strongly resembling the pits of A. annulata (Fig. 16n). However, Wicklow mentioned no details about this stage so that it is unknown whether these patches are invaginated or flat. Thus, reinvestigation of the morphogenesis of the type species is needed. In most hypotrichs the oral anlage develops on the cell surface (for review, see Foissner 1996). In euplotids, strombidiids, strobilidiids, and few non-euplotid hypotrichs, e.g., Psilotricha succisa (Foissner 1983), the anlage originates in a more or less distinct subsurface pouch (= hypoapokinetal stomatogenesis). In Pseudoamphisiella lacazei morphogenesis commences with the proliferation of loosely arranged basal bodies below the cortex and therefore parental cirri and fibres remain intact (Song et al. 1997; for review, see Berger 2006). According to molecular data, the oligotrichs and the euplotids are the nearest relatives of the remaining hypotrichs (e.g., Bernhard et al. 2001, Modeo et al. 2003). This indicates that the hypoapokinetal stomatogenesis is a plesiomorphy for the hypotrichs. Unfortunately, we do not know whether or not the roundish pits of A. annulata are homologous with the subsurface pouches of the groups mentioned above or with the subcortical formation of the anlagen field in Pseudoamphisiella lacazei. If they are not homologous then the anlagen pits can be interpreted as apomorphy of Amphisiella. A further interesting morphogenetic feature of A. annulata is the formation of an additional cirral anlage between the ordinary anlagen IV and V. Since all anlagen of A. annulata – except this additional one – can be unequivocally homologised with the anlagen of A. capitata, I kept the ordinary numbering (I–VI) and designated the additional anlage, more or less arbitrarily, as anlage IVa (Berger 2004). This anlage produces only a transverse cirrus so that A. annulata has six transverse cirri (Table 15). In contrast, Amphisiella capitata lacks such an additional anlage and therefore has the ordinary number of five transverse cirri (see Fig. 16m–t). Amphisiella turanica also has six transverse cirri, indicating that it forms, like A. annulata, an additional anlage (Fig. 20a). Molecular data: The 18S rDNA of A. annulata was sequenced by Yi & Song in 2006 (GenBank accession number DQ832260; length 1773 bp). Further details see Addenda. Occurrence and ecology: Amphisiella annulata is a salt water species. Kahl (1928b) discovered it in the Brennermoor, a saline (25‰), silt peat bog near the village of Bad Oldesloe, north Germany (Kahl 1928a). Later, he found it in the harbour of the city of Kiel (Germany), Baltic Sea (Kahl 1932). Alekperov & Asadullayeva (1999) isolated their population from the periphyton of the South Apsheron coast of the Caspian Sea. Due to the neotypification by Berger (2004), the type locality of A. annulata is now the sandy beach of the northern Adriatic Sea ahead of the camp-
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ground Pra' delle Torri (45°34'N 12°49'E) near the village of Duna Verde, Italy. I found it there in a sample containing mainly washed up sea grass and sand; water temperature was about 20°C. It occurred together with Uroleptopsis citrina (see Berger 2004a), Pseudoamphisiella sp., and some euplotids. In the cultures the abundance of A. annulata was rather low, while U. citrina grew very well. Hu et al. (2004) found A. annulata twice (1996 and 2000) in mariculture water near the coast of the city of Qingdao (Tsingtao, 36°08'N 120°43'E; China) at 4°C and 15°C, 29‰ and 33‰ salinity, and pH 8.0 and 8.3. Records from the Gulf of Mexico: rare in diatom detritus from the mouth of a saltmarsh near the Florida State University Marine Laboratory at Alligator Harbor, USA (Borror 1962, 1963); interstitial of Enmedio Island and Laguna de Mandinga, Veracruz, Mexico (Aladro Lubel 1985, Aladro Lubel et al. 1988, 1990). Records not substantiated by morphological data: during summer and winter in the sublittoral (Stoller Grund, Großer Belt) of the Bay of Kiel, Baltic Sea (Bock 1952); Schlei, a polluted, brackish fjord in the western Baltic (Bock 1960, Jaeckel 1962; see also Hartwig 1974); with a frequency of 2.7% at 21–22°C and 17–18‰ salinity at the Bulgarian coast of the Black Sea (Detcheva 1982, 1983; for review, see Detcheva 1992, p. 100); sediment of Loch Eil on the west coast of Scotland, Atlantic Ocean (Wyatt & Pearson 1982); coastal areas of the Sea of Cantabria (Spain), Bay of Biscay, Atlantic Ocean (Fernandez-Leborans & Novillo 1993, p. 216). Amphisiella annulata feeds on small diatoms (Kahl 1932, Borror 1963; for review, see Fenchel 1968). In cultures, it also ingests wheat starch (Berger 2004) or bacteria (Hu et al. 2004).
Amphisiella milnei (Kahl, 1932) Horváth, 1950 (Fig. 19a–f) 1932 Amphisiella milnei spec. n. – Kahl, Tierwelt Dtl., 25: 590, Fig. 1123 (Fig. 19a; original description; no formal diagnosis provided and no type material available; see nomenclature for correct name). 1933 Amphisiella milnei Kahl 1932 – Kahl, Tierwelt N.- u. Ostsee, 23: 112, Fig. 17.10 (Fig. 19b; guide to marine ciliates; see nomenclature for correct name). 1950 Amphisiella milnei – Horváth, Annls Inst. Biol. pervest. hung., 19: 155 (combination with Amphisiella, see nomenclature). 1972 Amphisiella milnei Kahl, 1932 – Agamaliev, Acta Protozool., 10: 22, Fig. 12 (Fig. 19c, d; redescription; possibly, a voucher slide is deposited in the Institute of Zoology, Academy of Sciences, Baku, Azerbaijan). 1972 Amphisiella milnei Kahl, 1932 – Borror, J. Protozool., 19: 9 (see nomenclature; revision of hypotrichs). 1974 Amphisiella milnei Kahl – Stiller, Fauna Hung., 115: 95, Fig. 58A (Fig. 19a; revision). 1983 Amphisiella milnei Kahl, 1932 – Agamaliev, Ciliates of Caspian Sea, p. 107, Fig. 56a, b (redrawing of Fig. 19c, d; review). 1986 Amphisiela milnei Kahl, 1932 – Aladro-Lubel, Martínez-Murillo, Mayén-Estrada, HernándezAnaya & Sánchez-Calderón, Revta lat.-am. Microbiol., 28: 239, Lámina III, Fig. 5 (Fig. 19e; illustrated record; incorrect subsequent spelling of Amphisiella).
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1990 Amphisiella milnei Kahl, 1932 – Aladro Lubel, Martínez Murillo & Mayén Estrada, Manual de ciliados, p. 127, Figure on p. 127 (Fig. 19f; review). 1992 Amphisiella milnei Kahl, 1930–5 – Carey, Marine interstitial ciliates, p. 179, Fig. 700 (guide; the illustration is a redrawing of Fig. 19a). 2001 Amphisiella milnei (Kahl, 1932) ?Horvath, 1950 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 33 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs).
Nomenclature: Kahl (1932) dedicated this species to William Milne (England; e.g., Milne 1886). Kahl (1932, 1933) classified Amphisiella as subgenus of Holosticha. Thus, the correct name in his reviews is Holosticha (Amphisiella) milnei Kahl, 1932. This was obviously overlooked by all later authors, who assumed that Kahl (1932) had established the present species in the genus Amphisiella (see list of synonyms). For the sake of simplicity I suggest fixing Horváth (1950) as combining author because he was the first who used the present combination. Incorrect subsequent spellings: Amphysiella milnei Kahl (Stiller 1974, p. 94); Amphisiella milenei Kahl, 1932 (Dini et al. 1995, p. 69). Holosticha (Amphisiella) milnei Kahl, 1932 is a primary homonym of Holosticha (Holosticha) milnei Kahl, 1932 (note that different subgeneric names are irrelevant; ICZN 1999, Article 57.4). Kahl (1932, p. 590) suggested synonymy of these two species (although his illustrations look rather different) and therefore used the same species-group name. Berger (2001, p. 35) introduced a replacement name, Holosticha holomilnei, for H. (Holosticha) milnei, which is now considered as junior synonym of Anteholosticha oculata (Mereschkowsky, 1877) Berger, 2003 (for review, see Berger 2006, p. 449). Remarks: The classification in the subgenus/genus Amphisiella by Kahl (1932) and later authors seems correct because the cirral pattern matches that of A. capitata, type of the genus, very well. Kahl (1932) provided a detailed illustration of the cirral pattern, which shows that A. milnei has, like A. capitata, only two cirri left of the anterior portion of the amphisiellid median cirral row. However, the type species is more or less distinctly cephalised and lacks (i) the additional cirri between the left frontal cirrus and the buccal cirrus, and (ii) the two conspicuous ring-shaped structures (hollow spheres?) in the anterior, respectively, posterior body portion. Thus, synonymy of A. milnei and A. capitata, as suggested by Hemberger (1982), can be rejected. For a separation of A. milnei from A. annulata, see A. annulata. Anteholosticha oculata, the senior synonym of Holosticha (Holosticha) milnei Kahl, 1932 – which has, like the present species, one ring each in the anterior and posterior body portion – has a midventral complex and is therefore classified in the urostyloids (for review, see Berger 2006, p. 448). The illustration provided by Agamaliev (1972, 1983) is obviously not very exact, possibly due to the inappropriate preparation method. Thus, his data should not be over-interpreted. The cirral pattern observed by Aladro-Lubel et al. (1986) is basically identical with that described by Kahl, but they did not provide more detailed data. Thus, a thorough redescription of A. milnei is needed. The identification should be done exclusively according to Kahl (1932). The two “hollow spheres” (ring-
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shaped structures) are reminiscent of those of Tachysoma pellionellum, a common, limnetic 18-cirri oxytrichid (for review, see Berger 1999, p. 433). Morphology: This chapter is mainly based on the original description because it is the most detailed one. Body length of Kahl’s specimens 100–140 µm, body length:width ratio 3.0–3.5:1. Body outline elliptical. Body soft, flexible, and rather thick. Two macronuclear nodules slightly left of midline about in central body portion. Each macronuclear nodule with a micronucleus. Contractile vacuole neither illustrated nor mentioned, indicating that it is lacking. Obviously two types of cortical granules, namely (i) “dust-shaped colourless protrichocysts” (that is, these are obviously rather small granules), and (ii) in between loosely arranged yellowish granules (“pearls”). In anterior and posterior body portion one large hollow sphere (ringshaped structure) each; posterior sphere sometimes lacking. The following data about the oral apparatus and the infraciliature are mainly from Fig. 19a. Adoral zone occupies about 35% of body length (Fig. 19a), proximal portion sigmoidal, distal portion extends rather far posteriorly (DE-value of specimen illustrated about 0.44); membranelles narrow rather than wide. Buccal field obviously narrow, undulating membranes likely roughly straight. According to Kahl (1932) “8–10 frontal cirri arranged in three groups”. Three slightly enlarged frontal cirri; one buccal cirrus right of anterior half of undulating membranes. Between left frontal cirrus and buccal cirrus three additional cirri, which is unusual; however, a similar pattern is known from Caudiholosticha sylvatica (Foissner, 1982) Berger, 2003 (for review, see Berger 2006, p. 239), indicating that Kahl’s observation is correct. No cirrus immediately behind right frontal cirrus. Two cirri left of anterior portion of amphisiellid median cirral row (Fig. 19a, long arrow). Amphisiellid median cirral row extends from distal end of adoral zone to near right transverse cirri (composed of 16 cirri in specimen shown in Fig. 19a; value must not be over-interpreted). Five slightly enlarged transverse cirri, right two project somewhat beyond rear end of cell; transverse cirri arranged about at level of rear ring-shaped structure. Pretransverse ventral cirri neither mentioned nor illustrated (since they are usually rather small it cannot be excluded that Kahl has overlooked them). Right marginal row commences about at level of buccal cirrus, ends subterminally and therefore distinctly separated from rear end of left marginal row which commences left of proximal end of adoral zone. Marginal cirri and cirri of amphisiellid median row of ordinary width and length, that is, not as wide, short, and narrowly spaced as in A. annulata. Dorsal bristles likely of ordinary length (around 3 µm) because neither mentioned nor illustrated. Number and arrangement of dorsal kineties unknown; caudal cirri obviously lacking (Fig. 19a). Supplementary or deviating data from other populations (see also Fig. 19c–f): body length/size in life 100–125 µm (Münch 1956), 120 µm (after wet silver nitrate impregnation?; Agamaliev 1972); 108 × 29 µm (Aladro-Lubel et al. 1986); 70–108 × 21–28 µm (Aladro Lubel et al. 1990). Macronuclear nodules about 7 µm in size (if this is the length then it is very likely an underestimation; Aladro-Lubel et al. 1986).
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Fig. 19a–f Amphisiella milnei (a, from Kahl 1932; b, from Kahl 1933; c, d, from Agamaliev 1972; e, from Aladro-Lubel et al. 1986; f, from Aladro Lubel et al. 1990. a, b, e, f, from life; c, d, wet silver nitrate impregnation). a, b: Ventral views showing, inter alia, cirral pattern and nuclear apparatus, 100–140 µm. Short arrow in (a) denotes buccal cirrus, long arrow marks two cirri left of the anterior portion of the amphisiellid median cirral row. Arrows in (b) mark the anterior and posterior ring (“hollow” sphere?), which are characteristic for this species. c, d: Infraciliature of ventral side and nuclear apparatus of a Caspian Sea specimen, 87 µm. Note that the cirral pattern does not match very well Kahl’s observation. e, f: Ventral view of a specimen from the Gulf of Mexico, 108 µm. MA = macronuclear nodule, MI = micronucleus. Page 118.
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Usually two (one anterior, one posterior) ring-shaped structures (Münch 1956, Aladro-Lubel et al. 1986). Adoral zone about 35 µm (Agamaliev 1972), respectively, 38 µm long (Aladro-Lubel et al. 1986); 20–35 adoral membranelles (Agamaliev 1972). Nine cirri in frontal area and five transverse cirri (Aladro-Lubel et al. 1986). Specimen illustrated by Agamaliev (1972) with nine isolated cirri in frontal region (arrangement does not correspond with that observed by Kahl 1932), 38 cirri in amphisiellid median cirral row, five transverse cirri, 36 right marginal cirri, and 44 left marginal cirri (Fig. 19c). Occurrence and ecology: Sea and brackish water. The type locality of A. milnei is the Baltic Sea near the city of Kiel, Germany, where it is characteristic for certain sandy sites (Bülk, 5 cm deep; Kahl 1932). Further records substantiated by morphological data: benthal of an island in the Apseronskij and Bakinskij archipelagos of the Caspian Sea (Agamaliev 1972, 1983; further records see next paragraph); during April in 8–10 cm sediment depth at Boca del Rio, a beach in Veracruz, Gulf of Mexico (Aladro-Lubel et al. 1986; see also next paragraph for further Mexican records). Records from sea and brackish waters not substantiated by morphological data: sediment of the Bay of Kiel, Baltic Sea (Bock 1952, p. 81; ); Schlei, a large brackish water (partially polluted by sewage) near the city of Kiel, Germany (Bock 1960, p. 64; Jaeckel 1962, p. 13; see also Hartwig 1974, p. 17); coastal waters (salinity 5.4 to 7.8‰) of Hiddensee, an island in the Baltic Sea (Münch 1956, p. 434); brackish lake near Bay of Danzig (Poland), Baltic Sea (Czapik 1962, p. 374); with maxima (biomass up to 257 mg C m-3) during February and November in the sublittoral area in Castro Urdiales, Bay of Biscay, Atlantic Ocean (Fernandez-Leborans 2000, p. 416; 2001, p. 740; Fernandez-Leborans et al. 1999, p. 730); Italy (Dini et al. 1995, p. 69); saline groundwater of coast near the city of Kuçova (Albania), Adriatic Sea (Czapik 1952, p. 62); sediment of Loch Eil on the west coast of Scotland (Wyatt & Pearson 1982); western coast of the middle and southern part of the Caspian Sea (Agamaliev 1967, p. 369; 1974, p. 57); psammon of Kandalakshskij Bay, White Sea (Burkovsky 1970, p. 11; 1970a, p. 56; 1970b, p. 190); psammon of Ussuri Gulf, Japan Sea (Raikov 1963, p. 1757; Raikov & Kovaleva 1968, p. 331); benthal of the Laguna de La Mancha and Playa de Boca del Rio, Veracruz, Mexico (Mayén Estrada & Aladro Lubel 1987, p. 74; Aladro-Lubel et al. 1988, p. 436). Records from limnetic habitats and soil are very likely misidentifications: sewage treatment plant in Poland (Czapik 1961, p. 64); chernozem soils from the centre of the Great Hungarian Plain (Szabó 2000, p. 14). Whether the record of “Holosticha milnei Kahl” from the psammon of Lake Balaton (Hungary) by Tamás & Gellért (1960, p. 68) refers to H. (Holosticha) milnei or the present species remains unknown; anyhow, the record is probably incorrect because both Anteholosticha oculata, the synonym of H. (Holosticha) milnei, and the present species are very likely confined to marine or brackish habitats (for review of Anteholosticha oculata see Berger 2006). Feeds on diatoms and ciliates (Kahl 1932; Fenchel 1968, p. 114).
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Amphisiella turanica Alekperov & Asadullayeva, 1999 (Fig. 20a, b) 1999 Amphisiella turanica sp. nov. – Alekperov & Asadullayeva, Turkish J. Zool., 23: 219, Fig. 7 (Fig. 20a, b; original description; no formal diagnosis provided; the holotype slide [accession number M No 29/2] is deposited in the Protistological Laboratory, Institute of Zoology, Academy of Sciences of Azerbaijan, Baku).
Nomenclature: No derivation of the name is given in the original description. I do not know to which feature the species-group name refers; likely it is a geographical name. Amphisiella turanica was overlooked by Berger (2001), but is, of course, included in the updated version at www.protozoology.com. Remarks: The classification in Amphisiella is very likely correct as indicated by the cirral pattern and the saltwater habitat. Alekperov & Asadullayeva (1999) described “three complete and one short” dorsal kinety, which is a rather low value for Amphisiella. Further, “short” dorsal kineties are usually dorsomarginal rows. Thus a detailed redescription should concentrate, inter alia, on details of the cirral pattern (exact arrangement of frontal ciliature, presence/absence of pretransverse ventral cirri) and the dorsal ciliature (length of dorsal bristles; number of dorsal kineties, which is sometimes difficult to count even in silver preparations; exact arrangement; morphogenesis, to show whether or not the short kinety is a dorsomarginal row or a misobservation). Moreover, the presence/absence of cortical granules has to be checked. Amphisiella turanica is easily recognised by the four macronuclear nodules and the saline habitat. For other quadrinucleate amphisiellids, see Lamtostyla; however, these species are terrestrial. Morphology: Body length 170–210 µm in life, 130–180 µm after fixation, body length:width ratio of illustrated specimens 4.8:1 (Fig. 20a) to 6.5:1 (Fig. 20b). Body elongated, flattened dorsoventrally. Anterior body portion curved leftwards (Fig. 20a, b). Nuclear apparatus composed of four, pair-wise arranged macronuclear nodules; between each pair a single micronucleus (Fig. 20a). Contractile vacuole close to left body margin near proximal end of adoral zone of membranelles. Cytoplasm transparent, distinctly vacuolised. Presence/absence of cortical granules and movement not described. Adoral zone occupies 36% of body length in specimen illustrated (Fig. 20a), proximal portion roughly sigmoidal as in A. capitata, A. annulata, and A. milnei, and proximal end somewhat spoon-shaped as in A. annulata. Adoral zone composed of 70–85 membranelles, distal portion extends distinctly posteriorly (DE-value = 0.37). Undulating membranes long, straight, and not distinctly optically intersecting. In total seven isolated cirri on frontal area, that is, three frontal cirri, one buccal cirrus, likely one cirrus (= cirrus III/2) behind right frontal cirrus, and two cirri left of anterior portion of amphisiellid median cirral row (Fig. 20a). Amphisiellid median cirral row composed of about 67 cirri, extends slightly sigmoidally from near distal end of adoral zone to near transverse cirri. Six transverse cirri, arranged in
124
SYSTEMATIC SECTION Fig. 20a, b Amphisiella turanica (from Alekperov & Asadullayeva 1999. Silver impregnation [method not clearly indicated]). Infraciliature of ventral and dorsal side and nuclear apparatus, size not indicated. Long arrow marks the two cirri left of the anterior portion of the amphisiellid median cirral row, short arrows mark micronuclei between macronuclear pairs. Broken lines connect cirri which very likely originate from the same anlage (only shown for anlagen I–III); dotted line connects frontal cirri. ACR = amphisiellid median cirral row, AZM = distal end of adoral zone of membranelles, E = endoral, LMR = rear end of left marginal row, RMR = rear end of right marginal row, TC = transverse cirri, III/2 = cirrus III/2 (very likely), 1–4 = dorsal kineties. Page 123.
slightly subterminal, hook-shaped pseudorow; no further details (presence or absence of pretransverse ventral cirri) given. If these six cirri are true transverse cirri, then the species must have seven frontal-ventraltransverse cirri anlagen (however, it cannot be excluded that, for example, the rightmost transverse cirrus is possibly a pretransverse ventral cirrus). Right marginal row commences about at level of the two cirri left of the amphisiellid median cirral row, terminates roughly at level of transverse cirri, composed of about 41 cirri. Left marginal row composed of about 46 cirri, begins left of adoral zone, distinctly ahead of its proximal end, terminates ahead of left transverse cirri (Fig. 20a). Dorsal bristles arranged in three rows of body length and one posteriorly distinctly shortened row (see remarks). Caudal cirri absent; length of dorsal bristles neither mentioned nor illustrated. Occurrence and ecology: The type locality of A. turanica is the psammon of the North Apsheron coast (salinity 6–10‰) of the Caspian Sea (Alekperov & Asadullayeva 1999). No further records published.
Amphisiella
125
Amphisiella ovalis Fernandez-Leborans & Novillo, 1992 (Fig. 21a) 1992 Amphisiella ovalis n. sp. – Fernandez-Leborans & Novillo, Proc. biol. Soc. Wash., 105: 165, 178, Fig. 2, Table 2 (Fig. 21a; original description; no formal diagnosis provided; type slides [reference numbers 1665a–l; silver carbonate preparations] are deposited in the Laboratorio de Biología, Facultad de Biología, Universidad Complutense de Madrid, Spain). 2001 Amphisiella ovalis Fernandez-Leborans and Novillo, 1992 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 8 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2001 Amphisiella ovalis Fernandez-Leborans & Novillo sp. n. – Fernandez-Leborans & Novillo, Acta Protozool., 40: 227, Fig. 2 (Fig. 21a; see nomenclature for explanation).
Nomenclature: No derivation of the species-group name is given in the original descriptions. The species-group name oval·is, -is, -e (Latin adjective [m, f, n]; eggshaped, ovoid) obviously refers to the (natural?) body shape. Fernandez-Leborans & Novillo (1992) did not provide a formal diagnosis, which is not necessary according to the relevant code (ICZN 1985). By contrast, the ICZN (1999) recommends to include a diagnosis, that is, a summary of the characters that differentiate the new nominal taxon from related or similar taxa (Recommendation 13A). Obviously for that reason, Fernandez-Leborans & Novillo (2001, p. 225) provided a second original description, which was, however, superfluous in my opinion. Remarks: Fernandez-Leborans & Novillo (1992, 2001) provided a rather comprehensive description based on silver carbonate preparations. However, this method distorts the arrangement of the cirri so heavily that it is almost impossible to recognise the species. Moreover, no live observations have been published so that it is unknown whether or not cortical granules are present. Thus, it will be rather difficult to re-identify it. However, since the present species has many (32–45) macronuclear nodules I preliminarily accept A. ovalis, although its classification in Amphisiella is very uncertain because it lacks the cirri left of the anterior portion of the amphisiellid median cirral row. Detailed redescription from life and after protargol preparations needed. Morphology: This chapter is mainly based on the second paper by FernandezLeborans & Novillo, supplemented with some details provided in the original description. Body size 50–63 × 27–47 µm (from life? after silver preparations?). Body outline oval, that is, posterior end more broadly rounded than anterior (likely this is the unnatural shape after silver carbonate preparation). 32–45 macronuclear nodules; individual nodules 4.2–9.0 × 2.1–4.4 µm after silver carbonate preparation. 2–4 micronuclei, about 1.6–2.0 µm long. Presence/absence of contractile vacuole and cortical granules not known. Cytoplasm (inclusions, colour) and movement not described. Adoral zone 24–31 µm long, composed of 16–19 membranelles. Membranelles of middle region have four rows of basal bodies, namely, a short one with 3–4 basal bodies, an intermediate one with 9–10, and two long rows with 15–16 basal bodies each. Posteriormost membranelle composed of two rows of 6–8 basal bodies each.
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Endoral composed of a single row (stichomonad) of 22–25 basal bodies, paroral longer and composed of 24–26 basal body pairs. As mentioned above, the cirral pattern is distorted due to the preparation procedure, that is, the exact arrangement of the cirri cannot be reconstructed. For sizes of cirri see original description and Fig. 21a. Four frontal cirri (strictly speaking three frontal cirri and one buccal cirrus) right of paroral. Amphisiellid median cirral row composed of 18–22 cirri, extends from level of proximal portion of adoral zone to end of second body third. Two pretransverse ventral cirri 5–6 µm ahead of the 6–7 slightly enlarged transverse cirri, which are 6.6 to 9.6 µm away from rear body end (six, and especially seven transverse cirri is an unusual high, but not impossible numFig. 21a Amphisiella ovalis (from Fernandez-Leborans & Nober for an Amphisiella species; villo 1992. Silver carbonate impregnation). Infraciliature of ontogenetic data are needed ventral side, size not indicated. Pretransverse ventral cirri enfor correct interpretation, and circled. ACR = amphisiellid median cirral row, AZM = adoral zone of membranelles, BC = buccal cirrus, E = endoral, FC = consequently for a more right frontal cirrus, LMR = left marginal row, RMR = right proper classification of this marginal row, TC = transverse cirri. Page 125. species). 30–32 right marginal cirri and 20–24 left marginal cirri; both rows are obviously slightly shortened posteriorly. Four dorsal kineties, length of bristles and arrangement of kineties not mentioned. Occurrence and ecology: Type locality of Amphisiella ovalis is the Castro Urdiales Beach (43°22'N 0°28'W) in Spain, Cantabrian Sea, Atlantic Ocean. No further records published.
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Species indeterminata Amphisiella arenicola Fernandez-Leborans & Novillo, 1992 (Fig. 22a) 1992 Amphisiella arenicola n. sp. – Fernandez-Leborans & Novillo, Proc. biol. Soc. Wash., 105: 165, 178, Fig. 1, Table 1 (Fig. 22a; original description; no formal diagnosis provided; type slides [reference numbers 1663a–f; silver carbonate preparations] are deposited in the Laboratorio de Biología, Facultad de Biología, Universidad Complutense de Madrid, Spain). 2001 Amphisiella arenicola Fernandez-Leborans and Novillo, 1992 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 7 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2001 Amphisiella arenicola Fernandez-Leborans & Novillo sp. n. – Fernandez-Leborans & Novillo, Acta Protozool., 40: 225, Fig. 1 (Fig. 22a; see nomenclature for explanation).
Nomenclature: No derivation of the species-group name is given in the original descriptions. The species-group name arenicola (Latin adjective; living in the sand) is a composite of the Latin noun aréna (the sand) and the Latin verb cólere (inhabit), and refers to the habitat where the species was discovered. Fernandez-Leborans & Novillo (1992) did not provide a formal diagnosis. Such a diagnosis is not necessary according to the relevant code (ICZN 1985). By contrast, the ICZN (1999) recommends to include a diagnosis, that is, a summary of the characters that differentiate the new nominal taxon from related or similar taxa (Recommendation 13A). Obviously for that reason, Fernandez-Leborans & Novillo (2001, p. 225) provided a second original description, which was, however, superfluous in my opinion. Remarks: Fernandez-Leborans & Novillo (1992, 2001) provided a rather comprehensive description based on silver carbonate preparations. However, this method changes the arrangement of the cirri so heavily that it is almost impossible to recognise this species. Moreover, no live observations have been published so that we do not know whether or not cortical granules or ring-shaped structures are present. And last but not least, the morphometric characterisation does not agree with the illustration. For example, the specimen shown in Fig. 22a has 27 ventral cirri, whereas in the diagnosis a range of 52–56 ventral cirri is given. Consequently, I classify A. arenicola as species indeterminate, that is, a species that cannot be identified from its original description. However, since this is merely my opinion I present the (rather long) diagnosis from the second original description (Fernandez-Leborans & Novillo 2001) so that other workers can make their own decision (the first original description contains some further details, for example, the size of the cirral bases): “Oval elongated ciliates, with dorsal and ventral sides flattened (132–162 µm long, 37.5–67.5 µm width). Oral ciliature on the left side of ventral surface with an adoral organelles zone and a paroral formation. Adoral organelles zone comprises 36–42 organelles disposed in three parts: (a) an “anterior part” with 15–17 organelles having three parallel kineties (with 2, 6 and 6 basal bodies each); (b) an “intermediate
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Fig. 22a Amphisiella arenicola, a species indeterminata (from FernandezLeborans & Novillo 1992. Silver carbonate impregnation). Infraciliature of ventral side, size not indicated; p. 127.
part” of 16–19 organelles having three rows (with 2, 10–14 and 10–14 basal bodies each); (c) a “posterior part” of 5–6 organelles having 2 rows of 6–8 basal bodies each (the next to last organelle with 2 rows and 4 basal bodies, and the last 2 rows of 2 basal bodies each). Paroral formation composed of two components: (a) “paroral formation 1”, nearest the adoral zone of organelles, is a stichomonad of 60–68 basal bodies; (b) “paroral formation 2”, beside the frontal cirri, formed of 50–52 pairs (Fig. 22a shows only 31 pairs!) of basal bodies (diplostichomonad). Four anterior frontal cirri and three posterior or frontoterminal cirri. 50–54 right marginal cirri. 48–50 left marginal cirri. 52–56 ventral cirri with two cirri located directly above the transverse cirri. 5–6 transverse cirri. Two oval macronuclear nodules in the middle region of the body (19.5–31.5 µm long, 9.0–13.2 µm width each). Beside each macronucleus is an oval micronucleus (6.75–7.5 µm long). Dorsal surface with 5–6 kineties.” The type locality of Amphisiella arenicola is the littoral area of Gandía Beach (38°01'42"N 0°10'28"E) in Spain, Mediterranean Sea. No further records published.
Insufficient redescription
Amphisiella capitata (Perejaslawzewa, 1886) Kahl, 1930–5 – Carey, Marine interstitial ciliates, p. 179, Fig. 702 (Fig. 16f). Remarks: Kahl is not the combining author because he classified Amphisiella only as subgenus of Holosticha (details see species). Carey (1992) illustrated only the two marginal rows, that is, he omitted the amphisiellid median cirral row. Thus, the illustration is very misleading. Moreover, he illustrated six transverse cirri, which does not agree with other descriptions. It is not known whether or not Carey had original data.
Caudiamphisiella
129
Caudiamphisiella gen. nov. Nomenclature: Caudiamphisiella is a composite of the Latin noun caud·a (tail), the thematic vowel ·i-, and the genus-group name Amphisiella (see there for derivation). It refers to the fact that the type species has caudal cirri. Like Amphisiella feminine gender because ending with the Latin suffix -ella (ICZN 1999, Article 30.1.3). Characterisation (A = supposed apomorphy): Amphisiellidae with continuous adoral zone of membranelles. Undulating membranes straight and parallel. Three enlarged frontal cirri. Buccal cirrus present. Two or more cirri left of anterior portion of amphisiellid median cirral row. Postperistomial cirrus lacking. Two pretransverse ventral cirri. Five prominent transverse cirri. One left and one right marginal row. More than three bipolar dorsal kineties (A?), that is, dorsomarginal row and kinety fragmentation lacking. Caudal cirri present. Saltwater. Type species: Amphisiella antarctica Wilbert & Song, 2005. Remarks: See type species. Species included in Caudiamphisiella (alphabetically arranged basionyms are given): (1) Amphisiella antarctica Wilbert & Song, 2005. Incertae sedis: (2) Amphisiella oscensis Fernandez-Leborans, 1984.
Key to Caudiamphisiella and Caudiamphisiella-like species 1 Marine; several (about 5) cirri left of anterior portion of amphisiellid median cirral row (Fig. 23a–g). . . . . . . . . . . . . . . . . . . Caudiamphisiella antarctica (p. 129) - Limnetic; 1 cirrus left of anterior portion of amphisiellid median cirral row (Fig. 24a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amphisiella oscensis (p. 133)
Caudiamphisiella antarctica (Wilbert & Song, 2005) comb. nov. (Fig. 23a–g, Table 16) 2005 Amphisiella antarctica nov. spec.1 – Wilbert & Song, J. nat. Hist., 39: 969, Fig. 11E–K, 16A–F, Table VIII (Fig. 23a–g; original description; type slides are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria).
Nomenclature: No derivation of the name is given in the original description. The species-group name antarctic·us, -a, -um (Latin adjective [m, f, n]; living in the Antarctic Ocean) refers to the region where the species was discovered. 1 Wilbert & Song (2005) provided the following diagnosis: Slender marine Amphisiella 120–200 × 30–50 µm in vivo with narrowed caudal portion. Eight to twelve frontal and five transverse cirri; single ventral row extending to posterior third of cell length with about 29 cirri on average; two small ventral cirri close to transverse cirri; 25–32 adoral membranelles; 39–56 left and 42–55 right marginal cirri; four dorsal kineties and four caudal cirri; two macro- and two micronuclei. Cortical granules large and sparsely distributed on dorsal side.
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Remarks: Caudiamphisiella antarctica has the same ventral cirral pattern as Amphisiella species. Moreover, all these species are marine. However, the presence of distinct caudal cirri (Fig. 23g) separates C. antarctica clearly from Amphisiella species, which lack this part of the dorsal ciliature. Since the presence of this or another cirral group is generally used to distinguish genera, Caudiamphisiella is established although it is (still) monotypic. Whether the cirri shown in Fig. 23g are true caudal cirri or the posterior portion of the left or right marginal row can be unambiguously decided only by studying the dorsal morphogenesis. However, both N. Wilbert and W. Song are very experienced workers so that it is very unlikely that they misinterpreted the data. They compared their species with Holosticha (Amphisiella) thiophaga Kahl, 1928 – likely a junior synonym of Holosticha diademata (Rees, 1884) Kahl, 1932 (for review, see Berger 2006, p. 115) – and A. annulata, redescribed and neotypified by Berger (2004). It is interesting that Wilbert & Song (2005) did not discuss the presence/absence of the caudal cirri as distinguishing feature. The taxonomy of the amphisiellids is rather difficult so that we have to use all features which allow a grouping of the many species described so far. Caudal cirri are very likely part of the ground pattern of hypotrichs (Berger 2006, p. 78). Thus, their presence in Caudiamphisiella is only a plesiomorphy. Maregastrostyla also has caudal cirri, but differs from Caudiamphisiella, inter alia, in the number of cirri left of the anterior portion of the amphisiellid median cirral row (one vs. more than one). Very likely some ontogenetic differences also exist; however, the cell division of Caudiamphisiella is not yet known. Amphisiella oscensis is preliminarily classified in Caudiamphisiella because the infraciliature basically agrees with that of C. antarctica, except for the lack of distinct pretransverse ventral cirri and the single cirrus left of the anterior portion of the amphisiellid median cirral row in A. oscensis (vs. present and several in C. antarctica). Further details, see A. oscensis in this chapter. Amphisiellides atypicus, type of Amphisiellides, also has roughly the same cirral pattern as C. antarctica. However, there are some noteworthy differences which strongly indicate that Amphisiellides and Caudiamphisiella are not synonymous: (i) A. atypicus lacks pretransverse ventral cirri (vs. present in C. antarctica); (ii) A. atypicus has likely a dorsal kinety fragmentation (vs. lacking); and (iii) A. atypicus is terrestrial (vs. marine). Morphology: Body size 120–200 × 30–50 µm, on average 160 × 40 µm in life, that is, body length:width ratio about 4:1 in life (4.3:1 in protargol preparations, Table 16). Body outline slender with parallel margins; margins of posterior portion distinctly converging. Body flexible, slightly contractile, and often more or less distorted in middle portion (even twisted while gliding on debris; Fig. 23d). Body dorsoventrally flattened by about two thirds (Fig. 23c; likely this means that body height is two thirds of body width). Macronuclear nodules about 20 × 15 µm in life, slightly separated from one another and left of body-midline. Invariably two large (4–5 µm across) micronuclei, adjacent to macronuclear nodules (Fig. 23a, g). Contractile vacuole not observed. Pellicle soft and thin. Cortical granules colourless,
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Fig. 23a–e Caudiamphisiella antarctica (from Wilbert & Song 2005. a–d, from life; e, protargol impregnation). a: Ventral view of a representative specimen, 160 µm. b, c: Dorsal view showing cortical granulation and left lateral view showing dorsoventral flattening. d: Specimen bending around debris. e: Infraciliature of anterior body portion, length 63 µm. Arrow marks cirrus III/2, which originates from the same anlage as the right frontal cirrus (connected by broken line). Dotted line connects frontal cirri; cirri left of anterior portion of amphisiellid median cirral row encircled. ACR = amphisiellid median cirral row, AZM = adoral zone of membranelles, BC = buccal cirrus, E = endoral, FC = left frontal cirrus, LMR = left marginal row, P = paroral, DF = pharyngeal fibres, RMR = right marginal row. Page 129.
about 1 µm in size (shape not indicated), not grouped, and sparsely distributed on dorsal side (Fig. 23b). Cytoplasm greyish or colourless, often with refractive globules 2–5 µm across rendering cells dark at low magnification. Several to many food vacuoles. Movement relatively slow, crawling without pause on debris (Fig. 23d). Adoral zone occupies about 25% of body length, composed of 29 membranelles on average (Fig. 23a, e, f, Table 16); distal portion of zone slightly extending posteriorly. Largest membranelles about 7 µm wide. Buccal field narrow and inconspicuous. Undulating membranes roughly straight. Paroral two-rowed, parallel to endoral which is composed of zigzagging basal bodies. Along undulating membranes al-
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ways some narrowly spaced argentophilic granules (extrusomes?; Fig. 23f). Pharyngeal fibres conspicuous after protargol impregnation, about 30 µm long. Cirral pattern and number of cirri of usual variability (Fig. 23e, f, Table 16). Frontal cirri and cirrus III/2 (Fig. 23, arrow) distinctly enlarged. One buccal cirrus right of anterior portion of paroral. Usually five cirri left of anterior portion of amphisiellid median cirral row, which commences near distal end of adoral zone and extends sigmoidally to about 66% of body length. Invariably two pretransverse ventral cirri ahead of terminal transverse cirri. Right marginal row commences about at level of buccal cirrus, left row begins distinctly ahead of level of buccal vertex. Marginal rows distinctly separated posteriorly; however, gap largely occupied by terminally arranged transverse cirri (Fig. 23). Dorsal bristles about 5 µm long, arranged in four more or less bipolar kineties; each kinety composed of Fig. 23f, g Caudiamphisiella antarctica (from Wilbert & Song 2005. Protargol impregnation). Infraciliature about 10 dikinetids. Four distinct of ventral and dorsal side and nuclear apparatus, caudal cirri, that is, obviously each 105 µm. Arrowhead marks argyrophilic granules (exkinety forms a cirrus, strongly inditrusomes?) along undulating membranes, arrow decating that dorsal kinety fragmentanotes fibres extending from transverse cirri anteriad. tion and a dorsomarginal row are Asterisk in (g) marks anterior end of right marginal row. Detail labelling of cirri see Fig. 23e. ACR = amlacking (see comparison with Amphisiellid median cirral row, CC = caudal cirri, MA = phisiellides atypicus). macronuclear nodule, Mi = micronucleus, TC = transOccurrence and ecology: Maverse cirri (note the two small pretransverse cirri ahead rine. Type locality of Caudiamof transverse cirri!), 1–4 = dorsal kineties. Page 129. phisiella antarctica is the periphyton on rocks and in tidal pools in the littoral zone of Potter Cove, King George Island (62°14'S, 58°40'W), Antarctic area (Wilbert & Song 2005). It feeds on small diatoms and flagellates (Fig. 23; Wilbert & Song 2005).
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Incertae sedis in Caudiamphisiella Amphisiella oscensis Fernandez-Leborans, 1984 (Fig. 24a, Table 41) 1984 Amphisiella oscensis sp. nov.1 – Fernandez-Leborans, J. nat. Hist., 18: 25, Fig. 1–16 (Fig. 24a; original description; site where type slides are deposited not mentioned). 2001 Amphisiella oscensis Fernandez-Leborans, 1984 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 8 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs).
Nomenclature: No derivation of the name is given in the original description. The species-group name oscensis is a composite of Osca (= Huesca; province in Spain) and the Latin suffix -ens·is (geographical area of life), and refers to the area (Huesca) where the species was discovered. Remarks: Fernandez-Leborans (1984) found this species in a limnetic habitat in Spain. Unfortunately, the description – especially the illustration – is not very exact, making a serious classification difficult. Anyhow, the assignment to Amphisiella is very likely incorrect, inter alia, because this group is confined to marine habitats and lacks caudal cirri. The cirral pattern of A. oscensis more or less perfectly matches that of Amphisiellides atypicus, especially as concerns the distinct transverse cirri and the presence of caudal cirri. The presence of only one cirrus (= cirrus III/2) left of the anterior portion of the amphisiellid median cirral row is a distinct difference to Amphisiellides, which has two or more such cirri. In addition, Amphisiella oscensis has only three dorsal kineties each forming caudal cirri, that is, lacks a kinety fragmentation which is very likely present in Amphisiellides atypicus. Mainly for that reason I attach Amphisiella oscensis to Caudiamphisiella, whose type species also has only bipolar dorsal kineties. However, because of uncertainties in the description of A. oscensis and some differences to C. antarctica (one cirrus left of anterior portion of amphisiellid median cirral row vs. several; pretransverse ventral cirri lacking vs. present; limnetic vs. marine) the assignment is only provisional and without formal combination. To get a better idea of the phylogenetic position of A. oscensis, a detailed redescription showing the exact cirral and kinety pattern and its ontogenesis is needed. Further, life data (e.g., presence/absence of cortical granules, location of contractile vacuole) are required for a complete morphological characterisation.
1
Fernandez-Leborans (1984) provided the following diagnosis: Fresh-water ciliates measuring 77.7–81 µm × 31–33 µm. Elongate oval shape with a short posterior tail region (8.5 µm × 2.6 µm). The peristome (20 µm long) has a membranellar zone and a paroral formation consisting of two kineties. There are four frontal cirri, one buccal cirrus, two right marginal cirral rows, a single left marginal cirral row, five transverse cirri, six or seven caudal cirri and three dorsal rows of bristles. There are two macronuclei (17 × 9.1 µm), each with one spherical and adjacent micronucleus (1.36 µm of diameter). Morphogenesis of bipartition with five fan-shaped rows of cirral primordia.
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SYSTEMATIC SECTION
Fig. 24a Amphisiella oscensis (from FernandezLeborans 1984. Method not indicated, likely silver carbonate impregnation). Infraciliature of ventral and dorsal (broken lines) side, individual size not indicated (around 80 µm). Arrowhead marks buccal cirrus at anterior end of undulating membranes, arrow (very likely) denotes cirrus III/2. Frontal cirri are connected by dotted line. Since five transverse cirri are present, Amphisiella oscensis very likely has six frontalventral-transverse cirri anlagen. ACR = rear end of amphisiellid median cirral row, AZM = adoral zone of membranelles, CC = caudal cirri on rear portion of tail, DB = dorsal bristles (about 4 µm long), E = endoral, FC = right frontal cirrus, LMR = left marginal row, P = paroral, RMR = right marginal row, TC = transverse cirri, 1–3 = dorsal kineties. Page 133.
Morphology: Unfortunately, live data are lacking. Body size about 80 × 30 µm (method not indicated). Natural body outline not known; outline of illustrated specimen elongate oval-shaped with a short tail about 9 µm long and 3 µm wide. However, this outline is very likely from a cell prepared with the silver carbonate method, which changes the cell shape drastically. Two macronuclear nodules. One small micronucleus (only 1.5 µm across!) attached to each macronuclear nodule. Contractile vacuole (location) and details on cortex (flexible [very likely] or rigid), cortical granules (present or absent), and movement not described. Adoral zone (“oral region” in original description) about 20 µm long, that is, 25% of body length (according to Fernandez-Leborans 1984 the mean length of the adoral zone is 30.4 µm; very likely this is the full length – including the curve – of the zone), composed of membranelles of usual fine structure, that is, of four kineties of three different lengths, namely, 4.8 µm (two rows), 2.6 µm, and 1.1 µm. Endoral slightly curved and about 18 µm long on average and bipartite; anterior portion about 13 µm long, composed of a row of about 82 basal body pairs (rather high, possibly incorrect value); rear portion about 5 µm long, composed of a row of about 16 basal bodies. Paroral roughly parallel to endoral, on average about 16 µm long; anterior third composed of a row of basal body pairs; middle and rear third composed of three posteriorly slightly converg-
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Table 16 Morphometric data on Caudiamphisiella antarctica (from Wilbert & Song 2005) Characteristics a Body, length Body, width Adoral zone of membranelles, length Macronuclear nodule, length Macronuclear nodule, width Macronuclear nodules, number Micronuclei, number Adoral membranelles, number Cirri on frontal field, number b Amphisiellid median cirral row, number of cirri Pretransverse ventral cirri, number Transverse cirri, number Left marginal cirri, number Right marginal cirri, number Dorsal kineties, number Caudal cirri, number
mean 139.6 32.8 41.9 17.3 11.8 2.0 2.0 28.9 9.6 29.3 2.0 5.0 44.4 46.2 4.0 4.0
M – – – – – – – – – – – – – – – –
SD 17.4 6.8 5.6 3.2 2.9 0.0 0.0 2.2 1.3 3.3 0.0 0.0 5.2 4.4 0.0 0.0
SE 3.9 1.5 1.3 0.7 0.7 0.0 0.0 0.5 0.4 0.9 0.0 0.0 1.3 1.2 0.0 0.0
CV
Min
Max
n
12.5 112.0 181.0 20.8 23.0 46.0 13.3 34.0 52.0 18.4 12.0 23.0 24.8 7.0 17.0 0.0 2.0 2.0 0.0 2.0 2.0 7.5 25.0 32.0 16.5 8.0 12.0 11.1 21.0 36.0 0.0 2.0 2.0 0.0 5.0 5.0 11.7 39.0 56.0 9.6 42.0 55.0 0.0 4.0 4.0 0.0 4.0 4.0
20 20 20 20 20 >30 >30 16 9 12 >20 >20 13 13 13 12
c
a
All measurements in µm. CV = coefficient of variation in %, M = median, Max = maximum value, mean = arithmetic mean, Min = minimum value, n = number of individuals investigated, SD = standard deviation, SE = standard error of arithmetic mean. Data based on protargol-impregnated specimens. b
Comprising three frontal cirri, one buccal cirrus, and the cirri left of the anterior portion of the amphisiellid median cirral row (encircled in Fig. 23e).
c
Note that the specimen shown in Fig. 23f, g is only 105 µm long.
ing rows of basal bodies (Fig. 24a). For a brief description of the oral apparatus, see Fernandez-Leborans (1985, p. 371). The description of the cirri and the cirral pattern is rather comprehensive in the original description. In spite of this, the exact arrangement of the individual cirri is not known, due to the silver carbonate method. Three frontal cirri, one buccal cirrus right of anterior portion of undulating membranes, and one cirrus (very likely cirrus III/2) slightly behind these cirri. Bases of all these cirri large, that is, 3.8 × 3.4 µm on average; posteriormost cirrus with a thick, about 5 µm long fibre extending from the anterior right side of the cirrus. Amphisiellid median cirral row (designated as right marginal row 2 in original description) extends from about level of cirrus III/2 to near transverse cirri; bases of cirri about 2.8 × 1.3 µm, composed of two kineties with each five basal bodies. Transverse cirri arranged in slightly curved, distinct transverse pseudorow which is about at 73% of body length in specimen illustrated (Fig. 24a); cirri of about same size as marginal cirri. Right marginal row (termed right marginal row 1 in original description) commences near anterior body end, terminates – like left row – about at base of tail. Left marginal row commences, as is usual, near proximal end of adoral zone. Three dorsal kineties, according to Fig. 24a distinctly shortened anteriorly and posteriorly. Length of bristles not mentioned, according to illustration about 4 µm
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SYSTEMATIC SECTION
long (Fig. 24a; body length assumed 80 µm). Caudal cirri U-shaped arranged at tip of tail; each cirrus composed of four basal bodies. The high number of cirri indicates that each kinety forms 2–3 cirri. Cell division: Fernandez-Leborans (1984) described five fan-shaped frontalventral-transverse cirral anlagen (obviously without anlage I). However, no illustration, but only a not very detailed micrograph is provided. The presence of five distinct transverse cirri implies that six (I–VI) anlagen are formed although the remaining cirral pattern would require only five anlagen. One possibility is that anlage IV forms only the transverse cirrus, if we assume that the amphisiellid median cirral row is formed in the ordinary manner, that is, from the two rightmost anlagen (V and VI). However, details are needed to understand the formation exactly. Occurrence and ecology: Limnetic. Unfortunately, the type locality is not described in detail. Fernandez-Leborans (1984) discovered Amphisiella oscensis in a “water-zone” in the province of Huesca, Spain. He cultivated the specimens in 1 litre filtered water from the sample site and added six wheat grains. FernandezLeborans & Antonio-García (1988) recorded it from the Manzanares river (La Pedriza, Madrid, Spain) during a study about the effects of lead and cadmium on the protozoan community; however, no details have been provided. Food not known.
Maregastrostyla gen. nov. Nomenclature: Maregastrostyla is a composite of mare (Latin noun; the sea) and the genus-group name Gastrostyla Engelmann, 1862. No derivation of the name Gastrostyla is given in the original description and the review by Berger (1999). It is a composite of the Greek nouns he gaster (stomach) and he stylos (slate pencil; cirrus in present case). Likely it refers to the increased number of ventral cirri. Maregastrostyla refers to the fact that the type species, which was previously classified in Gastrostyla, lives in the sea. Like Gastrostyla of feminine gender (Aescht 2001, p. 283). Characterisation (A = supposed apomorphy): Amphisiellidae with continuous adoral zone of membranelles. Undulating membranes long and straight to slightly curved. Three enlarged frontal cirri. Buccal cirrus present. One cirrus (= III/2) left of anterior portion of amphisiellid median cirral row, which originates from anlagen V (forms posterior portion), IV (middle portion), and VI (anterior portion). Postperistomial cirrus indistinct. Two pretransverse ventral cirri. Five transverse cirri. One left and one right marginal row which originate de novo (A). Three dorsal kineties, which divide via intrakinetally originating primary primordia (A?) in mid-body (several parental dorsal kineties retained in neotype population1). Caudal cirri present. Proximal half of parental adoral zone completely reorganised (A?). Saltwater.
1
Borror (1963a) counted only four bipolar kineties in M. pulchra (Fig. 206j in Berger 1999); according to Kattar (1970) five dorsal kineties are present.
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Type species: Stilonichia pulchra Pereyaslawzewa, 1886. Remarks: See single species and Addenda where Maregastrostyla has to be synonymised with Protogastrostyla Gong et al., 2007! Species included in Maregastrostyla (basionym given): (1) Stilonichia pulchra Pereyaslawzewa, 1886.
Single species Maregastrostyla pulchra (Pereyaslawzewa, 1886) comb. nov. (Fig. 25a–e, 26a–r, Table 17, Addenda) 1886 Stilonichia pulchra n. sp. – Pereyaslawzewa, Zap. novoross. Obshch. Estest., 10: 90, Fig. 14 (Fig. 206a in Berger 1999; no formal diagnosis provided and likely no type material available; incorrect subsequent spelling of Stylonychia). 1888 Holosticha coronata, nov. spec. – Gourret & Roeser, Archs Biol., 8: 182, Planche XV, Fig. 1 (Fig. 25a; original description of junior synonym; see remarks; no formal diagnosis provided and likely no type material available). 1929 Gastrostyla (Stylonychia) pulchra Perejasl. 1885 – Hamburger & Buddenbrock, Nord. Plankt., 7: 93 (combination with Gastrostyla; see nomenclature). 1929 ? Gastrostyla (Holosticha) coronata – Hamburger & Buddenbrock, Nord. Plankt., 7: 93 (combination with Gastrostyla). 1932 Keronopsis (Holosticha) coronata (Gourret u. R., 1888) – Kahl, Tierwelt Dtl., 25: 576, Fig. 10121 (Fig. 25b; revision; see nomenclature). 1936 Gastrostyla (Stylonychia) pulchra Perejaslawzewa 1885 – Kiesselbach, Thalassia, 2: 20, Abb. 45 (Fig. 25c; illustrated record from northern Adriatic Sea). 1970 Gastrostyla pulchra (Perejaslawzewa, 1885) – Kattar, Zoologia e biologica marinha, 27: 186, Fig. 32 (Fig. 25e; redescription of Brazilian population). 1974 Gastrostyla (Stylonychia) pulchra (Perejaslawzewa, 1886) Wahlgren, 1890 – Jones, Univ. South Alabama Monogr., 1: 41, Fig. XXIX-1 (Fig. 25d; brief redescription; incorrect spelling of Wallengren and incorrect year). 1992 Gastrostyla pulchra (Perejaslawzewa, 1886) Kahl, 1930-5 – Carey, Marine interstitial ciliates, p. 190, Fig. 755 (redrawing of Fig. 206a in Berger 1999; guide). 1999 Gastrostyla pulchra (Pereyaslawzewa, 1886) Kahl, 1932 – Berger, Monographiae biol., 78: 818, Fig. 206a–n (detailed review). 2000 Gastrostyla pulchra (Perejaslawzewa, 1885) Kahl, 19321 – Hu & Song, Europ. J. Protistol., 36: 201, Fig. 1a–h, 2a–f, 3a–f, 4a, b, 5–15, Table 1 (Fig. 26a–r; detailed redescription, morphogenesis, and neotypification; one neotype slide and two additional slides are deposited in the Laboratory of Protozoology of the Ocean University of Qingdao, China). 2001 Gastrostyla pulchra (Pereyaslawzewa, 1886) Kahl, 1932 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 82 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs).
1
Hu & Song (2000) provided the following new diagnosis (based on neotype material only): Marine Gastrostyla in vivo measure 120–250 × 40–70 µm with elongated, slightly cephalized body shape and highly developed AZM; 44–65 membranelles; 28–40 left and 29–39 right marginal cirri; consistently 16 frontoventral cirri forming a typical Gastrostyla-pattern; 5 transverse and 3 caudal cirri; 9–11 dorsal kineties; 2 macro- and 3–4 micronuclei; cortical granules present.
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SYSTEMATIC SECTION
2002 Gastrostyla (Gastrostyla) pulchra – Foissner, Agatha & Berger, Denisia, 5: 720 (classification in subgenus Gastrostyla (Gastrostyla)). 2003 Gastrostyla pulchra Kahl, 1932 – Hu, Gong & Song, Pathogenic protozoa, p. 170, Fig. 5-8A–C (Fig. 26a, h, i; guide; incorrect authorship). 2006 Holosticha coronata Gourret & Roeser, 1888 – Berger, Monographiae Biol., 85: 95 (brief comment about exclusion from Holosticha and synonymy with G. pulchra).
Nomenclature: No derivation of the names are given in the original descriptions. The species-group name pulcher, pulchr·a, -um (Latin adjective [m, f, n]; beautiful) likely refers to beautiful general appearance. Type species of Maregastrostyla. The species-group name coronat·us, -a, -um (Latin adjective [m, f, n]; having a crown) likely refers to the prominent adoral zone which extends far onto the right body margin. Previously it was assumed that Kahl (1932) transferred the present species to Gastrostyla; however, obviously this act was done already by Hamburger & Buddenbrock (1929). Kahl (1932) classified Keronopsis as subgenus of Holosticha; thus, the correct name in his paper is Holosticha (Keronopsis) coronata Gourret & Roeser, 1888. Hu et al. (2003) assigned this species to Kahl (1932), which is certainly incorrect. In addition, they incorrectly assigned Gastrostyla to Lepsi (1928), who described Gastrocirrhus. Gastrostyla was established by Engelmann (1862). Details see Berger (2001). See Addenda for combination with Protogastrostyla! Remarks: In the review on oxytrichids I classified this species in Gastrostyla because the cirral pattern agrees very well with that of the type species G. steinii (Berger 1999). In addition, no cell division data, including dorsal kinety formation, were available at that time. Later, we studied various Gastrostyla species and established three subgenera because of ontogenetic differences (Foissner et al. 2002; see below). We assigned the present species to the nominotypical subgenus Gastrostyla (Gastrostyla) because we obviously assumed that no (detailed) ontogenetic data are available. Our assignment was uncertain because on p. 723 we wrote that G. pulchra possibly belongs to Gastrostyla (Kleinstyla). We did not consider the data of the neotype population by Hu & Song (2000) in detail because it is rather difficult to estimate the ventral row formation from their illustrations. The most interesting ontogenetic feature found by Hu & Song (2000) is the lack of dorsal kinety fragmentation, the main morphological apomorphy of the oxytrichids. This strongly indicates that G. pulchra is not an oxytrichid. By contrast, Gastrostyla steinii (type of Gastrostyla (Gastrostyla)), Gastrostyla bavariensis (type of Gastrostyla (Kleinstyla)), and G. mystacea, the type species of G. (Spetastyla), have this feature and therefore certainly belong to the oxytrichids (Hemberger 1982, Berger 1999, Foissner et al. 2002). According to molecular data, Gastrostyla steinii clusters in the stylonychines, that is, the rigid oxytrichids (Foissner et al. 2004). This position is supported by morphological data (rigid body, lack of cortical granules). By contrast, the type species of the other subgenera of Gastrostyla have flexible bodies and cortical granules (Berger 1999, Foissner et al. 2002) proving that they do not belong to the Stylonychinae. Consequently, the subgenera have to be raised to genus rank and the in-
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139
Fig. 25a–e Maregastrostyla pulchra (a, from Gourret & Roeser 1888; b, after Gourret & Roeser 1888 from Kahl 1932; c, from Kiesselbach 1936; d, from Jones 1974; e, from Kattar 1970. a–d, from life; e, protargol impregnation). a, b: Ventral view of synonym Holosticha coronata, size not indicated. The contractile vacuole in the posterior body end is very likely a misobservation. c–e: Ventral views (c = 200 µm, d = 183 µm, e = 158 µm) showing, inter alia, cirral pattern and nuclear apparatus. Note that the transverse cirri in the population described by Jones (1974) are very likely illustrated too far posteriorly. Page 137.
cluded species have to be combined. For reviews and descriptions of the species mentioned below see Berger (1999), Foissner et al. (2002), and Shi et al. (2003). Gastrostyla Engelmann, 1862. Type species (by monotypy): Gastrostyla steinii Engelmann, 1862. Species included: Gastrostyla steinii Engelmann, 1862; Gastrostyla setifera (Engelmann, 1862) Kent, 1882 (basionym: Pleurotricha setifera; according to the detailed redescription by Shi et al. [2003], the body is inflexible and cortical granules are lacking); Gastrostyla muscorum Kahl, 1932 (detailed data are lacking and thus the assignment is only preliminary). Remarks: According to morphological, ontogenetic, and molecular data (see above) Gastrostyla belongs to the Stylonychinae. Shi et al. (1999, p. 122) and Shi (2000, p. 12) established the family Gastrostylidae (see also Shi et al. 2003, p. 1419, for English diagnosis). At first it comprised Gastrostyla, Ancystropodium, Parastylonychia, and Hemisincirra (Shi et al. 1999, Shi 1999). Later it was confined to Gastrostyla (Shi et al. 2003), with G. steinii as type species. Since this species is unequivocally a Stylonychinae (see above), the gastrostylids would be a subgroup of the stylonychines. However, at present the gastrostylids are redundant.
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SYSTEMATIC SECTION
Kleinstyla Foissner, Agatha & Berger, 2002 stat. nov. Type species (by original designation): Gastrostyla (Kleinstyla) bavariensis Foissner, Agatha & Berger, 2002. Species included: Kleinstyla bavariensis (Foissner, Agatha & Berger, 2002) Foissner, Agatha & Berger comb. nov.; Kleinstyla dorsicirrata (Foissner, 1982) Foissner, Agatha & Berger comb. nov. (basionym: Gastrostyla dorsicirrata). Remarks: According to morphological (flexible body, cortical granules present) and morphogenetic data (kinety fragmentation and dorsomarginal kineties present), Kleinstyla belongs to the flexible oxytrichids, that is, it is a non-stylonychine oxytrichid (Fig. 9a). Spetastyla Foissner, Agatha & Berger, 2002 stat. nov. Type species (by original designation): Oxytricha mystacea Stein, 1859. Species included: Spetastyla mystacea (Stein, 1859) Foissner, Agatha & Berger comb. nov.; Spetastyla mystacea mystacea (Stein, 1859) Foissner, Agatha & Berger comb. nov.; Spetastyla mystacea minima (Hemberger, 1985) Foissner, Agatha & Berger comb. nov. (basionym: Gastrostyla minima). Remarks: According to morphological (flexible body, cortical granules present) and morphogenetic data (kinety fragmentation and dorsomarginal kineties present), Spetastyla is a non-stylonychine oxytrichid (Fig. 9a). Maregastrostyla does not belong to the oxytrichids because a dorsal kinety fragmentation and true dorsomarginal kineties are lacking. At the anterior end of the new right marginal row a short bristle row is formed. However, this short row does not migrate dorsally (Fig. 26k, q). By contrast, Hu & Song (2000, p. 209) supposed that G. pulchra has “a lower systematic position in the family Gastrostylidae and all three species mentioned above [G. pulchra, G. steinii, G. opisthoclada] seem to derive from different ancestors”. In addition, Maregastrostyla differs from Gastrostyla, Kleinstyla, and Spetastyla by the origin of the marginal rows (de novo vs. intrakinetal) and the fate of the parental adoral zone (proximal half completely reorganised vs. not [conspicuously] reorganised; Hu & Song 2000). In M. pulchra the frontoventral row is formed from three anlagen: anlage V forms the posterior portion; anlage IV forms the middle portion; and anlage VI forms the anterior portion. This is highly reminiscent of Spiroamphisiella hembergeri, strongly indicating that M. pulchra is also an amphisiellid. Consequently, Maregastrostyla is (preliminary) assigned to the amphisiellids. This is a further example that species from different major habitats (sea, freshwater, soil) usually do not belong to the same genus. A further major difference between Gastrostyla and Maregastrostyla is the flexibility of the cell. Maregastrostyla has the plesiomorphic, flexible body, whereas Gastrostyla has a rather rigid body which is in agreement with the classification in the Stylonychinae (Foissner et al. 2004), a subgroup of the Oxytrichidae (Berger 1999). Hemigastrostyla enigmatica is reminiscent of Maregastrostyla pulchra because the proximal portion of the adoral zone is reorganised during cell division and at least the right marginal row of the opisthe originates de novo (Song & Wilbert 1997, Song & Hu 1999). However, Hemigastrostyla enigmatica is an 18-cirri hypotrich with two fragmenting dorsal kineties (left and middle kinety); this indicates that it is an oxytrichid, although the fragmentation of the middle kinety is somewhat different
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of the typical oxytrichid fragmentation because the posterior portion migrates leftwards in Hemigastrostyla whereas rightwards in the ordinary oxytrichids. In addition, the dorsal kineties originate via primary primordia (Fig. 1g in Song & Hu 1999), as in Maregastrostyla (Fig. 26m). Of course one cannot exclude that M. pulchra is also an oxytrichid which, however, has lost dorsal kinety fragmentation. Unfortunately, Song & Wilbert (1997) fixed the little known Oxytricha stenocephala Borror, 1963 as type of Hemigastrostyla. Since its dorsal kinety pattern and formation is not known, a serious classification of Hemigastrostyla is not yet possible (see also remarks at Trachelostyla). If a redescription of H. stenocephala shows that it is closely related to M. pulchra, then Maregastrostyla is a junior synonym of Hemigastrostyla and for H. enigmatica a new genus has to be established. Only the morphology of the neotype population studied by Hu & Song (2000) is described. In addition, only the most important stages of cell division are reproduced in the present book. For a detailed description and documentation of the neotype population, see Hu & Song (2000). For a detailed review of most pre-2000 data, see Berger (1999). The list of synonyms above contains the original description, my review (Berger 1999), the description of the neotype population, and descriptions which I had overlooked (Berger 1999). The population described by Kiesselbach (1936) has, like several other populations, four macronuclear nodules; according to Hu & Song (2000) this is due to different physiological states. Jones (1974) illustrated the transverse cirri rather near the posterior body end, whereas in all other populations they are distinctly displaced anteriad (Berger 1999, Hu & Song 2000). Very likely, this is a misobservation (Fig. 25d). Synonymy of Holosticha coronata and M. pulchra is very likely and was already proposed by Borror (1972, p. 14). A transfer from Holosticha to Gastrostyla was already done by Hamburger & Buddenbrock (1929). Morphology: The neotype population described by Hu & Song (2000) agrees very well with the previous descriptions reviewed by Berger (1999). Thus, in the present chapter the neotype population is described (Fig. 26a–r, Table 17). Furthermore, some additional and/or deviating data of descriptions overlooked by Berger (1999) are reviewed (Fig. 25a–e). Description by Gong et al. (2007), see Addenda. Body size of neotype material 150–200 × 50–60 µm in life. Body outline elliptical with body length:width ratio 2.5–4.0:1; both ends rounded, frontal portion slightly head-shaped narrowed; margins of body proper more or less convex (Fig. 26a); outline rather variable, depends on nutritional and physiological stage (Fig. 26d, e). Body dorsoventrally flattened about 2:1 (Fig. 26f); consistence (flexible or rigid) not described, but very likely M. pulchra is flexible because cortical granules are present. Invariably two large macronuclear nodules left of midline in central body portion; individual nodules ellipsoidal and with many spherical chromatin bodies. Micronuclei 2.5–3.0 µm across, usually close to macronuclear nodules. Contractile vacuole not observed. Cortical granules less than 1 µm in size, colour not mentioned (thus likely colourless), usually 2–4 arranged in short row; after protargol im-
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SYSTEMATIC SECTION
Fig. 26a–i Maregastrostyla pulchra (neotype population from Hu & Song 2000. a, c–g, from life; b, protargol impregnation?; h, i, protargol impregnation). a: Ventral view of representative specimen, 198 µm. b: Extrusomes (ejected cortical granules after protargol impregnation?). c, g: Cortical granules are less than 1 µm in size. d, e: Shape variants. f: Left lateral view. h, i: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 111 µm. Arrows in (i) mark some short, parental kineties. For detailed labelling of cirri, see Fig. 26j. Page 137.
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Fig. 26j Maregastrostyla pulchra (neotype population from Hu & Song 2000. Protargol impregnation). Infraciliature of ventral side (same specimen as shown in Fig. 26h). Broken lines connect cirri which originate from the same anlage (see Fig. 26q for representative divider). Frontal cirri connected by dotted line. Short arrow marks rear end of amphisiellid median cirral row (three portions connected by dotted lines); long arrow marks postperistomial cirrus. ACR = anterior end of amphisiellid median cirral row, AZM = adoral zone of membranelles, BC = buccal cirrus, E = endoral, FC = left frontal cirrus, LMR = anterior end of left marginal row, P = paroral, PTVC = pretransverse ventral cirri, RMR = posterior end of right marginal row, TC = leftmost transverse cirrus, I–VI = frontal-ventraltransverse cirri anlagen, 1 = leftmost dorsal kinety. Page 137.
pregnation (extrusomes? 1 of Hu & Song 2000) about 2.5 µm long (Fig. b, c, g). Cytoplasm colourless to slightly greyish. Food vacuoles large. Movement moderately fast, rotates about main body axis when swimming. Adoral zone occupies about one third of body length in life (39% of body length in specimen shown in Fig. 26a), but about 44% on average in protargol preparations (Table 17), extends far posteriorly on right body margin; composed of an average of 54 membranelles (Fig. 26a, h). Undulating membranes slightly curved and rather long, optically intersecting in posterior half. Buccal field large. Pharyngeal fibres extend obliquely backwards. Cirral pattern rather constant as shown in Fig. 26h, j; cirri generally rather strong, about 15–25 µm long. Three frontal cirri along distal portion of adoral zone with right cirrus behind distal end of adoral zone. Buccal cirrus right of paroral between level of middle and right frontal cirrus. One cirrus (= cirrus III/2) left of anterior portion of adoral zone. Postperistomial cirrus (= IV/2) near buccal vertex and only indistinctly set off from amphisiellid median cirral row which extends from near distal end of adoral zone to near left pretransverse ventral cirrus; usually com1
Hu & Song (2000, p. 203) wrote that the extrusomes are only recognisable after impregnation. By contrast, Fig. 26b shows the extrusomes “in vivo”. I suppose that the “extrusomes” are the ejected cortical granules after protargol impregnation. For further data on these organelles see review by Berger (1999).
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SYSTEMATIC SECTION
Maregastrostyla
145
posed of eight cirri (postperistomial cirrus and pretransverse cirri not included). Transverse cirri about 20 µm long, but distinctly displaced anteriad and thus not projecting beyond rear body margin; rearmost cirrus at 81% of body length in specimen illustrated (Fig. 26h); slightly enlarged and arranged in hook-shaped pseudorow. Right marginal row commences slightly behind level of right frontal cirrus, extends to near midline at rear end of cell and thus optically overlapping with left marginal row which begins left of buccal vertex and ends often on dorsal side and therefore difficult to distinguish from caudal cirri. Dorsal bristles about 3 µm long, arranged in about 9–11 kineties of different length because new and parental rows are present. Three caudal cirri which are, however, difficult to recognise in life (Fig. 26i). Additional and/or deviating data from other descriptions: body size 160–200 × 40–50 µm (Kiesselbach 1936), 160 × 50 µm (Kattar 1970), body length 100–250 µm (Jones 1974); macronuclear nodules 17 × 11 µm (Katter 1970); 4–5 micronuclei (Jones 1974); contractile vacuole terminal (Gourret & Roeser 1888, Jones 1974; very likely a misobservation); adoral zone composed of 40–45 membranelles (Kattar 1970); five dorsal kineties (Kattar 1970). Cell division (Fig. 26k–r): Hu & Song (2000) studied this part of the life cycle in detail. In the present review only the most important stages are shown and the major events are described. For a more comprehensive description and documentation, see Hu & Song (2000). Stomatogenesis (Fig. 26k, l, n, p, q): The oral primordium of the opisthe is formed (apokinetally?) ahead of the leftmost transverse cirri. Later it extends to near the buccal vertex. Anlage I originates from the oral primordium and the new membranelles are formed from anterior to posterior. The proximal half of the parental adoral zone is completely reorganised. In late dividers it fuses with the parental distal portion. The parental undulating membranes are modified to anlage I of the proter which later forms the two undulating membranes and the left frontal cirrus. Formation of frontal-ventral-transverse cirri (Fig. 26l, n, p, q): The left anlagen for the proter and opisthe originate independent while the right anlagen (IV?, V, VI) very likely are primary primordia, that is, at first common anlagen for the proter and the opisthe are formed which later divide (Fig. 26l). The exact origin of each anlage is not known. The streaks I–VI form the following number of cirri (Fig. 26j, q): anlage I forms the undulating membranes and the left frontal cirrus (I/1); anlage II forms the left transverse cirrus, the buccal cirrus (II/2), and the middle frontal cirrus (II/3); anlage III forms transverse cirrus III/1, the cirrus (= III/2) left of the anterior
b Fig. 26k–o Maregastrostyla pulchra (neotype population from Hu & Song 2000. Protargol impregnation).
Infraciliature of ventral (k, l, n) and dorsal (m, o) side of very early, early, and middle dividers, k = size not indicated, l, m = 180 µm, n, o = 198 µm. Arrow in (k) marks some basal body pairs ahead of the right marginal row (note that this is not a true dorsomarginal kinety). Arrows in (m) mark primary primordia for dorsal kineties. Note that only three kineties show intrakinetal proliferation. The remaining kineties present in post-dividers are remnants of the parental ciliature. Arrows in (n) mark marginal row primordia which are formed de novo. MA = macronuclear nodules, OP = oral primordium. Page 137.
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SYSTEMATIC SECTION
Fig. 26p–r Maregastrostyla pulchra (neotype population from Hu & Song 2000. Protargol impregnation). Infraciliature of ventral (p, q) and dorsal (r) side of late dividers, p = 190 µm, q = 200 µm, r = 182 µm. Note that in Maregastrostyla pulchra the frontal-ventral-transverse cirri are formed from the ordinary six (I–VI) anlagen (see Fig. 26j). Arrow in (q) denotes the short row of basal body pairs ahead of the right marginal row. Arrows in (r) mark new caudal cirri. I–VI = frontal-ventraltransverse cirri anlagen which invariably produce the following number of cirri: 1 (plus paroral and endoral); 3; 3; 4; 5; 5. Page 137.
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Table 17 Morphometric data on Maregastrostyla pulchra (neotype population from Hu & Song 2000) Characteristics a
mean
M
SD
SE
CV
Max
n
Body, length Body, width Adoral zone of membranelles, length Adoral membranelles, number Macronuclear nodules, number Macronuclear nodules, length Macronuclear nodules, width Frontal cirri, number b Amphisiellid median cirral row, number of cirri c Transverse cirri, number Left marginal cirri, number Right marginal cirri, number Dorsal kineties, number d Caudal cirri, number
152.6 71.9 66.5 54.1 2.0 27.5 14.5 5.0 11.0 5.0 35.7 33.5 10.0 3.0
– – – – – – – – – – – – – –
15.5 10.1 6.7 4.2 0.0 6.8 3.1 0.0 0.0 0.0 3.1 6.3 0.8 0.0
3.4 2.2 1.5 0.9 0.0 1.5 0.7 0.0 0.0 0.0 0.7 1.3 0.2 0.0
10.2 117.5 180.0 14.1 50.0 87.5 10.0 50.0 77.5 7.8 44.0 65.0 0.0 2.0 2.0 24.7 18.0 35.0 21.5 10.0 20.0 0.0 5.0 5.0 0.0 11.0 11.0 0.0 5.0 5.0 7.9 28.0 40.0 18.8 29.0 39.0 8.2 9.0 11.0 0.0 3.0 3.0
21 21 21 21 21 21 21 21 21 21 21 21 13 21
Min
a All measurements in µm. CV = coefficient of variation in %, M = median, Max = maximum value, mean = arithmetic mean, Min = minimum value, n = number of individuals investigated, SD = standard deviation, SE = standard error of arithmetic mean, ? = sample size not indicated. Data based on protargolimpregnated specimens. b
Buccal cirrus and cirrus III/2 included.
c
Pretransverse ventral cirri and postperistomial cirrus included.
d
Some short parental kineties not counted.
portion of the amphisiellid median cirral row, and the right frontal cirrus (III/3); anlage IV forms transverse cirrus IV/1, the indistinct postperistomial cirrus (= cirrus IV/2), and the middle portion (two cirri) of the amphisiellid median cirral row; anlage V forms transverse cirrus V/1, the left pretransverse ventral cirrus (V/2), and the rear portion (three cirri) of the amphisiellid median cirral row; anlage VI forms the rightmost transverse cirrus (VI/1), the right pretransverse ventral cirrus (VI/2), and the anterior portion (three cirri) of the amphisiellid median cirral row. This anterior portion is of course homologous to the frontoterminal cirri. Marginal row formation (Fig. 26l, n, p, q): Marginal cirri primordia originate de novo near the parental rows. Parental cirri are resorbed. The anterior portion of the right marginal anlagen are not modified to cirri, but remain as a short row of basal body pairs. However, they are not true dorsomarginal kineties because they keep this position. Dorsal kinety formation (Fig. 26m, o, r): Dorsal kinety formation proceeds rather uncommonly in M. pulchra. Within three kineties, which are likely from the new (not parental) generation, each one primary primordium is formed intrakinetally about in mid-body (Fig. 26m). Later, these anlagen obviously divide so that each three anlagen are present in proter and opisthe. Each anlage forms one bipolar kinety and one caudal cirrus at the posterior end. Many parental kineties are retained so that interphasic specimens have a rather high number of dorsal kineties; however, the ex-
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act number is difficult to estimate because of some irregularities. Dorsal kinety fragmentation does not occur and no dorsomarginal kineties are formed. The short row of basal body pairs ahead of the new right marginal row does not migrate dorsally, but remains ahead the marginal row in non-dividers (Fig. 26k, q). Division of nuclear apparatus (Fig. 26m, o, r): This process proceeds in the plesiomorphic manner, that is, the macronuclear nodules fuse to a single mass and divide again in late dividers. The micronuclei divide mitotically. Occurrence and ecology: Maregastrostyla pulchra is confined to saline waters (Berger 1999). Due to the neotypification by Hu & Song (2000), the type locality is now the coast off Qingdao (China), Yellow Sea (22 °C, pH 7.9, salinity 28‰). Clones were cultured in boiled seawater (salinity about 31‰) with squeezed rice grains to support bacterial growth. The type locality of the synonym H. coronata is the harbour of the city of Bastia, Corsica (France). Further records substantiated by morphological data: aufwuchs of sea wall of harbour of the city of Rovignj, Croatia and in littoral mud of Island of Torcello, Venice, Italy (Kiesselbach 1936); Mobile Bay (Point Clear Light) in Alabama, USA (Jones 1974); beach in the Santos-Sao area (23°56'27''S 46°22'25''W), Brazil (Kattar 1970). Records not substantiated by morphological data: Mediterranean Sea, Italy (Dini et al. 1995, p. 70); Bay of Biscay at Castro Urdiales, Spain (Fernandez-Leborans 2000, p. 416; Fernandez-Leborans & Novillo 1994, p. 201). Further data, see Berger (1999). Neotype population feeds mainly on flagellates and bacteria (Hu & Song 2000); according to Katter (1970) and Jones (1974) diatoms, algae, Cyclidium, and Aspidisca are ingested. The synonym Holosticha coronata fed mainly on bacteria, algae, and diatoms (Gourret & Roeser 1888).
Spiroamphisiella Li, Song & Hu, 2007 (Table 18) 2007 Spiroamphisiella gen. nov.1 – Li, Song & Hu, Acta Protozool., 46 (2): 108 (original description). Type species (by original designation): Spiroamphisiella hembergeri Li, Song & Hu, 2007. 2007 Metastrongylidium n. gen.2 – Xu & Lei, Acta Protozool., 46 (2): 122 (original description of new synonym; see nomenclature). Type species (by original designation): Metastrongylidium distichum Xu & Lei, 2007.
Nomenclature: Spiroamphisiella is, according to Li et al. (2007), a composite of spiro (spiral) and the genus-group name Amphisiella (see there for derivation). It obviously refers to the fact that the type species has a twisted body. Like Amphisiella of 1
Li et al. (2007) provided the following diagnosis: Amphisiellidae with twisted body shape; one left and more than one right marginal rows spirallized along twisted body; a single spiral ventral row, which might be segmented in structure and is generated from 3 FVT-cirral anlagen; frontal, buccal, pretransverse, transverse and caudal cirri differentiated. 2 Xu & Lei (2007) provided the following diagnosis: Spirofilidae with two parallel ventral cirral rows. Buccal and frontal cirri distinctly enlarged. Without isolated postperistomial cirrus. Transverse and caudal cirri present.
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Table 18 Parameters to evaluate the precedence of one of the synonyms Metastrongylidium or Spiroamphisiella (including their type species) whose original descriptions were published in the same issue of Acta Protozoologica. “Earlier” dates/pages in bold (details see nomenclature at genus section) Taxa a
Parameter
Spiroamphisiella Li, Song & Hu, Metastrongylidium Xu & Lei, 2007 and M. distichum Xu & Lei, 2007 and S. hembergeri Li, Song & Hu, 2007 2007 Collecting material Submission of manuscript
March 2000
15 March 2006
15 January 2007
23 November 2006
Submission of revised version of manuscript
11 April 2007
17 January 2007
Acceptation of manuscript
20 April 2007
8 March 2007
121–129
107–120
Pages within issue b a
Taxa are arranged alphabetically from left to right.
b
A so-called “position precedence” (page or line) is used as criterion, for example, to fix the type species of a nominal genus or subgenus (Recommendation 69A.10 of the ICZN 1999).
feminine gender because ending with the Latin suffix -ella (ICZN 1999, Article 30.1.3). Metastrongylidium is a composite of the Greek prefix meta (next to, after) and the genus-group name Strongylidium (Xu & Lei 2007), and possibly indicates a resemblance (relationship) with Strongylidium. Neuter gender. Spiroamphisiella and Metastrongylidium are subjective synonyms according to the authors1 of the taxa and according to my opinion. Unfortunately, both original descriptions have been published in the same issue of the Acta Protozoologica resulting in a rather tricky nomenclatural situation. Such a rare case is (unfortunately very insufficiently) regulated by Article 24 (Precedence between simultaneously published names, spellings or acts) of the ICZN (1999). Since Article 24.1 (Automatic determination of precedence of names) does not apply, the determination of the first reviser (Article 24.2) is relevant. According to Recommendation 24A of the Code, the first reviser “should select the name, spelling or nomenclatural act that will best serve stability and universality of nomenclature”. Both descriptions fulfil all criteria necessary for a publication and are of the same quality. Thus other parameters are used to determine the precedence of one name over the other. According to Table 18, Spiroamphisiella has precedence over Metastrongylidium in four of 1
Weibo Song informed me that he discussed the synonymy of Spiroamphisiella and Metastrongylidium with Kuidong Xu after the publication of the original descriptions. In an e-mail he asked the scientific community how to solve the tricky nomenclatural situation. No simple solution could be found because such a situation is not clearly regulated by the ICZN (1999). On the fringes of the V European Congress of Protistology in St. Petersburg (July 23–27, 2007) the problem was again briefly discussed but no usable, practicable solution could be found. Thus, Article 24.2 (Determination by the First Reviser) has to be applied.
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SYSTEMATIC SECTION
five parameters. Accordingly I fix Metastrongylidium and M. distichum as junior synonyms of Spiroamphisiella and S. hembergeri. If it is subsequently shown that the precedence of the names can be objectively determined, my action as first reviser is nullified (Article 24.2.5). Perhaps the species-group name distichum becomes valid when ontogenetic, genetic, and/or molecular data show that S. hembergeri and M. distichum are subspecies or sibling species. Characterisation (A = supposed apomorphy): Body twisted about main axis and slightly cephalised (A). Adoral zone of membranelles continuous. Undulating membranes curved. Three enlarged frontal cirri. Buccal cirrus present. Usually one cirrus (= cirrus III/2) left of anterior portion of amphisiellid median cirral row which originates from anlagen IV (forms middle portion of row), V (forms rear portion), and VI (forms anterior portion). Postperistomial cirrus present, enlarged, and part of amphisiellid median cirral row (A). Two pretransverse ventral cirri. Usually five distinct transverse cirri. One left and more than one right marginal row (A). Three bipolar dorsal kineties, that is, dorsomarginal row and kinety fragmentation lacking. Caudal cirri present. Saltwater. Remarks: See type species. Species included in Spiroamphisiella: (1) Spiroamphisiella hembergeri Li, Song & Hu, 2007.
Single species Spiroamphisiella hembergeri Li, Song & Hu, 2007 (Fig. 27a–i, 28a–k, 29a–n, Tables 18, 19) 2007 Spiroamphisiella hembergeri spec. nov.1 – Li, Song & Hu, Acta Protozool., 46: 108, Fig. 1A–I, 2A–K, 3A–M, Table 1 (Fig. 27a–i, 28a–k; original description; the holotype slide is deposited in the Natural History Museum, London, UK; a paratype slide is deposited in the Laboratory of Protozoology, Ocean University of China, Qingdao). 2007 Metastrongylidium distichum n. sp.2 – Xu & Lei, Acta Protozool., 46: 122, Fig. 1–23, Table 1 (Fig. 29a–n; original description of new synonym; the holotype slide [accession number IC-000306-01]
1
Li et al. (2007) provided the following diagnosis: Colorless marine Spiroamphisiella, in vivo 100–190 × 25–50 µm; ca. 50 adoral membranelles and a single buccal cirrus; 3–7 frontal, 4–6 transverse and often 2 pretransverse cirri; one left and two right marginal rows; ventral row in two or three slightly detached segments, which is strongly shortened posteriorly; always 2 macronuclear nodules with up to 3 micronuclei. Constantly 3 dorsal kineties; 3 indistinctly caudal cirri. 2 Xu & Lei (2007) provided the following diagnosis: Marine Metastrongylidium about 170 × 40 µm in vivo, clavate to elongate ellipsoidal with bluntly pointed posterior end. Two macronuclear nodules with 2 or 3 micronuclei. Adoral zone composed of about 56 membranelles; one buccal and five frontal cirri in two rows; left ventral row short and composed of about 29 cirri; right ventral row long and extending on dorsal surface anteriorly and posteriorly, composed of about 53 cirri; left marginal row mostly extends on dorsal surface posteriorly and contains about 42 cirri; right marginal row distinctly shortened and composed of about 23 cirri; 3 or 4 transverse cirri; 3 delicate caudal cirri; and 3 dorsal kineties.
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and one paratype slide [IC-000306-03] are deposited in the Marine Biological Museum of the Chinese Academy of Sciences, Qingdao).
Nomenclature: Li et al. (2007) dedicated this species to Horst Hemberger, who provided, inter alia, a detailed revision of hypotrichs, including many data about cell division (Hemberger 1982). The species-group name distich·us, -a, -um ([m, f, n]; having two rows) is a composite of the Greek prefix di- (two) and the Greek substantive stich- (row, line) and refers to the two rows of buccal and frontal cirri (Xu & Lei 2007); note that this species actually has only one buccal cirrus (the other two cirri forming this pseudorow are cirrus III/2 and the postperistomial cirrus). Spiroamphisiella hembergeri is type species of Spiroamphisiella and M. distichum is type species of Metastrongylidium. Remarks: Spiroamphisiella was establish for a highly interesting marine hypotrich with a strongly twisted body (Li et al. 2007). Since the frontoventral row is formed by three anlagen they assigned it to the amphisiellids, a classification which seems appropriate. The outer right marginal row, which is probably part of a parental row, indicates an isolated position of Spiroamphisiella within the amphisiellids according to Li et al. (2007). Practically simultaneously, Xu & Lei (2007) undoubtedly described the same species and assigned it to the new spirofilid genus Metastrongylidium. Since they did not have ontogenetic data, they came to a different higher level classification than Li et al. (2007). Interestingly, Li et al. (2007) did not compare Spiroamphisiella with Amphisiella, the most similar (related?) taxon in my opinion. Significant differences to Amphisiella are: (i) the anlagen IV-cirri including the postperistomial cirrus form the middle part of the amphisiellid median cirral row (vs. anlagen IV-cirri not included); (ii) the enlarged postperistomial cirrus (vs. not enlarged, respectively present); (iii) the twisted body (vs. not twisted); (iv) the second right marginal row (vs. second row lacking); (v) the presence of caudal cirri (vs. lacking); and (vi) the low, plesiomorphic number of three dorsal kineties (vs. usually more than three bipolar kineties). Caudiamphisiella antarctica, which also has caudal cirri, has, inter alia, cortical granules (vs. lacking in S. hembergeri), four dorsal kineties (vs. three), and a less conspicuous cirral pattern on the frontal area likely because the postperistomial cirrus is lacking, respectively, not enlarged. Li et al. (2007) compared Spiroamphisiella with Pseudouroleptus (see present book), Gastrostyla Engelmann, 1862 (for review, see Berger 1999, p. 789), and Apoamphisiella Foissner, 1997 (for review, see Berger 1999, p. 781). However, all these taxa have dorsal kinety fragmentation and Gastrostyla and Apoamphisiella also have dorsomarginal rows, showing that these three genera belong to the oxytrichids (Berger 1999, 2006). In addition, all of them are limnetic or terrestrial, that is, do not contain marine species because the marine Gastrostyla stenocephala is now the type species of Hemigastrostyla Song & Wilbert, 1997 and G. pulchra is transferred to a new genus (Maregastrostyla; see present book) because it differs significantly from G. steinii, type of Gastrostyla.
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SYSTEMATIC SECTION
Spiroamphisiella
153
Maregastrostyla pulchra, redescribed, ontogenetically analysed, and neotypified by Hu & Song (2000), has a very similar cirral pattern and forms the amphisiellid median cirral row from three anlagen too. However, it lacks a twisted body, has cortical granules, and many parental kineties are retained after cell division so that M. pulchra and S. hembergeri can be easily separated. Mucotrichidium hospes, a limnetic species, has a similar general appearance, but differs distinctly from the present species in several features (Fig. 91a–h). Main differences are: limnetic vs. marine; single micronucleus between macronuclear nodules vs. 2–3 micronuclei; postperistomial cirrus distinctly behind buccal vertex vs. at level of buccal vertex; many normal-sized cirri on frontal area vs. six large cirri; one right marginal row vs. two. Cossothigma dubium (Fig. 76a–d) is much more slender than S. hembergeri, has less prominent cirri on the frontal area, has a more or less straight (not helical) right marginal row, lacks the outer right marginal row, and the head region is much more slender. Morphology: Li et al. (2007) provided a detailed description of S. hembergeri, including each one plate with micrographs of live and protargol-impregnated specimens documenting the main features. Metastrongylidium distichum matches the description of S. hembergeri more or less perfectly so that only additional and/or deviating data of the synonym are provided below. Body size of type population of S. hembergeri in life 100–190 × 25–50 µm, usually 150 × 40 µm; in protargol preparations extremely variable, namely 80–268 × 40–80 µm (Table 19), although the cell is not contractile. Body outline elongate or slenderly oval with anterior portion more or less distinctly cephalised; posterior body portion often bluntly pointed and bent slightly side-ways. Body distinctly twisted about main axis; slightly flexible, but not contractile; slightly flattened dorsoventrally with ventral side plane and dorsal side distinctly vaulted in middle portion (Fig. 27a–c, 28a–d). Invariable two macronuclear nodules left of midline behind buccal vertex; individual nodules about 20 × 10 µm in life(?); 1–3 micronuclei attached at variable positions. No contractile vacuole and cortical granules observed. Cytoplasm colourless containing large food vacuoles. Movement rather slow, obviously usually lying edgeways on substrate; sometimes jerking back and forth.
b
Fig. 27a–g Spiroamphisiella hembergeri (from Lin et al. 2007. a–c, from life; d–g, protargol impregnation). a: Ventral view of a representative specimen, 156 µm. b: Ventral view of a slightly oval (contracted?) specimen, 150 µm. c: Right lateral view (in original description designated as left lateral view). d, e: Infraciliature of ventral and dorsal side and nuclear apparatus, 144 µm. Short arrow marks postperistomial cirrus, long arrow designates outer right marginal row; asterisk marks cirrus III/2. Pretransverse ventral cirri circled. Frontal cirri connected by dotted line, anteriormost cirri originating from anlagen II and III connected by broken lines. f: Infraciliature of oral region; note that this specimen has eight (instead of the usual six) enlarged cirri. g: Infraciliature of ventral side and nuclear apparatus of a slender specimen. ACR = amphisiellid median cirral row, AZM = adoral zone of membranelles, BC = buccal cirrus, CC = caudal cirri, E = endoral, FC = left frontal cirrus, LMR = left marginal row, MA = macronuclear nodule, MI = micronucleus, P = paroral, RMR = right marginal row, TC = transverse cirri, IV, V, VI = frontal-ventral-transverse cirri anlagen, 1–3 = dorsal kineties. Page 150.
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SYSTEMATIC SECTION
Fig. 27h, i Spiroamphisiella hembergeri (from Lin et al. 2007. Protargol impregnation). Infraciliature of ventral side of a middle and a late reorganiser, sizes not indicated. The specimen shown in (h) has a supernumerary frontal-ventral-transverse cirri anlage. Cirri originating from the same anlage are connected by broken lines. I–VI = frontal-ventraltransverse cirri anlagen. Page 150.
Adoral zone prominent, occupies 36% and 43% of body length in specimens illustrated (Fig. 27a, b) and 43% on average in protargol preparations (Table 19); according to text of original description it occupies almost half of body length; zone composed of 50 membranelles on average. Cilia of membranelles up to 20 µm long. Distal end of zone extends rather far posteriorly; DE-values of specimens shown in
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155
Fig. 27d, g are 0.50 and 0.58, respectively. Undulating membranes rather long and distinctly curved, optically intersecting near rear end in specimen shown in Fig. 27d. Pharyngeal fibres extend posteriorly (Fig. 27b, 28f). Cirral pattern and number of cirri rather variable (Table 19) and difficult to interpret without ontogenetic data; fortunately, Li et al. (2007) described two reorganisers so that the cirral pattern can be explained in detail. Interestingly, Li et al. (2007) did not homologise the cirri with those of other amphisiellids or 18-cirri hypotrichs. Frontal cirri, buccal cirrus, cirrus III/2, and cirrus IV/2 distinctly enlarged, cilia about 20 µm. Frontal cirri arranged in very oblique pseudorow along distal portion of adoral zone with right cirrus (III/3), as is usual, at distal end of zone; distance between right cirrus and middle cirrus greater than that between middle and left. Buccal cirrus somewhat behind anterior end of paroral (Fig. 27d, 28h–j). Cirrus III/2 rather large and thus prominent, distinctly behind right frontal cirrus. Specimens with few additional enlarged cirri right of proximal portion of adoral zone occur only rarely (Fig. 27f). The ordinary pattern, however, is that shown, for example, in Fig. 27d, where the frontal cirri, cirrus III/2, and cirrus IV/2 (= postperistomial cirrus) form a bow. Amphisiellid median cirral row (termed ventral row in original description) extends from near distal end of adoral zone to 50–69% of body length (Fig. 27d, g); row somewhat irregular because composed of cirri originating from three anlagen (details see chapter reorganisation). Anterior portion1 terminates at enlarged cirrus IV/2, that is, the postperistomial cirrus is not behind the buccal vertex as is usual, but more or less in line with the remaining cirri of the amphisiellid median cirral row; posterior portion commences slightly behind level of buccal vertex. Cirri of amphisiellid median cirral row and marginal cirri of same size (except of cirrus IV/2 which is distinctly enlarged), about 15 µm long, and arranged in shallow, spiral grooves (Fig. 27a, b, d, 28e). Usually two fine pretransverse ventral cirri, one (VI/2) ahead of transverse cirrus VI/1, the other (V/2) ahead of left transverse cirrus. Usually five slightly enlarged transverse arranged in hook-shaped pattern at base of “tail”; in life about 20 µm long (Fig. 27d, g, Table 19). Inner right marginal row commences dorsally ahead of level of distal end of adoral zone, extends helically onto ventral side and terminates ahead of transverse cirri. Outer right marginal row begins about at mid-body and terminates near cell end; cirri of outer row not distinctly more widerly spaced than those of inner row (Fig. 27d). Left marginal row commences at proximal end of adoral zone, extends helically onto dorsolateral surface posteriorly and terminates near rear cell end. Dorsal bristles 3–5 µm long, invariably arranged in three helical, bipolar kineties. Three inconspicuous caudal cirri at tip of rear cell end (Fig. 27d, e, 28g, k, Table 19). Additional and/or deviating data from the synonym M. distichum (Fig. 29a–n, Table 19): body size 140–200 × 30–50 µm in life, usually about 170 × 40 µm as estimated from measurements from live and protargol-impregnated specimens (Table 19). Length:width ratio 4–5:1 in life, while 3.0–4.2 in protargol preparations. Body 1
Note that this portion is composed of cirri of anlagen IV and VI (see “Reorganisation” below).
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SYSTEMATIC SECTION
Spiroamphisiella
157
Fig. 28h–k Spiroamphisiella hembergeri (from Lin et al. 2007. Protargol impregnation). h: Ventral view showing cirral pattern and nuclear apparatus. i: Infraciliature of anterior body portion. j: Ventral view of middle body portion. Arrow marks anterior end of posterior portion of amphisiellid median cirral row. k: Dorsal view showing dorsal kineties (arrows) and anterior end of inner right marginal row (double arrowhead). MA = macronuclear nodule. Bar 100 µm. Page 150.
flattened about 1.5:1 dorsoventrally. Body outline usually clavata to elongate elliptical with bluntly pointed posterior end, often widest in mid-body in protargol preparations. Chromatin bodies up to 4 µm across. Contractile vacuole at left body margin ahead of mid-body (Fig. 29a–c). Cortex very flexible, especially in anterior body half. No cortical granules recognisable. Cytoplasm colourless, with numerous lipid droplets; occasionally colourful due to food. Movement without peculiarities; may attach to substrate with rearmost cirri (especially transverse cirri) while swinging with very flexible anterior body half (Fig. 29c). Adoral zone occupies about 40% of body length on average in protargol preparations (Table 19). Undulating membranes about of same length, optically intersecting in anterior quarter and arranged in parallel. Cirral pattern as in population studied by Li et al. (2007). Buccal cirrus about at optical intersection of undulating membranes. Cilia of enlarged cirri about 20 µm long. Dorsal bristles about 6 µm long. Caudal cirri delicate; middle one composed of four basal bodies only, left and right cirrus composed of six basal bodies (Fig. 29g, i). Reorganisation (Fig. 27h, i): Li et al. (2007) found several reorganisers showing the main events of cirral pattern formation (for some micrographs, see Fig. 3I–M in Li et al. 2007). Accordingly, the plesiomorphic number of six (I–VI) anlagen is formed. These anlagen produce the following cirri: anlage I forms the left frontal cirrus (I/1); anlage II forms the leftmost transverse cirrus (II/1), the buccal cirrus
b Fig. 28a–g Spiroamphisiella hembergeri (from Lin et al. 2007. From life). a, b: Ventral and dorsal view of a representative specimen. Arrows indicate cirral rows extending onto dorsal side. c: Lateral view showing dorsoventral flattening. d: Shape variant. e: Ventral view showing distinctly twisted body. Arrows mark inner right marginal row. f: Anterior body portion. g: Dorsal side with 3–5 µm long dorsal bristles (arrows). Bars 80 µm. Page 150.
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Fig. 29a–f Spiroamphisiella hembergeri (from Xu & Lei 2007. a–c, from life; d–f, protargol impregnation). a: Ventral view of a representative specimen, 170 µm. b: Lateral view showing, inter alia, the contractile vacuole. c: Attached specimen with swinging body. d, e: Schematic representation of infraciliature of type specimen (details see Fig. 29j–n). Arrow marks outer right marginal row. f: Right lateral view of anterior body portion. ACR = amphisiellid median cirral row, AZM = adoral zone of membranelles, CV = contractile vacuole, RMR = right marginal row. Page 150.
(II/2), and the middle frontal cirrus (II/3); anlage III forms the transverse cirrus III/1, the paramalar cirrus (III/2), and the right frontal cirrus (III/3); anlage IV forms transverse cirrus IV/1, the postperistomial cirrus IV/2 (not behind buccal vertex, but in line with amphisiellid median cirral row in S. hembergeri!), and about four cirri forming the middle portion of the amphisiellid median cirral row (one of these cirri is homologous with cirrus IV/3, usually the rearmost frontoventral cirrus of the 18cirri hypotrichs); anlage V forms transverse cirrus V/1 (= rearmost transverse cirrus), the left pretransverse ventral cirrus (V/2), and the posterior portion of the amphisiellid median cirral row; and anlage VI forms the rightmost transverse cirrus (VI/1), the right pretransverse ventral cirrus (VI/2), and the anterior portion of the amphisiellid median cirral row (homologous to the frontoterminal cirri of the 18cirri hypotrichs). Rarely, an additional anlage producing some supernumerary cirri occurs (Fig. 27h). Each one marginal row anlage occurs right of the parental rows forming the new left and the new inner right marginal row (Fig. 27i, h). The origin of the outer right marginal row remains unclear; according to Li et al. (2007) the outer row seems to be the intact parental row since no primordium for this row is produced. I somewhat doubt this assumption because the cirri of the outer row are as narrowly spaced as
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Fig. 29g–i Spiroamphisiella hembergeri (from Xu & Lei 2007. Protargol impregnation). g, h: Infraciliature of ventral and dorsal side of rear body portion of same specimen. i: Infraciliature of ventral side showing, inter alia, transverse cirri and caudal cirri. ACR = amphisiellid median cirral row, CC = caudal cirri, LMR = left marginal row, RMR = right marginal row, TC = transverse cirri, 1–3 = dorsal kineties. Page 150.
Fig. 29j–n Spiroamphisiella hembergeri (from Xu & Lei 2007. Protargol impregnation). Infraciliature of ventral and dorsal side and nuclear apparatus of type specimen of synonym Metastrongylidium distichum, j, k = 178 µm. ACR = amphisiellid median cirral row, E = endoral, LMR = left marginal row, MA = macronuclear nodule, P = paroral, RMR = right marginal row, 2, 3 = dorsal kineties. Page 150.
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Table 19 Morphometric data on Spiroamphisiella hembergeri (hem, from Li et al. 2007; dis, synonym Metastrongylidium distichum from Xu & Lei 2007) Characteristics a
Population mean
Body, length Body, width Body length:width, ratio Adoral zone of membranelles, length Adoral zone length:body length, ratio Macronuclear nodules, length Macronucleus nodules, width Macronuclear nodules, number Micronuclei, diameter Micronuclei, number Adoral membranelles, number Cirri on frontal field, number b Buccal cirri, number Amphisiellid median cirral row, number of cirri in anterior portion Amphisiellid median cirral row, number of cirri in posterior portion Amphisiellid median cirral row, total number of cirri c Pretransverse ventral cirri, number Transverse cirri, number Left marginal cirri, number Inner right marginal row, number of cirri Outer right marginal row, number of cirri Dorsal kineties, number Caudal cirri, number
hem dis hem dis dis hem dis dis dis dis hem dis dis hem dis hem dis hem dis hem dis hem
M
192.4 192.0 172.8 177.5 55.2 52.0 50.2 46.5 3.5 3.4 83.5 84.0 75.2 78.0 0.4 0.4 22.0 20.0 11.6 12.0 2.0 2.0 2.0 2.0 3.6 3.5 2.4 2.0 2.3 2.0 49.2 50.0 55.2 56.0 4.7 5.0 5.0 5.0 1.0 1.0 1.0 1.0 16.2 16.0
SD 38.1 19.9 12.1 7.8 0.5 14.2 6.2 – 4.2 1.4 0.2 0.0 0.7 0.6 0.5 3.2 3.1 1.0 0.0 0.0 0.0 2.7
SE
CV
Min
Max
n
– 198.8 80.0 268.0 8.1 11.5 145.0 200.0 – 21.9 40.0 80.0 3.2 15.5 43.0 60.0 0.2 13.8 3.0 4.2 – 17.0 56.0 116.0 2.5 8.3 65.0 80.0 – 4.4 0.4 0.5 1.5 19.3 18.0 30.0 0.5 12.1 10.0 13.0 – 10.0 2.0 3.0 0.0 0.0 2.0 2.0 0.3 20.5 3.0 5.0 – 25.0 1.0 3.0 0.2 20.6 2.0 3.0 – 6.5 44.0 54.0 1.2 5.5 51.0 58.0 – 21.3 3.0 7.0 0.0 0.0 5.0 5.0 – 0.0 1.0 1.0 0.0 0.0 1.0 1.0 – 16.7 9.0 23.0
19 6 19 6 6 19 6 6 8 8 19 8 8 11 8 19 6 19 8 19 8 19
hem
15.7
16.0
2.4
–
15.3
11.0
19.0
19
dis
29.3
29.5
2.8
1.4
9.4
26.0
32.0
4
hem hem dis hem dis hem dis d hem dis hem dis hem dis
1.8 4.9 3.6 21.9 42.5 46.4 53.0 37.8 22.8 3.0 3.0 3.0 3.0
2.0 5.0 4.0 21.0 42.0 46.0 52.5 39.0 22.5 3.0 3.0 3.0 3.0
0.6 0.5 0.5 7.7 3.0 6.5 1.4 7.4 1.7 0.0 0.0 0.0 –
– – 0.2 – 1.5 – 0.7 – 0.9 – 0.0 – –
33.3 10.2 15.2 35.2 7.1 14.0 2.7 19.6 7.5 0.0 0.0 0.0 –
0.0 4.0 3.0 8.0 40.0 35.0 52.0 17.0 21.0 3.0 3.0 3.0 3.0
2.0 6.0 4.0 42.0 46.0 61.0 55.0 50.0 25.0 3.0 3.0 3.0 3.0
19 19 5 19 4 19 4 19 4 19 5 19 3
a All measurements in µm. CV = coefficient of variation in %, M = median, Max = maximum value, mean = arithmetic mean, Min = minimum value, n = number of individuals investigated, SD = standard deviation, SE = standard error of arithmetic mean. Data based on protargol-impregnated specimens. b
Comprising three frontal cirri, cirrus III/2 (= paramalar cirrus), cirrus IV/2 (= postperistomial cirrus) and 0–2 enlarged cirri between cirri III/2 and IV/2 (Fig. 27f).
c
Designated left ventral cirri in original description. Enlarged cirrus (= postperistomial cirrus) dividing the row likely included.
d
Designated as right ventral row in original description.
Lamtostyla
161
those of the inner, newly formed row. However, if parental rows are split into two parts then the distance between the individual cirri must increase. Since several cirri of the parental inner right marginal row are dissolved during reorganisation (Fig. 27h, i) it is also unlikely that the new outer right marginal row is the anterior or posterior half of the parental inner right marginal row. Likely, a specific mode is responsible for the formation of the outer row. The dorsal kineties obviously develop within the parental rows, that is, plesiomorphically (Fig. 27h, i). Occurrence and ecology: Likely confined to saline water. Type locality of Spiroamphisiella hembergeri is a scallop-farming pond (salinity 25‰) near the city of Laizhou (37°6'N 119°54'E), eastern China (Li et al. 2007). They discovered it during spring when the water temperature was 5–10°C. The type locality of the synonym Metastrongylidium distichum is the coastal water of Inchon Harbour in Korea (37°27'N 126°35'E) where Xu & Lei (2007) discovered it in March 2000 by using polyurethane foam units. The population was collected at the following conditions: water temperature 3.5°C, salinity around 31.5‰, pH 8.4, dissolved oxygen 12.2 mg l-1. The water was quite nutrient-rich due to the discharge of domestic sewage, industrial effluents, and waste loadings from the estuary (Xu & Lei 2007). Spiroamphisiella hembergeri feeds on diatoms (Li et al. 2007, Xu & Lei 2007) and greenish cyanobacteria, indicating that it is primarily herbivorous (Xu & Lei 2007).
Group II: Terrestrial Amphisiellids with six (I–VI) Frontalventral-transverse Cirri Anlagen This group comprises three relatively large taxa (Lamtostyla, Uroleptoides, Hemiamphisiella) which form, like the members of group I, their frontal-ventral-transverse cirri from six anlagen. However, group II species occur only in terrestrial habitats and are therefore usually rather slender. Unfortunately, the type species of both main genera (Lamtostyla, Uroleptoides) are not described in detail so that some uncertainty about their phylogenetic position remains.
Lamtostyla Buitkamp, 1977 1977 Lamtostyla lamottei n. gen. n. spec. – Buitkamp, Acta Protozool., 16: 270 (original description; no formal diagnosis provided). Type species (by original designation and monotypy): Lamtostyla lamottei Buitkamp, 1977. 1979 Lamtostyla Buitkamp, 1977 – Trudy zool. Inst., 86: 84 (generic catalogue of hypotrichs and euplotids). 1979 Lamtostyla Buitkamp, 1977 – Corliss, Ciliated protozoa, p. 309 (revision). 1979 Lamtostyla Buitkamp, 1977 – Tuffrau, Trans. Am. microsc. Soc., 98: 526 (revision). 1983 Lamtostyla Buitkamp, 1977 – Curds, Gates & Roberts, British and other freshwater ciliated protozoa, p. 410 (guide to ciliate genera). 1985 Lamtostyla – Small & Lynn, Phylum Ciliophora, p. 456 (guide to ciliate genera).
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SYSTEMATIC SECTION
1986 Lamtostyla Buitkamp, 1977 – Dragesco & Dragesco-Kernéis, Faune tropicale, 26: 456 (guide to ciliates of tropical Africa). 1987 Lamtostyla1 – Berger & Foissner, Zool. Jb. Syst., 114: 217 (improved diagnosis). 1987 Lamtostyla Buitkamp, 1977 – Tuffrau, Annls Sci. nat. (Zool.), 8: 116 (revision). 1988 Lamtostyla Buitkamp, 19772 – Berger & Foissner, Zool. Anz., 220: 114 (revision and improved diagnosis). 1996 Lamtostyla Buitkamp, 19773 – Petz & Foissner, Acta Protozool., 35: 277 (improved diagnosis). 1999 Lamtostyla Buitkamp, 1977 – Shi, Acta Zootax. sin., 24: 255 (revision of hypotrichs). 1999 Lamtostyla Buitkamp, 1977 – Shi, Song & Shi, Progress in Protozoology, p. 103 (revision of hypotrichs). 1999 Lamtostyla Buitkamp, 1977 – Berger, Monographiae biol., 78: 893 (brief note on systematic position). 2001 Lamtostyla Buitkamp 1977 – Aescht, Denisia, 1: 90 (catalogue of generic names of ciliates). 2001 Lamtostyla Buitkamp, 1977 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 45 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Lamtostyla Buitkamp, 1977 – Lynn & Small, Phylum Ciliophora, p. 459 (guide to ciliate genera). 2006 Lamtostyla Buitkamp, 1977 – Berger, Monographiae biol., 85: 1210 (brief note on systematic position).
Nomenclature: No derivation of the name is given in the original description. Lamtostyla is a composite of Lamto, a village in Ivory Coast, and the Greek noun ho stýlos (style, pillar; cirrus, an important feature of the hypotrichs); it obviously alludes to the fact that this hypotrich was discovered near Lamto. Feminine gender (Aescht 2001, p. 287). Characterisation (A = supposed apomorphy): Amphisiellidae with continuous adoral zone of membranelles. Three more or less distinctly enlarged frontal cirri. Buccal cirrus present. Two or more cirri left of anterior portion of amphisiellid median cirral row which terminates usually ahead of mid-body and originates from anlage V (posterior portion) and VI (anterior portion). Postperistomial cirrus sensu stricto lacking. Pretransverse ventral cirri in some species present. Transverse cirri present, sometimes arranged in U-shaped pattern. One right and one left marginal row. Four (type species), two, three, or five dorsal kineties; dorsomarginal row and kinety fragmentation lacking. Caudal cirri lacking (A?). Oral primordium originates apokinetally, that is, without contact to parental cirri. Terrestrial.
1 Berger & Foissner (1987) provided the following improved diagnosis: Usually small, wide to long ellipsoid Oxytrichidae without caudal cirri and postoral ventral cirri. Number of transverse cirri and adjacent ventral cirri usually reduced. 1 short frontal row, which originates from 2 streaks. 2 Berger & Foissner (1988) provided the following improved diagnosis: Small to medium sized Oxytrichidae. Frontoventral infraciliature restricted to the area right of the buccal cavity and at most 2 cirri adjacent to the transverse cirri. 1 frontal row which originates from 2 streaks. Number of transverse cirri usually reduced (<5). Infraciliature of dorsal surface consists of dorsal kineties only (Caudal cirri absent). 3 Petz & Foissner (1996) provided the following improved diagnosis: The oral primordium originates apokinetally near the transverse cirri. The amphisiellid median cirral row commences anlagen formation at its posterior end and originates from two rightmost anlagen. All dorsal kineties develop intrakinetally. At least one cirrus left of amphisiellid median cirral row. Transverse cirri obliquely arranged, originate from more than one anlage. Caudal cirri lacking.
Lamtostyla
163
Additional characters: body flexible; two or four macronuclear nodules in left body portion; contractile vacuole about in mid-body or slightly ahead of it; dorsal bristles short, that is, usually less than 5 µm long. Remarks: Buitkamp (1977) supposed that Lamtostyla belongs to the Holostichidae. He considered the shortened ventral row as genus feature. Corliss (1979) and Tuffrau (1979) accepted this proposal. However, a classification in the urostyloids, to which the holostichids belong (Berger 2006), is incorrect because a midventral complex – the main morphological apomorphy of the urostyloids – is very likely lacking in L. lamottei, type of the present genus (since we do not know the ontogenesis of L. lamottei we cannot completely exclude that Lamtostyla is a urostyloid). Hemberger (1982, p. 277) did not consider Lamtostyla in his revision; he mentioned L. lamottei in a list of species which are either indeterminable or not assignable to a certain higher taxon. Small & Lynn (1985) classified Lamtostyla in the Cladotrichidae, whereas Lynn & Small (2002) assigned it to the Trachelostylidae. Jankowski (1979), Dragesco & Dragesco-Kernéis (1986), Tuffrau (1987), Berger & Foissner (1988), Foissner & Foissner (1988, p. 89), and Tuffrau & Fleury (1994, p. 143) put it into the Oxytrichidae. We explained this classification by the six typical frontalventral-transverse cirri primordia which are characteristic for the oxytrichids (Berger & Foissner 1988). However, in the meantime we know that the six primordia are a feature already present in the last common ancestor of the hypotrichs and therefore cannot be used to define a group within the hypotrichs (Berger 2006). Petz & Foissner (1996) were the first who included Lamtostyla in the Amphisiellidae because the ontogenesis of L. edaphoni (now Lamtostylides edaphoni) proceeds basically as in Amphisiella, except for the origin of the oral primordium, namely, apokinetally in Lamtostyla against in close contact to parental cirri in Amphisiella (Wicklow 1982b). Accordingly, this detail of the cell division is the sole morphological difference between Amphisiella and Lamtostyla (Petz & Foissner 1996). However, the present review also shows that Amphisiella usually has more dorsal kineties and is obviously confined to saltwater, whereas Lamtostyla inhabits the terrestrial soil. Shi (1999) and Shi et al. (1999) accepted the classification of Lamtostyla in the Amphisiellidae. Eigner & Foissner (1994) did not consider it in their Amphisiellidae paper, likely because the cell division of the type species is not known. I consider, like Petz & Foissner (1996), the classification of Lamtostyla in the amphisiellids as the most parsimonious solution because both Amphisiella and the present genus have a distinct cirral row formed by the two rightmost anlagen. Further, Lamtostyla very likely lacks, like Amphisiella, a dorsomarginal row and a dorsal kinety fragmentation, preventing a classification in the Dorsomarginalia (with dorsomarginal row; Berger 2006), respectively, the Oxytrichidae (a subgroup of the Dorsomarginalia with kinety fragmentation; Berger 1999, 2006). Unfortunately, we do not know the exact dorsal kinety pattern (including its formation) of L. lamottei (type species) so that this is not 100% certain.
164
SYSTEMATIC SECTION
Several species previously assigned to Amphisiella are transferred to Lamtostyla in the present review (e.g., Amphisiella elegans). The main reasons for these transfers are the short amphisiellid median cirral row (vs. long in Amphisiella), the lower number of transverse cirri and dorsal kineties as well as the different habitat (Amphisiella – marine against Lamtostyla – terrestrial). Shi (1999) and Shi et al. (1999) consider Terricirra Berger & Foissner, 1989 as junior synonym of Lamtostyla. However, Terricirra has a special type of food vacuoles and very conspicuous cortical granules, making a synonymy with Lamtostyla very unlikely (further details, see Terricirra). Interestingly, some lamtostylid species (Lamtostyla decorata, L. longa, L. granulifera, Lamtostylides hyalinus) have the transverse cirri arranged in a roughly U-like pattern with a pretransverse(?) ventral cirrus in the U-cavity. Large Lamtostyla species have a deep buccal cavity and the undulating membranes arranged side by side and distinctly curved (Foissner et al. 2002, p. 719). I am almost convinced that Lamtostyla in the present form is a non-monophyletic group as indicated by the rather different cirral and dorsal kinety pattern. Most species so far included in Lamtostyla have more than one cirrus left of the anterior portion of the amphisiellid median cirral row. This is due to the fact that these species form their frontal-ventral-transverse cirri from the anlagen I–VI. By contrast, there are four species which have only one cirrus (= cirrus III/2; right frontal cirrus in Lamtostyla hyalina?) left of the anterior portion of the amphisiellid median cirral row. This reduced number is caused by the lack of anlage IV (Fig. 64p, r, t, 66q). Since the number of cirri left of the amphisiellid median cirral row is now generally used as main feature to define amphisiellid taxa (Foissner 1988, Eigner & Foissner 1994, Petz & Foissner 1996), I separate these species from Lamtostyla and put them in the new genus Lamtostylides. Within the remaining Lamtostyla species three groups can be distinguished: (i) the Lamtostyla lamottei-group, which has an amphisiellid median cirral row composed of more than four cirri and very likely lacks cortical granules. This group comprises the following species: L. lamottei (presence/absence of cortical granules not known), L. australis, L. procera, L. vitiphila, L. quadrinucleata, L. elegans, L. islandica, L. perisincirra; (ii) the L. granulifera-group, which basically has a cirral pattern like 18-cirri hypotrichs, except that the postoral ventral cirri are displaced anteriad. Thus, the amphisiellid median cirral row is composed of only four cirri (from anterior: VI/4, VI/3, V/4, V/3; see Fig. 40n). This group has, like many species from the L. lamottei-group, three dorsal kineties. In addition, both species included (L. granulifera, L. decorata) have cortical granules; (iii) the Lamtostyla longa-group, which has basically the same cirral pattern as the L. granulifera group, but five rather than three dorsal kineties, including a short one on the right cell margin. This group comprises L. longa and L. raptans. The short dorsal kinety indicates that at least a dorsomarginal row is present, and the number of five kineties suggests that even a kinety fragmentation is present. How-
Lamtostyla
165
ever, since an illustration of the dorsal infraciliature is not provided and the formation of the dorsal kineties is not known in either species, they are preliminarily retained in Lamtostyla. Whether cortical granules are present or not is not known because Hemberger (1982, 1985) did not study live specimens. The differences between the L. lamottei-group and the two other groups could be used to establish (sub)genera. However, this would result in three (sub)genera which basically differ only in the length of the amphisiellid median cirral row: more than 50% of body length in Uroleptoides; less than 50% of body length, but more than four cirri in the Lamtostyla lamottei-group; and only four cirri in the L. granulifera group and the L. longa group. Preliminarily I refrain from this step and recommend awaiting further data (e.g., ontogenetic data of the 18-cirri species; molecular data) to get a better, more well-founded idea of the phylogenetic relationships. As already mentioned above, Lamtostyla longa and L. raptans have five dorsal kineties, making the classification in the present genus rather uncertain (details, see individual species). Lamtostyla species do not have a postperistomial cirrus sensu stricto. However, especially in the species of the L. granulifera-group and the L. longa-group the homologous cirrus IV/2 is present, but inconspicuous because not behind but right of the adoral zone and part of the “cirri left of the amphisiellid median cirral row”. Species included in Lamtostyla (alphabetically arranged basionyms are given): (1) Amphisiella australis Blatterer & Foissner, 1988; (2) Amphisiella elegans Foissner, Agatha & Berger, 2002; (3) Amphisiella procera Foissner, Agatha & Berger, 2002; (4) Amphisiella quadrinucleata Berger & Foissner, 1989; (5) Lamtostyla decorata Foissner, Agatha & Berger, 2002; (6) Lamtostyla granulifera Foissner, 1997; (7) Lamtostyla islandica Berger & Foissner, 1988; (8) Lamtostyla lamottei Buitkamp, 1977; (9) Tachysoma longa Hemberger, 1985; (10) Tachysoma perisincirra Hemberger, 1985; (11) Tachysoma raptans Hemberger, 1985; (12) Uroleptoides vitiphila Foissner, 1987. In addition Amphisiella australis sensu Foissner (1988), which differs distinctly from the type population, is included. Species misplaced in Lamtostyla: The species listed below have been originally assigned or transferred to Lamtostyla. However, new data or distinct differences to the cirral pattern of L. lamottei show that they are very likely misplaced in the present genus. Lamtostyla abdita Foissner, 1997. Remarks: Now Afroamphisiella abdita (p. 377). Lamtostyla edaphoni Berger & Foissner, 1987. Remarks: Now Lamtostylides edaphoni (p. 324). Lamtostyla halophila Foissner, Agatha & Berger, 2002. Remarks: Now Lamtostylides halophilus (p. 336). Lamtostyla hyalina (Berger, Foissner & Adam, 1984) Berger & Foissner, 1987. Remarks: Now Lamtostylides hyalinus (p. 347). Lamtostyla kirkeniensis Berger & Foissner, 1988. Remarks: Now Lamtostylides kirkeniensis (p. 333).
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SYSTEMATIC SECTION
Key to Lamtostyla species Identification of Lamtostyla species is not quite simple. Unfortunately, for some species live data are not available so that the feature presence/absence of cortical granules is only of limited use. Anyhow, the differences between some species are rather sophisticated (e.g., two or three dorsal kineties) so that protargol impregnation is sometimes indispensable. If you cannot identify your specimen/population with the key below, see also Lamtostylides (only one cirrus left of anterior portion of amphisiellid median cirral row), Terricirra (like Lamtostylides, but with spindle-shaped food vacuoles and prominent cortical granules), Afroamphisiella (transverse cirri lacking), and Uroleptoides (amphisiellid median cirral row extends beyond 50% of body length). 1 Body length 80 µm or less (Fig. 37a, 39a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 - Body length usually more than 80 µm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2 One micronucleus between two narrowly spaced macronuclear nodules (Fig. 39a–f); body length usually 60 µm or less. . . . . . Lamtostyla perisincirra (p. 200) - Usually each one micronucleus attached to the 2 macronuclear nodules (Fig. 37a–d); body length usually 60–80 µm. . . . . . . . . . Lamtostyla islandica (p. 196) 3 (1) Four macronuclear nodules (Fig. 34a, 35a, 36a). . . . . . . . . . . . . . . . . . . . . . . 4 - Two macronuclear nodules. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 4 Body slender (Fig. 36a); body length 140–200 µm. . Lamtostyla elegans (p. 192) - Body elongate elliptical (Fig. 34a, 35a); body length usually below 130 µm. . . . 5 5 Usually 1 (rarely 2) buccal cirrus; 3 dorsal kineties (Fig. 34c, d). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lamtostyla vitiphila (p. 187) - Usually 3 (sometimes 2) buccal cirri; 2 dorsal kineties (Fig. 35d, e). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lamtostyla quadrinucleata (p. 190) 6 (3) Four or 5 dorsal kineties . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 - Two or 3 dorsal kineties. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 7 Two transverse cirri; amphisiellid median cirral row composed of about 10 cirri; 4 dorsal kineties (Fig. 30a). . . . . . . . . . . . . . . . . . . . . Lamtostyla lamottei (p. 167) - Five transverse cirri (plus 1 or 2 pretransverse ventral cirri); amphisiellid median cirral row composed of only about 4 cirri; 5 dorsal kineties (Fig. 43a, 44a). . . . 8 8 Body length around 200 µm (Fig. 44a). . . . . . . . . . . . Lamtostyla raptans (p. 221) - Body length around 85–130 µm (Fig. 43a). . . . . . . . . . Lamtostyla longa (p. 218) 9 (6) Cortical granules present (Fig. 40d, 41e, f, n). . . . . . . . . . . . . . . . . . . . . . . . 10 - Cortical granules lacking. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 10 Cortical granules arranged in plaques around dorsal bristles and some granules around cirri; individual granules colourless and only about 0.3 µm across (Fig. 41e–g, n, 42e, f). . . . . . . . . . . . . . . . . . . . . . . . . . . . Lamtostyla decorata (p. 212) - Cortical granules arranged in narrowly spaced rows, leaving blank only cirral and bristle rows; individual nodules colourless and 1–4 µm, usually 2 µm across (Fig. 40d, h–k). . . . . . . . . . . . . . . . . . . . . . . . . . . . Lamtostyla granulifera (p. 206)
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167
11 (9) Body almost vermiform; ratio of body length:width about 9:1 in life (Fig. 33a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lamtostyla procera (p. 183) - Body elongate elliptical; ratio of body length:width about 3–5:1 (Fig. 31a, g, 32a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 12 Usually 2 transverse cirri; buccal cirrus near anterior end of undulating membranes; distance between dorsal kineties 1 and 2 enlarged (Fig. 31a, e, f, h, i). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lamtostyla australis (p. 169) - Usually 5 transverse cirri; buccal cirrus distinctly behind anterior end of undulating membranes; distance between dorsal kineties 2 and 3 enlarged (Fig. 32a, f, g). . . . . . . . . . . . . . . . . . . . . . . . . Amphisiella australis sensu Foissner (1988; p. 179)
Lamtostyla lamottei-group This group comprises all Lamtostyla species, except for L. granulifera, L. decorata, L. longa, and L. raptans. All members have an amphisiellid median cirral row which is shorter than 50% of body length, but consists of more than four cirri.
Lamtostyla lamottei Buitkamp, 1977 (Fig. 30a, b) 1977 Lamtostyla lamottei n. gen. n. spec. – Buitkamp, Acta Protozool., 16: 270, Abb. 14 (Fig. 30a, b; original description; no formal diagnosis provided; type slides are deposited in the Institut für landwirtschaftliche Zoologie, Universität Bonn, Germany). 1983 Lamtostyla Buitkamp, 1977 – Curds, Gates & Roberts, British and other freshwater ciliated protozoa, p. 410, Fig. 240 (redrawing of Fig. 30a; guide to ciliate genera). 1985 Lamtostyla lamottei – Small & Lynn, Phylum Ciliophora, p. 456, Fig. 26 (Fig. 30a; guide to ciliate genera). 1986 Lamtostyla lamottei Buitkamp, 1977 – Dragesco & Dragesco-Kernéis, Faune tropicale, 26: 457, Planche 135A, B (Fig. 30a, b; guide to ciliates of tropical Africa). 1987 Lamtostyla lamottei Buitkamp, 1977 – Berger & Foissner, Zool. Jb. Syst., 114: 217 (brief review of Lamtostyla). 1988 Lamtostyla lamottei Buitkamp, 1977 – Berger & Foissner, Zool. Anz., 220: 122, Fig. 2, Table 1 (Fig. 30a; revision of Lamtostyla). 2001 Lamtostyla lamottei Buitkamp, 1977 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 45 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs).
Nomenclature: Buitkamp (1977) dedicated this species to Maxime Lamotte, Chief of the Station for Tropical Ecology in Lamto, Ivory Coast. Lamtostyla lamottei is the type species of Lamtostyla. Remarks: For discussion of Lamtostyla, see genus section. Lamtostyla lamottei is basically described only after protargol preparations. Thus, some important features, for example, presence/absence of cortical granules, are not known. It is very similar to Lamtostyla australis (Fig. 31e, h), which has, however, fewer dorsal kineties (four in L. lamottei vs. three in L. australis) and a longer amphisiellid median
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SYSTEMATIC SECTION
cirral row (about 25% of body length vs. 38–45%; Buitkamp 1977, Blatterer & Foissner 1988, Foissner 1988, Voß 1992). Thus, a redescription (life data, morphometry, ontogenesis) of L. lamottei is needed to show whether or not further differences exist. Morphology: Body length in life (?) about 100 µm. Body outline with parallel margins, both ends broadly rounded. Two ellipsoidal macronuclear nodules, three micronuclei (Fig. 30b). Contractile vacuole, as is usual, near left cell margin slightly ahead of mid-body (Fig. 30a). Cytoplasmic inclusions, presence/absence of cortical granules, movement not mentioned. Adoral zone relatively short, that is, occupies only about 23% of body length (Fig. 30a), composed of 19 membranelles on average. Cilia of membranelles up to 12 µm long. Paroral and endoral composed of monokinetids, cilia of endoral finer than those of paroral. Cirral pattern as shown in Fig. 30a. Three enlarged frontal cirri in common position. Buccal cirrus slightly behind of anterior end of paroral. Three cirri left of anterior portion of amphisiellid median cirral Fig. 30a, b Lamtostyla lamottei (from Buitrow; anteriormost cirrus likely homologous kamp 1977. Protargol impregnation). Infrawith cirrus III/2. Amphisiellid median cirral ciliature of ventral side and nuclear apparatus, 73 µm. Frontal cirri connected by dotted line; row composed of about 10 cirri, comcirri left of anterior portion of amphisiellid mences near anterior end of right marginal median cirral row circled (anteriormost cirrus row, terminates slightly behind level of = cirrus III/2). Broken lines connect cirri proximal end of adoral zone (about at 26% which very likely originate from same anlage (only shown for anlagen I–IV). ACR = amof body length in specimen illustrated). Prephisiellid median cirral row, AZM = adoral transverse cirri obviously lacking (Fig. zone of membranelles, CV = contractile vacu30a). Two subterminal transverse cirri, ole, FC = frontal cirri, LMR = left marginal about 16 µm long. Right marginal row row, TC = transverse cirri. Page 167. commences near anterior end of amphisiellid median cirral row, composed of 28–35 cirri, terminates, like left row, at level of transverse cirri. Left row commences near proximal end of adoral zone, composed of 24–32 cirri; marginal cirri about 10 µm long, ahead of mid-body composed of 3 × 2
Lamtostyla
169
cilia, behind mid-body usually composed of 2 × 2 cilia. Dorsal cilia about 3 µm long, arranged in four kineties. Caudal cirri lacking. Occurrence and ecology: Very likely confined to terrestrial habitats (Berger & Foissner 1988, Foissner 1987, p. 125; 1998, p. 205). Type locality of L. lamottei is the Plateau du Grand Nord, Ivory Coast, where Buitkamp (1977) discovered it sparsely in soil from the savane brûlée. Not found since then. Buitkamp (1979, p. 230) counted 54 active specimens g-1 dry soil at 15°C and six days after wetting the sample. This abundance corresponds a biomass of 1056 g ha-1 (cell volume about 2.3 × 10-8 cm3; Buitkamp 1979). Lamtostyla lamottei feeds on ciliates and diatoms (Buitkamp 1977). Biomass of 6 10 specimens about 26 mg (Foissner 1987, p. 125).
Lamtostyla australis (Blatterer & Foissner, 1988) Petz & Foissner, 1996 (Fig. 31a–z, Tables 20–22) 1988 Amphisiella australis nov. spec.1 – Blatterer & Foissner, Stapfia, 17: 36, Abb. 10a–f, Tabelle 8 (Fig. 31a–f; original description; the holotype slide [accession number 1989/56; Aescht 2003, p. 381] and a paratype slide [1989/57] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria). 1992 Amphisiella australis Blatterer and Foissner, 1988 – Voß, Europ. J. Protistol., 28: 405, Fig. 1–32, Tables 1–3 (Fig. 31g–z; redescription and cell division). 1994 Amphisiella australis Blatterer & Foissner, 1988 – Eigner & Foissner, J. Euk. Microbiol., 41: 255, Fig. 61 (brief revision). 1996 Lamtostyla australis (Blatterer and Foissner, 1988) nov. comb. – Petz & Foissner, Acta Protozool., 35: 277 (combination with Lamtostyla). 2001 Lamtostyla australis (Blatterer and Foissner, 1988) Petz and Foissner, 1996 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 7 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs).
Nomenclature: No derivation of the name is given in the original description. The species-group name australis (Latin adjective; southern) refers to the continent (southern hemisphere; Australia), where the species was discovered. Remarks: The present species is very well studied. It was originally assigned to Amphisiella mainly because of the cirral row and because it has more than one cirrus left of the anterior portion of the amphisiellid median cirral row. Foissner (1988) found it at various sites outside from Australia. However, these populations differ in several features from the type population and the population described by Voß (1992), strongly indicating that they do not belong to L. australis (details see Amphisiella australis sensu Foissner 1988). By contrast, the European population studied by Voß (1992) agrees very well with the type material. 1
Blatterer & Foissner (1988) provided the following diagnosis: In vivo etwa 90–130 × 30–40 µm große, farblose Amphisiella mit 2 Makronucleus-Teilen, 3 Dorsalkineten sowie durchschnittlich 2 Transversalcirren und 22 adoralen Membranellen.
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Fig. 31a–f Lamtostyla australis (from Blatterer & Foissner 1988. a–d, from life; e, f, protargol impregnation). a: Ventral view, 98 µm. b: Feeding specimen. c: Shape variant in dorsal view. d: Ventral view of a specimen of population II. e, f: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 107 µm. Arrowhead in (e) marks buccal cirrus, arrow denotes cirrus III/2. Frontal cirri connected by dotted line, cirri left of anterior portion of ACR circled. Note wide distance (bipolar arrow in f) between kineties 1 and 2, and widely spaced bristles (asterisk) in posterior portion of kinety 1. ACR = amphisiellid median cirral row, LMR = left marginal row, MA = macronuclear nodule, MI = micronucleus, P = paroral, RMR = right marginal row, TC = transverse cirri, 1–3 = dorsal kineties. Page 169.
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171
Table 20 Origin of the frontal-ventral-transverse cirri anlagen I–VI in Lamtostyla australis (after Voß 1992; terminology of some cirri modified). Daughter cell Proter
Anlagen I II III IV V, VI
Opisthe
I–III IV–VI
Parental structures associated with origin of the primordia Undulating membranes Buccal cirrus Cirrus III/2 (= cirrus behind right frontal cirrus) cirri left of anterior portion of amphisiellid median cirral row (originated from anlage IV in previous generation) Cirri of amphisiellid median cirral row (this row originated from anlagen V [posterior portion] and VI [anterior portion] in previous generation) Oral primordium Cirri of amphisiellid median cirral row and partially from oral primordium
Table 21 Number of new cirri formed from the frontal-ventral-transverse cirri anlagen I–VI in Lamtostyla australis (after Voß 1992) Anlage a I II III IV V VI
mean
M
SD
SE
CV
Min
Max
n
1.3 3.0 3.0 3.5 7.9 8.6
1.0 3.0 3.0 4.0 8.0 9.0
0.4 0.5 0.5 0.6 1.0 1.1
0.1 0.1 0.1 0.1 0.2 0.3
30.8 16.7 16.7 15.0 12.7 12.2
1.0 2.0 2.0 3.0 6.0 7.0
2.0 4.0 4.0 5.0 10.0 10.0
19 19 17 18 20 15
a
Data based on protargol-impregnated specimens. CV = coefficient of variation in %, M = median, Max = maximum value, mean = arithmetic mean, Min = minimum value, n = number of individuals investigated, SD = standard deviation, SE = standard error of arithmetic mean.
Petz & Foissner (1996) transferred the present species from Amphisiella to Lamtostyla because the stomatogenesis commences near the transverse cirri. I basically agree with this transfer although it is uncertain because the ontogenesis of L. lamottei, type of the genus, is not known. Eigner (1999, p. 46) did not accept the transfer to Lamtostyla. Lamtostyla australis is difficult to distinguish from the type species; the most important differences are the length of the amphisiellid median cirral row (38–46% of body length vs. about 26%), the number of cirri forming the amphisiellid median cirral row (around 12–14 vs. about 10), and the number of dorsal kineties (three vs. four). Morphology: The following description is based on the original description. For morphometric data on the European population studied by Voß (1992), see Table 22. Body size about 90–130 × 30–40 µm in life, body length:width ratio of type population 3.5:1 on average in protargol preparations (Table 22). Body outline elongate elliptical, slightly sigmoidal; anterior body portion of type population conspicuously pointed and front end narrowly rounded, margins of posterior portion slightly
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Lamtostyla
173
converging, rear end broadly rounded (Fig. 31a–c); in a second population, the apical portion is not so distinctly pointed (Fig. 31d). Body very flexible, about 30% contractile, dorsoventrally slightly flattened. Distinctly S-shaped when grabbing the prey (Fig. 31b). Two macronuclear nodules in ordinary position, that is, slightly left of midline, one ahead of and one behind mid-body (Fig. 31a, e, g); individual nodules ellipsoidal, with many globular to irregular-shaped chromatin bodies. Micronuclei globular to slightly ellipsoidal; usually each one attached to a macronuclear nodule. Contractile vacuole near left cell margin about in mid-body, during diastole with short collecting canals (Fig. 31a, c, g). Pellicle colourless, cortical granules lacking. Cytoplasm with a moderately high number of colourless, greasily shining globules 1–3 µm in diameter. Food vacuoles 6–8 µm across. Movement rapid, without peculiarities. Adoral zone occupies about 20–33% of body length in life, 24–30% on average in protargol preparations, composed of an average of about 22 membranelles of ordinary fine structure (Fig. 31a, e, Table 22). Largest membranelles about 5 µm wide in life. Buccal field narrow and deep. Pharyngeal fibres inconspicuous. Undulating membranes of about equal length and arranged in parallel, each membrane likely composed of two rows of basal bodies; endoral commences slightly more anteriorly than paroral. Invariable three frontal cirri arranged in oblique row (Fig. 31e, Table 22). Buccal cirrus near anterior end of paroral. Usually four cirri left of anterior portion of amphisiellid median cirral row, namely cirrus III/2 (usually slightly enlarged) and three cirri originating from anlage IV (details of origin see Fig. 31w); rarely up to 10 cirri arranged in up to three rows. Amphisiellid median cirral row commences close to distal end of adoral zone, terminates on average at 38–46% of body length in protargol preparations (Table 22). According to original description, frontoterminal cirri lacking; however, cell division shows that the anterior portion of the amphisiellid median cirral row is homologous to the frontoterminal cirri (see cell division and general section). Transverse cirri about 15 µm long, project by about half their length beyond rear body end. Marginal cirri and cirri of amphisiellid median row about 10 µm long. Right marginal row commences about at level of buccal cirrus, ends slightly subterminal; left row begins slightly ahead of level of proximal end of adoral zone, terminates subterminal so that marginal rows distinctly separated posteriorly.
b
Fig. 31g–n Lamtostyla australis (from Voß 1992. g, from life; h–n, protargol impregnation; g–i, interphase; j–n, cell division). g: Ventral view, 95 µm. h, i: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 105 µm. Note increased distance between dorsal kineties 1 and 2 (bipolar arrow) and widely spaced bristles in posterior portion of kinety 1 (asterisk). These are two important differences to Amphisiella australis sensu Foissner (1988). j–l: Origin of oral primordium near/from transverse cirri, j = 107 µm. m, n: Infraciliature of ventral side of early dividers, m, n = 85 µm. Arrow in (m) marks rearmost two cirri of amphisiellid median cirral row, which form a primordium. ACR = amphisiellid median cirral row, E = endoral, P = paroral, RE = replication band, 1–3 = dorsal kineties. Page 169.
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SYSTEMATIC SECTION
Lamtostyla
175
Fig. 31s, t Lamtostyla australis (from Voß 1992. Protargol impregnation). Infraciliature of ventral and dorsal side and nuclear apparatus of a middle divider, 73 µm. Both the ventral and dorsal morphogenesis proceed rather unspectacularly. I, VI = frontal-ventral-transverse cirri primordia. Page 169.
Dorsal cilia about 2 µm long, arranged in three more or less bipolar kineties. Distance between kinety 1 and 2 larger than that between kineties 2 and 3; bristles of kinety 1 rather widely spaced in posterior portion. Caudal cirri lacking (Fig. 31f, i). Cyst: This stage of life cycle is described by Voß (1992). It is spherical, in life 30 µm across on average (M = 29.9; SD =1.7; SE = 0.3; CV = 5.7%; Min = 26; Max = 32; n = 25), and has a smooth wall covered with a thin (2–4 µm) mucous layer to which some particles are sometimes attached (micrographs see Fig. 22, 23 in Voß 1992). Cell division (Fig. 31j–z): Voß (1992) studied ontogenesis of Lamtostyla australis in great detail.
b
Fig. 31o–r Lamtostyla australis (from Voß 1992. Protargol impregnation). Infraciliature of ventral side of middle dividers, o, p = 130 µm, q = 104 µm, r = 115 µm. Arrow in (r) marks separation of primary primordia. I–VI = frontal-ventral-transverse cirri primordia I–VI. Page 169.
176 SYSTEMATIC SECTION Fig. 31u–w Lamtostyla australis (from Voß 1992. Protargol impregnation. Parental structures white, new black). Infraciliature of ventral and dorsal side and nuclear apparatus of late dividers, u = 67 µm, v = 62 µm, w = 99 µm. Broken lines in (w) connect opisthe’s cirri which originate from the same anlage. Arrow in (w) marks anteriorly migrating cirri (except for transverse cirrus) of anlage VI to form the anterior portion of the amphisiellid median cirral row; thus, the anterior portion is homologous to the frontoterminal cirri of, for example, the urostyloids and oxytrichids. Dorsal side of specimen shown in (w), see Fig. 31x. AZM = parental adoral zone of membranelles, MA = fused macronucleus, MI = pre-division micronucleus. Page 169.
Lamtostyla 177
Fig. 31–z Lamtostyla australis (from Voß 1992. Protargol impregnation. Parental structures white, new black). Infraciliature of dorsal and ventral side and nuclear apparatus of late and very late dividers, x = 99 µm (same specimen as shown in w), y = 130 µm, z = 114 µm. Broken lines in (y) mark cirri which originate from same anlage. MA = macronuclear nodule, MI = dividing micronucleus, 1–3 = dorsal kineties. Page 169.
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SYSTEMATIC SECTION
Stomatogenesis commences with the formation of an oral primordium. In 43% of specimens the two transverse cirri dedifferentiate; in 40% the primordium originates de novo in close contact to the left transverse cirrus, and in 16% the development is associated with the dedifferentiation of the left transverse cirrus (Fig. 31j–l). The oral primordium forms into a long small field with a broadened anterior end (Fig. 31m, n). The field becomes wider and, as is usual, the formation of new membranelles proceeds from anterior to posterior. Right of the new adoral zone, the anlage for the undulating membranes originate (Fig. 31o–r). The old adoral zone is, as in most hypotrichs, retained. The anterior sections of the parental undulating membranes reorganise simultaneously, that is, show some basal body proliferation at their anterior ends; later, this anlage produces, as is usual, the left frontal cirrus. Development of cirral primordia. The last two or three cirri of the amphisiellid median cirral row dedifferentiate into small groups of basal bodies when the oral primordium of the opisthe begins to elongate (Fig. 31m). This anlage becomes larger and later contacts the anterior end of the oral primordium. Six anlagen are formed: three originate from the oral primordium and three from the primordium of the ventral row (Fig. 31n–r). In the meantime, the buccal cirrus, cirrus III/2, the cirri left of the anterior portion of the amphisiellid median cirral row, and further cirri of the amphisiellid median cirral row modify to anlagen (Fig. 31o, p). The three primordia of the proter and the six anlagen of the opisthe form long anlagen of loosely arranged basal body pairs. The anlagen II, III, and IV of proter and opisthe join, so that five long primordia occur. Only the anlage I of the proter and the opisthe do not fuse (Fig. 31q). However, the primordia separate very soon, so that a set of six anlagen (I–VI) each for proter and opisthe is formed (Fig. 31r). Voß (1992) summarised the origin of the anlagen in Table 20. Development of marginal rows and dorsal kineties. Marginal row primordia and dorsal kinety anlagen are formed, as is usual for most hypotrichs, at two levels within the parental rows and kineties, that is, dorsomarginal rows and dorsal kinety fragmentation are lacking (Fig. 31s–z). Differentiation of new cirri. This process shows no peculiarities and thus the reader is mainly referred to Fig. 31s, u–w, y. The data show that the amphisiellid median cirral row is formed by the anlagen V (posterior portion of row) and VI (anterior portion which is homologous to the frontoterminal cirri). Occasionally, the alignment is not perfect, that is, the two portions forming the row are recognisable by a slight interruption. Only these two rightmost anlagen form transverse cirri, a feature also known from other genera, for example, Urosomoida spp. (for review see Berger 1999). Note that the anlagen I–IV usually form more cirri than present in the interphasic specimens (Table 21), indicating that one or two cirri are resorbed during the final stages of ontogenesis (Fig. 31u–w). Nuclear apparatus. The nuclear apparatus divides plesiomorphically, that is, each macronuclear nodule shows a reorganisation band, usually at its proximal end. The two nodules fuse to a single mass and later divide (Fig. 31j, t, v, y, z).
Lamtostyla
179
Occurrence and ecology: Confined to terrestrial habitats (Foissner 1998, p. 205). Type locality of Lamtostyla australis is the South Para Reservoir near Adelaide, Australia, where Blatterer & Foissner (1988) discovered it in the slightly mouldy litter (upper soil layer; 0–5 cm; pH 5.1; collected by W. Foissner on 17.02.1987) of a secondary pine forest. Population II is likely from the bush in Brisbane Water National park about 50 km north of Sydney, where Blatterer & Foissner (1988) found it in the upper soil layer with much black litter and some sand (pH 4.2; collected by H. Blatterer on 23.10.1986). A third population, although not studied in detail by Blatterer & Foissner (1988), was recorded from the upper soil layer (litter from small bushes, mosses, and lichens) near Lake Dobson in the Mt. Fields National Park, Tasmania. Voß (1992) found his population in an infusion of an air-dried moss sample collected from a dune near the lighthouse of the East-Frisian isle of Norderney, Germany in September 1990. Voß isolated and cloned several specimens and used noncarbonated mineral water (“Vittel”) as culture medium. He added few drops of a concentrated mixture containing a suspension of baker’s yeast and Chlorella algae as food. In these cultures encystment took place every 2–3 days. Excystment was induced by transferring the concentrated cysts in fresh culture medium or distilled water; the cells hatched after 8–10 hours. Foissner (1994, p. 23) recorded L. australis from the type locality of Bryometopus hawaiinesis, that is, grassland soil near the entrance to the sandalwood trail in the Volcano National Park, Big Island, Hawaiian Archipelago (155°20'W 19°26'N). Foissner et al. (2002, p. 60) recorded it several times in terrestrial habitats from Namibia. Lamtostyla australis feeds on heterotrophic flagellates (Polytoma sp.), ciliates (Colpoda steinii, Cyclidium sp.), coccale green algae, bacteria, and fungal spores (Blatterer & Foissner 1988). Biomass of 106 specimens about 50 mg (Foissner 1998, p. 205).
Amphisiella australis Blatterer & Foissner, 1988 sensu Foissner, 1988 (Fig. 32a–i, Table 22) 1988 Amphisiella australis Blatterer & Foissner, 1988 – Foissner, Stapfia, 17: 113, Abb. 9a–i, Tabelle 6 (Fig. 32a–i; description of three African populations and one population from Singapore; see remarks; voucher slides are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria).
Remarks: Foissner (1988) found Lamtostyla australis at various sites outside of Australia. Two populations from Africa differ distinctly from the Australian type population in some morphometric features, for example, number of transverse cirri (usually five vs. usually two) and length of amphisiellid median cirral row (50% of body length vs. 38–45%; Table 22). Furthermore, the undulating membranes are distinctly curved (against only slightly curved) and the buccal cirrus is distinctly be-
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SYSTEMATIC SECTION
Fig. 32a–e Amphisiella australis sensu Foissner (1988; from life. a, b, population I; c, population II; d, e, Singapore and African specimen). a: Ventral view of a representative specimen, 160 µm. b–e: Shape variants in dorsal (b, c, e) and ventral view (d). For details on taxonomy, see text. CV = contractile vacuole. Page 179.
hind the anterior end of the paroral (against at anterior end; differences in dorsal ciliature, see morphology). Thus, the African specimens are reminiscent of Uroleptoides binucleatus, which, however, has cortical granules (Foissner et al. 2002, their Table 131). Foissner (1988) discussed that his populations possibly do not belong to Amphisiella australis (= Lamtostyla australis in present book) and therefore provided a rather detailed description. I am sure that the populations studied by Blatterer & Foissner (1988) and Foissner (1988) are not conspecific and thus do not include Foissner’s (1988) data in the L. australis part, but keep them separate. Further data should be awaited for a formal separation at subspecies or species level. Since the amphisiellid median cirral row is about 50% of body length in Foissner’s (1988) populations, Amphisiella australis sensu Foissner (1988) is also included in the Uroleptoides key. Morphology: Foissner (1988) described four non-Australian populations (see occurrence section). He studied two Kenyan populations in detail, which differ not
Lamtostyla
181
Fig. 32f–i Amphisiella australis sensu Foissner (1988; protargol impregnation. f–i, population I). f, g: Infraciliature of ventral and dorsal side and nuclear apparatus of representative, interphasic specimen, 122 µm. Arrow in (f) marks cirrus III/2. Bipolar arrow shows the wide distance between kineties 2 and 3; asterisk marks the normally spaced bristles in the posterior portion of kinety 1; both features separate, besides other features (see text), Amphisiella australis sensu Foissner 1988 from the type population (Fig. 31a–f). h, i: Infraciliature of ventral side of a middle and a late divider, h = 143 µm, i = 86 µm. Note that the amphisiellid median cirral row is formed by the cirri of the anlagen V (posterior portion) and VI (anterior portion; homologous to the frontoterminal cirri). ACR = amphisiellid median cirral row, AZM = adoral zone of membranelles, E = endoral, LMR = left marginal row, MA = macronuclear nodule, MI = micronucleus, P = paroral, RMR = right marginal row, TC = leftmost transverse cirrus, I–VI = frontal-ventral-transverse cirri anlagen, 1–3 = dorsal kineties. Page 179.
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SYSTEMATIC SECTION
only from the Australian material (see remarks), but also from each other. Thus, he concluded that they are local variants. The description below is based on these two populations, unless otherwise indicated. Body size 130–160 × 30–40 µm in life. Body outline very slenderly elliptical, posteriorly usually slightly converging, at level of adoral zone often distinctly curved leftwards (Fig. 32a, b), which is, besides the cortical granulation, a difference to Uroleptoides magnigranulosus. However, elliptical and posteriorly rightwards curved specimens also occur (Fig. 32c–e). Difficult to fix and therefore much wider in protargol preparations than in life. Not to only slightly flattened dorsoventrally; specimens from Singapore slightly twisted about main body axis. Body very flexible, population I (see occurrence) contractile by about 20% of body length. Invariably two macronuclear nodules about in mid-body slightly left of midline. Contractile vacuole, as is usual, near left cell margin slightly ahead of or about in midbody (Fig. 32d, c, e). Micronuclei of the Singapore specimens 6 × 4 µm, that is, conspicuously large. Cortical granules lacking. One Kenyan population with colourless granules (1 µm across) in buccal field. Cytoplasm colourless, often packed with 1–10 µm-sized, globular to club-shaped, greasy-shining inclusions and 1–2 µmsized, lens-shaped crystals. Cirral pattern very similar to that of Uroleptoides magnigranulosus (Fig. 32f, Table 22). The most important differences to the Lamtostyla australis populations described by Blatterer & Foissner (1988) and Voß (1992) are (i) the higher number of transverse cirri (on average five vs. on average two); (ii) the shape of the undulating membranes (distinctly curved and optically intersecting vs. slightly curved and almost in parallel); (iii) the position of the buccal cirrus (distinctly behind anterior end of paroral vs. right of anterior end); (iv) length of amphisiellid median cirral row (50% vs. 38–45%; Table 22); and (v) details of dorsal ciliature (kineties 2 and 3 distinctly separated and bristles of kinety 1 more or less equally spaced [Fig. 32g] vs. kineties 1 and 2 slightly separated and bristles of kinety 1 widely spaced in middle and posterior portion [Fig. 31f, i]). One of the three African populations invariably with only three cirri (cirrus III/2 included) left of anterior portion of amphisiellid median cirral row (Foissner 1988). Population from Singapore with 60–70 cirri (n = 2) each in left and right marginal row and with 48 (n = 1) cirri in amphisiellid median cirral row, that is, number of cirri distinctly higher than in African and Australian populations (Table 22). In population II (see occurrence) two out of 13 specimens with four dorsal kineties. Buccal field in all populations deep, narrow, and anteriorly distinctly curved leftwards. Cell division (Fig. 32h, i): The rear portion of the amphisiellid median cirral row, the cirri left of the anterior portion of the amphisiellid median cirral row, the cirrus III/2, the buccal cirrus, and the undulating membranes are modified to primordia. The amphisiellid row originates, as is usual, from two anlagen (V and VI): the cirri of anlage V form the posterior portion, those of anlage VI, which are homologous to the frontoterminal cirri, form the anterior portion (Fig. 32i).
Lamtostyla
183
Occurrence and ecology: Terrestrial. Foissner (1988) studied four populations. Population I is from the mouldy litter (pH 6.7) of a leaf tree from near the Naivasha railway station near Lake Naivasha, Kenya. Population II is from a strongly saline soil (pH 6.9) grown with brown algae and grass near a geyser near the shore of Lake Baringo, Kenya. The third African population is also from Kenya, but no details are provided. The Singapore population is from the Bukit Timah National Park. Feeds on small ciliates (Cyclidium sp., Colpoda cf. aspera, Sathrophilus muscorum, Drepanomonas sp.), heterotrophic flagellates, and coccale green algae. Specimens of population I move rapidly, those of the other populations slowly, showing great flexibility (Foissner 1988).
Lamtostyla procera (Foissner, Agatha & Berger, 2002) comb. nov. (Fig. 33a–g, Table 22) 2002 Amphisiella procera nov. spec.1 – Foissner, Agatha & Berger, Denisia, 5: 670, Fig. 149a–g, Table 132 (Fig. 33a–g; original description; one holotype slide [accession number 2002/380], one paratype slide [2002/381], and one voucher slide [2002/382] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria).
Nomenclature: The species-group name procer·us, -a, -um (Latin adjective [m; f; n]; elongate) refers to the slender body shape (Foissner et al. 2002). Remarks: In the absence of ontogenetic data, which – according to Petz & Foissner (1996) – separate Amphisiella and Lamtostyla, we preliminarily assigned the present species to Amphisiella as defined by Eigner & Foissner (1994). Now we know that Amphisiella is a marine group, which has, inter alia, a more or less complete set of pretransverse ventral cirri and transverse cirri and more than three dorsal kineties. By contrast, terrestrial species likely belong to the Uroleptoides/Lamtostyla group. The type species of both genera are not known in detail and therefore the separation is preliminarily only pragmatic. Amphisiella procera is transferred to Lamtostyla because the amphisiellid median cirral row is less than 50% of body length (for details, see genus section of Uroleptoides). Lamtostyla procera is most similar to Uroleptoides binucleatus (Fig. 50a–h, 51a–i). It differs from this species in the location of the buccal cirrus (near anterior end of paroral vs. near posterior end or in middle portion), the cortical granules (absent vs. present), the number of dorsal kineties (two vs. three), and the length of the amphisiellid median cirral row (1/3 of body length vs. ≥2/3 or more). Further similar, but comparatively very distinct species are Uroleptoides magnigranulosus (with conspicuous cortical granules; Fig. 52a–l, 53a–e) and Lamtostyla australis, a 1
Foissner et al. (2002) provided the following diagnosis: Size about 170 × 18 µm in vivo, that is, almost vermiform with posterior fifth narrowed tail-like. 2 macronuclear nodules. Amphisiellid median cirral row (ACR) ends above mid-body, composed of an average of 15 cirri. On average 20 adoral membranelles, 55 cirri in left and 57 in right marginal row, 3 cirri left of anterior end of ACR, 1 buccal cirrus, 4 transverse cirri, and 2 dorsal kineties which extend along right body margin in posterior body portion.
184
SYSTEMATIC SECTION
Lamtostyla
185
smaller (length 90–160 µm), elongate ellipsoidal species with three dorsal kineties (e.g., Fig. 31a–f). In life, Lamtostyla procera is identified by the following combination of features (Foissner et al. 2002): very slender body with distinct tail, two macronuclear nodules, very inconspicuous transverse cirri, lack of cortical granules and postperistomial ventral cirri. Foissner et al. (2002) found two populations, which agree very well in all features. Only the average number of cirri left of the anterior end of the amphisiellid median cirral row is different, namely, three in the type population and four in the other population. Morphology: Body size 130–210 × 15–25 µm in life, length:width ratio about 9:1 in vivo and 6.1:1 on average in protargol preparations (Table 22). Body outline almost vermiform with posterior fifth narrowed tail-like in life; tail very fragile and thus rarely preserved in protargol preparations (Fig. 33a, b, e, g, Table 22); trunk dorsoventrally flattened up to 1.5:1. Body extremely flexible, but acontractile; often twisted about main body axis by half a turn. Usually two macronuclear nodules slightly left of midline; individual nodules elongate ellipsoidal, often dumb-bellshaped or of irregular outline, with many medium-sized and small chromatin bodies. Micronuclei difficult to recognise in life and protargol preparations, attached to macronuclear nodules, conspicuously small (Fig. 33a, c, d, g, Table 22). Contractile vacuole slightly ahead of mid-body at left cell margin, with two collecting canals during diastole. Cortical granules lacking. Cytoplasm colourless with many lipid droplets 1–3 µm across. Food vacuoles about 7 µm across. Moves slowly and rather clumsily. Adoral zone occupies only 11–18%, on average 15% of body length, of usual shape and structure, composed of 20 membranelles on average (Table 22). Buccal cavity deep and moderately wide; anterior portion of right cavity margin thickened, posterior forms prominent lip covering proximal portion of adoral zone. Exact structure and arrangement of undulating membranes not clearly recognisable; paroral and endoral slightly to distinctly curved, almost parallel to each other, paroral cilia about 5 µm long in life. Pharyngeal fibres clearly recognisable in life and after protargol impregnation, of ordinary length and structure, extend obliquely backwards (Fig. 33a, b, d, g). Cirral pattern rather constant, number of cirri of usual variability (Fig. 33a–e, g, Table 22). All cirri about 8 µm long in life, fine because usually composed of two or
b
Fig. 33a–f Lamtostyla procera (type population from Foissner et al. 2002. a, from life; b–f, protargol impregnation). a: Ventral view of a representative specimen, 190 µm. Arrow marks contractile vacuole. b, c: Infraciliature of ventral and dorsal side and nuclear apparatus of holotype specimen, 147 µm. Arrow marks rearmost cirrus of group left of anterior end of amphisiellid median cirral row. Right frontal cirrus and cirrus behind (= cirrus III/2) are connected by broken line. d: Ventral infraciliature of a specimen with a long amphisiellid median cirral row, which is composed of two slightly overlapping segments (arrow). e: Posterior body portion showing inconspicuous transverse cirri, which are easily misinterpreted as marginal cirri. f: Most cirri are composed of four or two cilia only. ACR = amphisiellid median cirral row, AZM = adoral zone of membranelles, BC = buccal cirrus, FC = right frontal cirrus, MA = macronuclear nodules, P = paroral, RMR = right marginal row, TC = transverse cirri, 1, 2 = dorsal kineties. Page 183.
186
SYSTEMATIC SECTION four cilia only (Fig. 33f). Frontal cirri slightly to distinctly enlarged, right cirrus, as is usual, at distal end of adoral zone. Buccal cirrus slightly behind anterior end of undulating membranes. Type population usually with three, specimens from Namibian site (9) usually with four cirri left of anterior portion of amphisiellid median cirral row (Fig. 33b, d, g) which begins close to right frontal cirrus and terminates at 30% of body length on average, composed of two segments of unequal length, often slightly overlapping at level of buccal vertex; anteriormost cirrus of amphisiellid median cirral row often slightly enlarged (Fig. 33d, g). Postperistomial cirrus lacking. Transverse cirri inconspicuous both in life and after protargol impregnation because of same size as marginal cirri and often close to marginal rows. Marginal rows end slightly to distinctly subterminally, right row extends onto dorsolateral surface anteriorly; left row commences left of proximal end of adoral zone. Dorsal bristles about 3 µm long in life, arranged in two sparsely ciliated rows. Both rows commence subapically left of body midline and course obliquely backwards ending at right posterior margin of cell. Caudal cirri absent (Fig. 33c). Occurrence and ecology: Lamtostyla procera is very likely confined to terrestrial habitats to which it is welladapted with its slender body (Foissner et al. 2002, p. 50). The type locality is in the surroundings of the Gariganus Guest Farm, about 50 km north of the town of Keetmanshoop (Namibia), in the dwarf shrub savanna (26°25'S 18°20'E), where it occurred sparsely in a soil sample (pH 5.1) under Aloe dichotoma (Quivertree). We also found it in the southern Namib Desert and succulent steppe, about 50 km east of the town of Lüderitz, at main road traversing the Namib Desert (26°40'S 15°40'E), where it oc-
Fig. 33g Lamtostyla procera (from Foissner et al. 2002. Protargol impregnation). Infraciliature of ventral side and nuclear apparatus of a specimen from Namibian site (9), 158 µm. Frontal cirri connected by dotted line, cirri left of anterior end of amphisiellid median cirral row circled. Right frontal cirrus and cirrus III/2 originate from the same anlage and are therefore connected by a broken line. AZM = distal end of adoral zone of membranelles, MA = macronuclear nodule, MI = micronucleus, RMR = rear end of right marginal row, TC = transverse cirri, 2 = dorsal kinety 2 (= right dorsal kinety). Page 183.
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curred in litter, mud, and sand of a flat, dried puddle near the road (pH 6.2). Abundance was very low in both cultures. Feeds on sporulating bacteria (Foissner et al. 2002).
Lamtostyla vitiphila (Foissner, 1987) comb. nov. (Fig. 34a–d, Table 22) 1987 Uroleptoides vitiphila nov. spec.1 – Foissner, Zool. Beitr., 31: 201, Abb. 5a–d, Tabelle 1 (Fig. 34a–d; original description; the holotype slide [accession number 1988/141] and three paratype slides [1988/132–134] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; Aescht 2003, p. 399). 1988 Uroleptoides vitiphilus nom. corr. – Foissner, Stapfia, 17: 113 (correction of name). 1988 Amphisiella vitiphila (Foissner, 1987) nov. comb. – Foissner, Stapfia, 17: 113 (combination with Amphisiella). 1994 Amphisiella – Eigner & Foissner, J. Euk. Microbiol., 41: 259, Fig. 61 (Fig. 34c, d; brief revision of the amphisiellids). 2001 Amphisiella vitiphila (Foissner, 1987) Foissner, 1988 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 97 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs).
Nomenclature: No derivation of the name is given in the original description. The species-group name vitiphil·us, -a, -um (m; f; n) is a composite of viti (likely from Viti Levu; see occurrence) and philos (Greek; loving) and refers to the site (Viti Levu, Fiji Islands) where the species was discovered. Remarks: The present species was established in Uroleptoides, which was classified in the Holostichidae by Foissner (1987a). Somewhat later, Foissner (1988) synonymised – in accordance with Jankowski (1979) – Uroleptoides with Amphisiella and therefore had to transfer the present species to Amphisiella. However, the cirral pattern of the present species is more similar to that of Lamtostyla lamottei, type of Lamtostyla, than to that of Amphisiella capitata, type of Amphisiella. Further, the present transfer based on the morphology agrees also with the habitat restriction (Amphisiella – marine vs. Lamtostyla – terrestrial). Thus, I transfer U. vitiphilus to Lamtostyla. Lamtostyla vitiphila closely resembles L. quadrinucleata (Fig. 35a–e) which, however, has 2–3 (vs. one) buccal cirri, but only two (vs. three) dorsal kineties. Since both features are usually difficult to recognise in life, protargol preparations should be made. Lamtostyla australis has more or less the same ventral and dorsal ciliature as L. vitiphila, but only two (vs. four) macronuclear nodules (Fig. 31e, f). Hemiamphisiella quadrinucleata (Fig. 62a–e) has the same type of nuclear apparatus, but differs from the present species by the presence of a postperistomial cirrus,
1
Foissner (1987) provided the following diagnosis: In vivo etwa 90–120 × 30–50 µm große, farblose Uroleptoides mit 4 Makronucleus-Teilen, 2 Transversalcirren, 3 Dorsalkineten und durchschnittlich 21 adoralen Membranellen.
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Fig. 34a–d Lamtostyla vitiphila (from Foissner 1987a. a, b, from life; c, d, protargol impregnation). a: Ventral view of a representative specimen, 104 µm. b: Right lateral view showing dorsoventral flattening. c, d: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 99 µm. Arrow marks cirrus (= cirrus III/2) behind right frontal cirrus; frontal cirri connected by dotted line; cirri left of anterior end of amphisiellid median cirral row circled by dotted line. ACR = amphisiellid median cirral row, BC = buccal cirrus, MA = macronuclear nodule, MI = micronucleus, P = paroral, RMR = right marginal row, TC = transverse cirri, 1–3 = dorsal kineties. Page 187.
caudal cirri, and cortical granules and the lack of transverse cirri (vs. present in L. vitiphila). Morphology: The type population is described first (Foissner 1987a), followed by some additional observations from a Costa Rican population. Body size 90–120 × 30–50 µm in life, body length:width ratio in protargol preparations 2.4:1 on average (Table 22). Body outline oblong, right margin slightly concave in mid-body, left margin distinctly convex behind adoral zone of membranelles; anterior body end conspicuously narrowed, posterior end broadly rounded (Fig. 34a). Body flattened 2:1 dorsoventrally (Fig. 34b), likely soft and flexible. Usually four macronuclear nodules, longitudinally arranged in pairs left of midline; individual nodules with a large central and several small peripheral chromatin bodies. Micronuclei globular, usually each one attached to a macronuclear nodule.
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Contractile vacuole about in mid-body near left cell margin, with two long collecting canals. Pellicle colourless, conspicuous cortical granules lacking. Food vacuoles large, often irregularly shaped. Movement moderately rapid, without peculiarities. Adoral zone occupies 20–25% of body length, average length in protargol preparations about 25% (Table 22), composed of an average of 21 membranelles of ordinary fine structure. Bases of largest membranelles about 6 µm wide in life. Buccal field small and flat. Undulating membranes rather short, of equal length, begin at same level, slightly curved and optically intersecting in anterior half (Fig. 34a, c). Cirral pattern lamtostylid (Fig. 34c, Table 22). Most cirri about 12 µm long. Frontal cirri moderately enlarged, arranged in slightly oblique pseudorow. Buccal cirrus somewhat behind anterior end of undulating membranes. Cirrus III/2, that is, cirrus behind right frontal cirrus, about at level of buccal cirrus. Usually three cirri (cirrus III/2 included) left of anterior portion of amphisiellid median cirral row. Amphisiellid median cirral row commences about at level of buccal cirrus, extends, as is usual, slightly obliquely to cell midline, terminates at 42% of body length on average (Table 22); distance among cirri becomes wider from anterior to posterior (Fig. 34c). Postperistomial cirrus lacking. Usually two, rarely only one slightly subterminal transverse cirrus. Right marginal row commences about at level of buccal cirrus, ends slightly subterminal. Left marginal row commences close to proximal end of adoral zone, ends – like right row – slightly subterminal so that rows are distinctly separated posteriorly. Dorsal bristles short (about 2–3 µm; estimated from Fig. 34d), arranged in three kineties; kinety 1 (incorrectly designated as right kinety in original description) slightly shortened anteriorly and posteriorly (Fig. 34d). Caudal cirri lacking. Additional observations from a Costa Rican population (kindly supplied by W. Foissner): body size in life about 100 × 30 µm; macronuclear nodules about 15 × 8 µm, micronuclei about 2.5 µm across in life, one between each two macronuclear nodules which form a pair; contractile vacuole distinctly ahead of mid-body; cytoplasm colourless; cortical granules and cytoplasmic crystals lacking; moderately many greasily shining globules 1–3 µm across; food vacuoles 7 µm across; rapidly gliding; two transverse cirri; dorsal bristles 3–4 µm long; caudal cirri lacking. Occurrence and ecology: Likely confined to terrestrial habitats (Foissner 1987, 1998). Type locality of L. vitiphila is a rain forest near Suva, Viti Levu, Fiji Islands, where Foissner (1987a) found it with very low abundance. Further records: soil (pH 4.8) from trees with epiphytes from the Braulio Carrillo National Park, Costa Rica (Foissner 1997, p. 320; some morphological data of this population see above); mud from dry rock-pools along the stream in the Aub Canyon (23°55'S 16°15'E) in the surroundings of the Büllsport Guest Farm in the escarpment of the southern Namib Desert (Foissner et al. 2002, p. 58). Lamtostyla vitiphila feeds on ciliates and flagellates (Foissner 1987a) and cysts of amoebas (W. Foissner, unpublished). Biomass of 106 specimens 92 mg (Foissner 1987, p. 128; 1998, p. 199).
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Lamtostyla quadrinucleata (Berger & Foissner, 1989) comb. nov. (Fig. 35a–e, Table 22) 1989 Amphisiella quadrinucleata nov. spec.1 – Berger & Foissner, Bull. Br. Mus. nat. Hist., 55: 32, Fig. 33–37, Table 8 (Fig. 35a–e; original description; the holotype slide [accession number 1988:2:1:3] and the paratype slide [1988:2:1:4] are deposited in the British Museum of Natural History, London, England). 2001 Amphisiella quadrinucleata Berger and Foissner, 1989 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 8 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Amphisiella quadrinucleata – Lynn & Small, Phylum Ciliophora, p. 454, Fig. 42A, B (Fig. 35d, e; guide to ciliate genera).
Nomenclature: No derivation of the name is given in the original description. The species-group name quadrinucleata is a composite of the Latin numeral quattuor (quadr- in compositions; four) and nucleat·us, -a, -um (Latin adjective [m; f; n]; [nut]nucleus-like) and refers to the four macronuclear nodules. Remarks: Berger & Foissner (1989) assigned the present species to Amphisiella because the cirral pattern and the dorsal ciliature (especially the lack of caudal cirri) basically match the situation in the type species A. capitata. We avoided a classification in Lamtostyla because the amphisiellid median cirral row is longer than the adoral zone. Now we know that Amphisiella is a marine group showing a more or less complete set of pretransverse ventral and transverse cirri and a rather high number of dorsal kineties. Since the cirral pattern of the present species strongly resembles that of L. lamottei, and because the amphisiellid median cirral row is less than 50% of body length, Amphisiella quadrinucleata is transferred to Lamtostyla. Lamtostyla quadrinucleata closely resembles L. vitiphila (Fig. 35a–d). However, this species has only one buccal cirrus (vs. 2–3), but three dorsal kineties (vs. two). Since both features are usually difficult to recognise in life, protargol preparations should be made. Hemiamphisiella quadrinucleata (Fig. 62a–e) has the same type of nuclear apparatus, but differs from the present species by the presence of a postperistomial ventral cirrus, caudal cirri, and cortical granules and the lack of transverse cirri. The two dorsal kineties, the straight row behind the right frontal cirrus, and the increased number of buccal cirri is reminiscent of Orthoamphisiella spp., which, however, lack transverse cirri (Eigner & Foissner 1991, 1993). Ontogenetic data are needed for a more proper classification. Morphology: Body size in life about 100–125 × 30 µm; body length:width ratio of protargol-impregnated specimens about 3.4:1 (Table 22). Body margins parallel, anterior part of cell usually slightly bent to the left, both ends rounded (Fig. 35a, c). Body flexible, flattened about 2:1 dorsoventrally (Fig. 35b). Macronuclear nodules form two longitudinally arranged, indistinct pairs left of midline; individual nodules 1
Berger & Foissner (1989) provided the following diagnosis: In vivo about 100–125 × 30 µm, long ellipsoid. 4 macronuclear segments, 2 dorsal kineties, 2–3 buccal cirri. 15 cirri in the right frontoventral row, 3 transverse cirri, and 17 adoral membranelles on average.
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Fig. 35a–e Lamtostyla quadrinucleata (from Berger & Foissner 1989. a–c, from life; d, e, protargol impregnation). a: Ventral view of a representative specimen, 120 µm. b: Right lateral view showing dorsoventral flattening. c: Dorsal view showing contractile vacuole. d, e: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 87 µm. Short arrow marks buccal cirri, long arrow denotes cirri behind right frontal cirrus (ontogenetic data are needed to show whether these cirri originate from one or two anlagen). ACR = rear end of amphisiellid median cirral row, AZM = adoral zone of membranelles, FC = right frontal cirrus, MA = macronuclear nodules, MI = micronucleus, RMR = right marginal row, TC = transverse cirri, 1, 2 = dorsal kineties. Page 190.
about 10 × 6 µm in life, with large chromatin bodies. Contractile vacuole slightly ahead of mid-body, distinctly displaced inwards, during diastole without collecting canals. Cortical granules lacking. Cytoplasm colourless, with numerous 1–7 µm large, greasily shining globules and some food vacuoles. Slow, trembling movement. Adoral zone occupies about 20% of body length, composed of 17 membranelles of common fine structure. Undulating membranes slightly curved, about of same
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length, superimposed or closely arranged side by side. Buccal field moderately wide (Fig. 35a, d, Table 22). Cirral pattern and number of cirri of usual variability (Fig. 35d, Table 22). Frontal cirri slightly enlarged, arranged in almost transverse pseudorow. Buccal cirri right of anterior half of paroral. Three, rarely four, cirri form short row behind right frontal cirrus (ontogenetic data are needed to show whether this row is formed from one or two [more likely] cirral anlagen), row ends at 13% of body length on average. Amphisiellid median cirral row commences near distal end of adoral zone, terminates at 38% of body length on average (Table 22). Usually three more or less terminally arranged transverse cirri, in life about 14 µm long and therefore distinctly projecting beyond rear body end. Right marginal row commences almost at same level as amphisiellid median cirral row, ends subterminal. Left marginal row commences slightly behind proximal end of adoral zone; rear end J-shaped, that is, row terminates almost in midline of cell. Dorsal bristles about 3 µm long in life, arranged in two kineties; kinety 1 slightly shortened anteriorly; distances between bristles become wider in posterior direction in kinety 2. Caudal cirri lacking (Fig. 35e). Occurrence and ecology: Likely confined to terrestrial habitats (Foissner 1998, p. 199). The type locality of Lamtostyla quadrinucleata is a yellowish soil (pH 3.8; 10–20 m above sea-level) with poorly decomposed needles from Mae-Shima, Amakusa, Kumamoto Prefecture, Japan (Berger & Foissner 1989). No further records published. Feeds on ciliates like Colpoda fastigata and Pseudoplatyophrya nana (Berger & Foissner 1989; for details on food organisms, see Foissner 1993). Biomass of 106 specimens about 70 mg (Foissner 1998, p. 199).
Lamtostyla elegans (Foissner, Agatha & Berger, 2002) comb. nov. (Fig. 36a–d, Table 22) 2002 Amphisiella elegans nov. spec.1 – Foissner, Agatha & Berger, Denisia, 5: 674, Fig. 150a–d, Tables 133, 134 (Fig. 36a–d; original description; one holotype slide [accession number 2002/594] and four paratype slides [2002/595–598] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; Foissner et al. 2002, p. 37 and Aescht 2003, p. 385).
Nomenclature: The species-group name elegans (Latin adjective; elegant) refers to the elegant appearance of the species (Foissner et al. 2002). Remarks: For a foundation of the transfer of the present species from Amphisiella to Lamtostyla, see genus section. 1
Foissner et al. (2002) provided the following diagnosis: Size about 170 × 35 µm in vivo; narrowly lanceolate and slightly sigmoidal. Usually 4 macronuclear nodules forming 2 pairs left of midline. Amphisiellid median cirral row (ACR) ends above mid-body, composed of about 12 cirri. On average 23 adoral membranelles, about 48 cirri each in right and left marginal row, 3 cirri left of ACR, 1 buccal cirrus, 4 transverse cirri, and 2 dorsal kineties extending along right body margin in posterior body portion.
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This species is highly variable, even in features which are usually constant in hypotrichs. Thus, Foissner et al. (2002) separated specimens with four, respectively, six or more macronuclear nodules in the morphometry (Table 22). However, no clear-cut relationships emerged, except that all specimens with more than four nodules have three dorsal kineties (vs. two). Furthermore, some observations on other populations show that the Jordanian specimens do not exhaust the whole range of variability (see below). Lamtostyla elegans is, like several congeners, difficult to preserve. Only if the fixative is amended with osmium tetroxide, do they stabilise; however, the specimens from Namibian site (47) even disintegrated in osmium-amended Stieve solution (Foissner et al. 2002). Lamtostyla elegans differs from other quadrinucleate Lamtostyla species (L. quadrinucleata, L. vitiphila) by the large (length 150 µm vs. <100 µm after protargol impregnation), slender body (5:1 vs. <3:1). Uroleptoides longiseries, which also has four macronuclear nodules, has (i) thrice the number of cirri in the amphisiellid median cirral row, which thus extends to more than 50% of body length; (ii) cortical granules; and (iii) three dorsal kineties (Fig. 46a–g). Thus, Lamtostyla elegans is usually easily identified in life. Other quadrinucleate soil hypotrichs belong to different genera: Terricirra matsusakai (oral apparatus differently organised, green cortical granules, spindle-shaped; this book); Hemisincirra quadrinucleata and H. namibiensis (ventral cirral row not longer than adoral zone; this book); Parakahliella halophila Foissner, Agatha & Berger, 2002 (four dorsal kineties with caudal cirri, no transverse cirri); Bistichella buitkampi (this book) and Fragmocirrus espeletiae Foissner, 2000 (ventral cirral rows extend to near transverse cirri, three or more buccal cirri and dorsal kineties; rather broad, massive species). Morphology: Body size 140–200 × 25–45 µm in life, usually about 170 × 35 µm, length:width ratio highly variable also in protargol preparations, that is, 3.4 to 7.6:1, on average 5:1 (Table 22). Body dorsoventrally flattened up to 2:1. Body outline slender and often slightly sigmoidal with posterior end more narrowly rounded than anterior; body very flexible but acontractile, often twisted about main axis by half a turn (Fig. 36a; Table 22). Nuclear apparatus in middle third of cell left of midline. Number and arrangement of macronuclear nodules highly variable, usually a pair of nodules each above and below mid-body; in specimens with six or eight nodules, they form a strand, with one half of the nodules usually still separated from the other by a slightly increased distance. Of 56 specimens analysed, 13 had more than the usual four macronuclear nodules, an unusually high percentage (23%) for this kind of hypotrich. Individual nodules ellipsoidal on average and often connected by a strand of lighter impregnated material; chromatin bodies small and numerous. Micronuclei near, or attached to, macronuclear nodules, about 4 × 3 µm in life, number highly variable, especially in specimens with more than four macronuclear nodules (Table 22). Contractile vacuole slightly ahead of mid-body at left cell margin, during diastole with two collecting canals. Cortical granules lacking. Cytoplasm colourless, contains many lipid droplets 1–4 µm across, mainly in posterior
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Fig. 36a–c Lamtostyla elegans (from Foissner et al. 2002. a, from life; b, c, protargol impregnation). a: Ventral view of representative, slightly twisted specimen, 135 µm. Note the sail-like paroral composed of very closely spaced, 7 µm long cilia. b, c: Infraciliature of ventral and dorsal side and nuclear apparatus of holotype specimen, 126 µm. Arrow marks break in amphisiellid median cirral
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Fig. 36d Lamtostyla elegans (from Foissner et al. 2002. Protargol impregnation). About 13% of the specimens of the type population have five, six, or eight macronuclear nodules and a third, loosely ciliated kinety (arrows) indicating that this row is a remnant of the previous generation. LMR = rear end of left marginal row, 1, 2 = dorsal kineties. Page 192.
half of cell. Food vacuoles up to 30 µm across. Glides slowly and, although being very flexible, rather clumsily, possibly due to the twisted body, on microscope slide and soil particles. Adoral zone occupies only 16–23%, on average 19%, of body length, of usual shape and structure, composed of an average of 23 membranelles; frontal scutum possibly unusually thick because often faintly impregnated. Buccal cavity moderately deep and wide, posterior cavity margin projects angularly, partially covering adoral zone. Paroral and endoral rather distinctly curved and optically intersecting in midportion, likely composed of dikinetids. Paroral cilia about 7 µm long and very closely spaced, forming compact, sail-like structure in life. Endoral cilia at least 15 µm long, beat into the pharynx, form conspicuous bundle with pharyngeal fibres extending obliquely backwards (Fig. 36a, b). Cirral pattern rather constant, while cirral number unusually variable, especially in specimens with six or more macronuclear nodules (Fig. 36a, b; Table 22). Most cirri 8–10 µm long in life and rather fine, especially in posterior portion of marginal rows and in median cirral row. Frontal cirri slightly enlarged, right cirrus, as is usual, at distal end of adoral zone. Buccal cirrus near summit of curve formed by paroral. On average three cirri left of anterior portion of amphisiellid median cirral row which commences close to the right frontal cirrus and on average terminates at 31% of body length right of cell’s midline; occasionally a distinct break in anterior third of row, showing that it is composed of (at least) two segments. Transverse cirri subterminally inserted, about 12 µm long in life, inconspicuous because fine, likely consisting of only four cilia each. Right marginal row extends onto dorsal side in anterior body quarter and ends far subterminally, that is, at or ahead of level of transverse cirri. Left marginal row extends onto dorsal side in posterior body fifth to end terminally in body midline; last cirri easily misinterpreted as caudal cirri.
b
row indicating that it is composed of (at least) two portions. Frontal cirri connected by dotted line, cirri left of anterior portion of amphisiellid median cirral row circled; right frontal cirrus and cirrus III/2 connected by broken line. ACR = amphisiellid median cirral row, AZM = adoral zone of membranelles, E = endoral, FC = left frontal cirrus, LMR = left marginal row, RMR = rear end of right marginal row, TC = transverse cirri, 1, 2 = dorsal kineties. Page 192.
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Dorsal bristles about 3 µm long in life, arranged in two common rows or in three rows with left row very sparsely ciliated, indicating that it is a remnant of a previous generation (Fig. 36c, d). Both rows commence subapically left of or in body midline and course obliquely backwards ending at right posterior margin of cell, an unusual and highly constant pattern found also in several congeners, for instance, Lamtostyla procera. Caudal cirri lacking. Observations on other populations: Foissner et al. (2002) found this or very similar species also in African soil samples (see occurrence and ecology section). Unfortunately, they did not withstand the preparation procedures, as explained above, and thus we have data only from live observations. The specimens from Namibian site (51) are only 120–130 µm long, very slender (length:width ratio about 7:1), and have minute (<0.2 µm), colourless cortical granules around bases of cirri and dorsal bristles. Furthermore, they possess some lithosomes about 3 µm across. The specimens from the highly saline sample from Saudi Arabia are very similar to those from Jordan, but have three buccal cirri, indicating that they might be a slender variant of Lamtostyla quadrinucleata. Considering the high variability of the Jordanian L. elegans, it seems reasonable to assign these populations to this species. Occurrence and ecology: Lamtostyla elegans is very likely confined to terrestrial habitats (Foissner et al. 2002, p. 50). Type locality is Wadi Ram (29°30'N 35°E) about 10 km east of Al Aqaba (Jordan), where we discovered it in a soil sample (pH 6.7, salinity <0.2%; collected by H. Blatterer, Linz) containing red sand mixed with litter from shrubs (possibly Chenopodiaceae) and grasses (Foissner et al. 2002). While L. elegans was rare at type locality, an abundant population developed in brownish soil mixed with plant litter and rabbit (?) droppings from the same area. Furthermore, we found L. elegans in a highly saline (2.1%) soil sample collected in Saudi Arabia near the village of Alqasab. Finally, we observed some specimens at two Namibian sites, that is, in bark from Combretum imberbe and Colophospermum mopane (detailed description of sites 47 and 51, see Foissner et al. 2002). Thus, Lamtostyla elegans is euryhaline and has a broad ecological range. In spite of this, W. Foissner did not find it in about 1000 other samples from terrestrial habitats collected world-wide (Foissner 1998). Lamtostyla elegans feeds on small ciliates (Pseudocohnilembus sp., Colpoda aspera, swarmers of Vorticella astyliformis) and naked amoebae (Foissner et al. 2002).
Lamtostyla islandica Berger & Foissner, 1988 (Fig. 37a–d, 38a, Table 22) 1982 Tachysoma perisincirra n. spec. – Hemberger, Dissertation, p. 251, pro parte, Abb. 46c, not Abb. 46a, b, d (Fig. 37d; see remarks). 1985 Tachysoma perisincirra n. spec. (pro parte) – Hemberger, Arch. Protistenk., 130: 413 (see remarks).
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1988 Lamtostyla islandica nov. spec.1 – Berger & Foissner, Zool. Anz., 220: 122, Fig. 15–18, Table 1 (Fig. 37a–c, 38a; original description; the holotype slide [accession number 1988/53; Aescht 2003, p. 388] and a paratype slide [1988/56] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria). 2001 Lamtostyla islandica Berger and Foissner, 1988 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 45 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs).
Nomenclature: No derivation of the name is given in the original description. The species group name islandic·us, -a, -um (Latin adjective [m; f; n]; occurring in/indigenous to Iceland) refers to the area (Iceland) where the species was discovered. Remarks: Hemberger (1985) described and figured Lamtostyla perisincirra with one micronucleus between the two macronuclear nodules (Fig. 39a–f). However, he mentioned that the number of micronuclei perhaps varies because he found specimens with an identical ventral and dorsal infraciliature, but two micronuclei and a more elongated body shape (Fig. 37d). Our population from Iceland showed the same state of this character pair, indicating the existence of two closely related, but clearly separable species (Fig. 38a). Thus, we described our population from Iceland as new species (Berger & Foissner 1988). Morphology: Body size in life about 60–80 × 20–25 µm; body length:width ratio of protargol-impregnated specimens 3.4:1 on average (Fig. 37a, 38a, Table 22). Body margins parallel, anterior body portion slightly narrowed, rear end broadly rounded. Two macronuclear nodules slightly left of midline about in body centre; individual nodules ellipsoidal with medium-sized chromatin bodies. Usually each macronuclear nodule with a single micronucleus, rarely only one nodule with a micronucleus (never between the two macronuclear nodules!). Contractile vacuole slightly ahead of mid-body near left cell margin. Cortical granules lacking. Cytoplasm and movement without peculiarities. Adoral zone occupies 27% of body length on average in protargol preparations (Table 22), composed of about 16 membranelles of ordinary fine structure (Fig. 37b). Buccal cavity narrow. Undulating membranes short, arranged roughly in parallel but overlapping only by about half of their length. Cytopharynx extends obliquely backwards. Cirral pattern of usual variability (Fig. 37b, c, Table 22). Frontal cirri arranged in slightly oblique pseudorow, only slightly larger than remaining cirri, right cirrus, as is usual, behind distal end of adoral zone. Buccal cirrus ahead of paroral. Three cirri left of anterior portion of amphisiellid median cirral row, including cirrus III/2 which is slightly shifted leftwards. Amphisiellid median cirral row usually composed of six cirri, terminates at 27% of body length on average, that is, as long as adoral zone. Usually three transverse cirri near rear body end, project distinctly beyond rear cell end. Right marginal row commences slightly ahead of level of buccal vertex, ends, like left row, subterminally; left row commences near buccal vertex. 1
Berger & Foissner (1988) provided the following diagnosis: In vivo c. 60–80 × 20–25 µm. About 16 adoral membranelles, 6 cirri in the frontal row, and usually 3 cirri left of the frontal row. 14 left and 15 right marginal cirri on the average. 3 dorsal kineties. Usually 2 micronuclei.
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Fig. 37a–d Lamtostyla islandica (a–c, from Berger & Foissner 1988; d, from Hemberger 1982. a, from life; b–d, protargol impregnation). a: Ventral view of representative specimen, 67 µm. b, c: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 73 µm. Arrow marks buccal cirrus ahead of paroral. Frontal cirri connected by dotted line, cirri left of anterior portion of amphisiellid median cirral row are circled, and broken line connects right frontal cirrus and cirrus III/2. d: Infraciliature of ventral side and nuclear apparatus of early divider, details see text. ACR = anterior end of amphisiellid median cirral row, AZM = adoral zone of membranelles, E = endoral, FC = right frontal cirrus, LMR = anterior end of left marginal row, MA = anterior macronuclear nodule, MI = micronucleus, P = paroral, RMR = anterior end of right marginal row, TC = transverse cirri, 1–3 = dorsal kineties. Page 196.
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Fig. 38a Plot of individual measurements for specimens of Lamtostyla islandica (O; n = 12) and Lamtostyla perisincirra (*; n = 28). “2” and “3” above or right of a symbol means that two or three specimens have the same size. The black dot is a combination of a ring and an asterisk (from Berger & Foissner 1988). Pages 196, 200.
Dorsal bristles about 2–3 µm long, arranged in three kineties; kinety 1 slightly shortened anteriorly and posteriorly, kinety 2 anteriorly only, and kinety 3 posteriorly only (Fig. 37c). Caudal cirri lacking. Cell division: Hemberger (1982) found an early divider (Fig. 37d). Very likely, the oral primordium is formed de novo in the postoral region, possibly near the transverse cirri. Occurrence and ecology: Lamtostyla islandica is very likely confined to terrestrial habitats and of cosmopolitan distribution (Foissner 1998, p. 205). Type locality is Gooa Foss, Bardårdalur, North Iceland, where we found it in soil of a heath with dwarf shrubs (Berger & Foissner 1988). Further records: two natural forest stands in eastern Austria (Foissner et al. 2005, p. 629); frequent (22%) in terrestrial habitats from Namibia (Foissner et al. 2002, p. 60); upper soil layer (0–5 cm) with roots and litter from a sand hill in the 99 mile desert (north of Lake Alexandrina), near the city of Adelaide, Australia (Blatterer & Foissner 1988, p. 8); litter, soil, and roots (0–5 cm; pH 5.1) from a terra firma secondary rain forest about 300 m off the Rio Negro (not flooded during high water periods of the river) in the surroundings of Tropical Hotel Manaus, outskirts of Manaus, Brazil (Foissner 1997, p. 322); Sphagnum moss under mire vegetation (pH 5.0) from Tafelkop, Gough Island, and red lava/scoria gravel (no vegetation; pH 6.6; 200 m above sea level) from scree slope of Junior’s Kopoil as well as red lava/scoria gravel/mud with little vegetation (pH 6.3; about 150 m above sea level) near base of Junior’s Kop, Marion Island, southern Indian Ocean (Foissner 1996a, p. 284). Lamtostyla islandica feeds on bacteria and heterotrophic flagellates (Berger & Foissner 1988). Biomass of 106 specimens about 20 mg (Foissner 1998, p. 205).
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Lamtostyla perisincirra (Hemberger, 1985) Berger & Foissner, 1987 (Fig. 38a, 39a–m, Table 22) 1982 Tachysoma perisincirra n. spec.1 – Hemberger, Dissertation2, p. 251, (pro parte), Abb. 46a, b, d, not Abb. 46c (Fig. 39a, b; see nomenclature and remarks). 1984 Tachysoma perisincirra Hemberger – Berger, Foissner & Adam, Zool. Jb. Syst., 111: 363, Abb. 58–68, Tabelle 8 (Fig. 39c–m; detailed redescription and morphogenesis; two voucher slides [accession numbers 1986/99, 100] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; Aescht 2003, p. 394, see nomenclature). 1985 Tachysoma perisincirra n. spec. – Hemberger, Arch. Protistenk., 130: 412, (pro parte), Abb. 20 (Fig. 39a, b; original description; no formal diagnosis provided; type slide is deposited in the Institut für landwirtschaftliche Zoologie, University of Bonn, Germany; see nomenclature and remarks). 1987 Lamtostyla perisincirra (Hemberger, 1985) nov. comb. – Berger & Foissner, Zool. Jb. Syst., 114: 216 (combination with Lamtostyla). 1988 Lamtostyla perisincirra (Hemberger, 1985) Berger and Foissner, 1987 – Berger & Foissner, Zool. Anz., 220: 119, Fig. 3–14, 18, Table 1 (Fig. 38a, 39a–m; revision of Lamtostyla). 1994 Amphisiella perisincirra (Hemberger, 1985) nov. comb. – Eigner & Foissner, J. Euk. Microbiol., 41: 255 (combination with Amphisiella). 2001 Lamtostyla perisincirra (Hemberger, 1985) Berger & Foissner, 1987 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 88 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Lamtostyla perisincirra – Lynn & Small, Phylum Ciliophora, p. 459, Fig. 57A (Fig. 39e; guide to ciliate genera).
Nomenclature: No derivation of the name is given in the original description. The species group name perisincirra (likely from the genus-group name Perisincirra Jankowski, 1978) obviously alludes to the fact that the cirral pattern of the present species resembles that of Perisincirra species (now most species are in Hemisincirra Hemberger, 1985). We deposited two protargol slides in the Upper Austrian Museum in Linz. Unfortunately, we forgot to mention the deposition and details about the slides in the publication (Berger et al. 1984). According to Aescht (2003, p. 394), we labelled the slides as neotypes. However, the designation as neotype is incorrect because the type slide(s) are deposited in the Institut für landwirtschaftliche Zoologie of the University of Bonn (Hemberger 1982, 1985; see list of synonyms). Consequently, our slides deposited in Linz are voucher slides. Remarks: According to Hemberger (1982, 1985), the present species resembles Tachysoma bicirratum (Foissner, Blatterer, Berger & Kohmann, 1991) Foissner, Blatterer, Berger & Kohmann, 1991 (= T. furcata Kahl in Hemberger 1982, 1985; for review, see Berger 1999, p. 457) in body size and nuclear apparatus, whereas the cirral pattern is similar to that of Perisincirra (Hemberger 1982) and Hemisincirra (Hemberger 1985) species (see present book). However, since ontogenesis commences near the transverse cirri in the present species he avoided a classification in Hemisincirra, where the first primordium originates at the rear end of the frontoven1 2
This name is disclaimed for nomenclatural purposes (ICZN 1999, Article 8.3). See next footnote. See same footnote at Uroleptoides binucleatus.
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tral row (however, note that the ontogenesis of the type species of Hemisincirra is not known). Furthermore, all Hemisincirra species have many macronuclear nodules (vs. two in L. perisincirra). Thus, he provisionally assigned it to Tachysoma because caudal cirri are lacking both in this genus and in the present species. Berger et al. (1984) provided a detailed redescription and some ontogenetic stages, but did not change the generic classification. Somewhat later we transferred it to Lamtostyla because it agrees with several Lamtostyla species in the following features (Berger & Foissner 1987): dorsal kineties without caudal cirri; small body length; reduced number of transverse cirri and pretransverse ventral cirri; lack of postoral ventral cirri (in ordinary postoral location), and possession of a short frontoventral row which terminates at about level of cytostome. The classification in Lamtostyla was retained by Berger & Foissner (1988), whereas Eigner & Foissner (1994) included it in Amphisiella. However, Petz & Foissner (1996, p. 277) supported Berger & Foissner’s (1987) assignment to Lamtostyla because the ontogenesis proceeds similar as in L. edaphoni (now Lamtostylides edaphoni) and L. australis; and, more important, the cirral pattern and habitat is much more similar to that of L. lamottei, type of Lamtostyla, than to that of A. capitata, type of Amphisiella. Hemberger (1982, 1985) described and figured the present species with one micronucleus between the two macronuclear nodules (Fig. 39a, b). However, he mentioned that the number of micronuclei perhaps varies because he found specimens with an identical ventral and dorsal ciliature, but two micronuclei and a more elongate body shape (Fig. 37d, 39a). Berger & Foissner (1988) found a population in Iceland showing the same state of this character pair, indicating the existence of two closely related, but clearly separable species: Lamtostyla perisincirra (stouter, one micronucleus between the macronuclear nodules; Fig. 38a, 39c, d, e) and L. islandica (longer, two micronuclei; Fig. 37a, c, 38a). Morphology: The population described by Berger et al. (1984) matches the cirral pattern and the ontogenesis described by Hemberger (1982, 1985) more or less perfectly so that conspecificity is beyond doubt. Thus, the description below is based on the data by Berger et al. (1984), unless otherwise indicated. Body size about 50–80 × 20–30 µm in life, body length according to Berger & Foissner (1988) up to 90 µm; body size according to Hemberger (1982, 1985) 60–80 × 20–23 µm (23 µm is likely incorrect because the average size of Hemberger’s specimens is usually 60 × 25 µm); body length:width ratio 2.8:1 in protargol preparations (Table 22). Body outline as shown in Fig. 39c, that is, right margin straight to slightly concave, left one distinctly vaulted about in mid-body; both ends broadly rounded. Body flattened about 2:1 dorsoventrally, ventral side flat, dorsal side slightly convex (Fig. 39d); translucent at low magnification; flexible and slightly contractile under cover glass. Macronuclear nodules slightly left of midline about in cell centre, in life about 7 × 4 µm, with some small to medium-sized chromatin bodies. One micronucleus between macronuclear nodules, ellipsoidal or globular (2–3 µm across), stains well with protargol (for Hemberger’s record of specimens with two micronuclei see remarks and L. islandica). Contractile vacuole at some distance
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from left cell margin about at level of micronucleus, that is, about in mid-body, without distinct collecting canals during diastole (Fig. 39c, d). Cortical granules lacking. Cytoplasm colourless, with many greenish globules 2–5 µm across, making cells bright at low magnification; posterior body portion often with some dark (refractive?) globules. Locomotion fast and erratic. Adoral zone occupies 30% of body length and composed of 16 membranelles of ordinary fine structure on average (Table 22). Buccal field small. Undulating membranes only slightly overlapping and not (or not distinctly) intersecting optically. Cytopharynx extends obliquely backwards (Fig. 39a, c, e). Cirral pattern and number of cirri of usual variability (Fig. 39a, e, Table 22). Frontal cirri slightly enlarged. Buccal cirrus right of anterior end of anterior undulating membrane (paroral?). Three cirri (including cirrus III/2) left of amphisiellid median cirral row. Amphisiellid median cirral row composed of six cirri on average, terminates about at level of buccal vertex. Usually three or four transverse cirri (including a possible pretransverse ventral cirrus) close to rear cell end; according to Hemberger (1982, 1985), the small cirrus ahead of the right transverse cirrus is a pretransverse ventral cirrus (Fig. 37a, arrow). Transverse cirri about 15 µm long (Hemberger 1982, 1985). Marginal cirri about 8–10 µm long in life, distance between cirri increases slightly from anterior to posterior. Right marginal row commences slightly ahead of level of buccal vertex, ends, like left row, at or somewhat ahead of level of transverse cirri; that is, marginal rows widely separated posteriorly; left marginal row commences about at level of buccal vertex (Fig. 39a, e). Dorsal bristles 2–3 µm long in life (according to Hemberger 1982, 1985, the length is 3–5 µm), arranged in three, rarely in four kineties, which are more or less distinctly shortened anteriorly and posteriorly; number of bristles per kinety increases from left (kinety 1) to right (kinety 3). Caudal cirri lacking (Fig. 39f, Table 22). Ryan et al. (1989) found Lamtostyla perisincirra in an Antarctic soil (see occurrence). Foissner (pers. comm.) checked the identification in protargol preparations and could not find any important differences to the populations described by Hemberger (1982, 1985) and Berger et al. (1984).
b
Fig. 39a–h Lamtostyla perisincirra (a, b, after Hemberger 1985; c–h, after Berger et al. 1984. a, b, e–h, protargol impregnation; c, d, from life). a, b: Infraciliature of ventral side and nuclear apparatus of a specimen from the type population, 58 µm. Arrow marks the pretransverse ventral cirrus. c, d: Ventral view and right lateral view of a representative specimen, 62 µm. Note the highly characteristic nuclear apparatus, which is composed of two macronuclear nodules and a micronucleus in between. e, f: Infraciliature of ventral and dorsal side and nuclear apparatus, e = 40 µm, f = 45 µm. The amphisiellid median cirral row is composed of two portions originating from frontal-ventral-transverse cirri anlagen V (posterior portion) and VI (anterior portion). Cirri left of amphisiellid median cirral row circled (cirrus III/2 included). Frontal cirri connected by dotted line. Cirri which originate from the same anlage are connected by a broken line (corresponding transverse cirri not included). g, h: Posterior portion of very early dividers, showing that the oral primordium originates from/near the transverse cirri. AZM = adoral zone of membranelles, ACR = amphisiellid median cirral row, CV = contractile vacuole, MA = macronuclear nodule, MI = micronucleus, LMR = left marginal row, OP = oral primordium, RMR = right marginal row, TC = transverse cirri, 1–3 = dorsal kineties. Page 200.
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Fig. 39i–m Lamtostyla perisincirra (from Berger et al. 1984. Protargol impregnation. Parental structures white, new black). i–k: Infraciliature of ventral side and nuclear apparatus of early (i, j) and middle (k) dividers. Arrow in (j) marks primordium at rear end of amphisiellid median cirral row. l, m: Infraciliature of ventral and dorsal side and nuclear apparatus of late divider, 38 µm. Note that the amphisiellid median cirral row is formed, as is usual, from the anlagen V (posterior portion) and VI (anterior portion; homologous to the frontoterminal cirri of, for example, the oxytrichids and urostyloids). I–VI = frontal-ventraltransverse cirri anlagen, 1–3 = new dorsal kineties of proter. Page 200.
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Cell division: Hemberger (1982) and Berger et al. (1984) found some dividers (Fig. 39g–m). Ontogenesis commences with the formation of an oral primordium near the transverse cirri (Fig. 39g–j). In addition, the rearmost cirrus of the amphisiellid median cirral row modifies to a primordium (Fig. 39j). In middle dividers the anlagen field widens and partially contacts the anlagen originating from parental cirri (Fig. 39k). Late dividers have six frontal-ventral-transverse cirri primordia per filial product. Very likely, the amphisiellid median cirral row is formed, as is usual, from the cirri of the anlagen V and VI (Fig. 39l). The new dorsal kineties originate in common manner, that is, two primordia are formed within each parental kinety (Fig. 39m). Division of the nuclear apparatus proceeds plesiomorphically (Fig. 39m). Occurrence and ecology: Very likely confined to terrestrial habitats (Foissner 1987, p. 125; 1998, p. 205). The type locality of Lamotstyla perisincirra is not given by Hemberger (1985), who mentions only “suspension of mull rendzina soil”. However, this soil sample is not from Peru, the sole country mentioned under materials, but from Germany (see Hemberger 1982, p. 2). According to a personal communication by Norbert Wilbert (University of Bonn, Germany), the site is near the village of Mechernich (50°36'N 6°39'E), Germany. It is a beech forest on the “Kakushöhle Nordhang” and the outcrops are a middle Devonian dolomite and a Pleistocene tufaceous limestone. Berger et al. (1984) found L. perisincirra in an alpine pasture (0–5 cm; 1965 m altitude) in the Schloßalm area, near the village of Bad Hofgastein, Austria (see also Berger 1985; Berger et al. 1985, p. 107; 1986, p. 268). Further records: with a dominance of up to 4% in an alpine soil (Festkogel near the village of Obergurgl, Tyrol, Austria) from a ski slope about 2800 m above sea level (Lüftenegger et al. 1986, p. 152); subalpine grassland in Styria, Austria (Foissner et al. 1990, p. 18); various Austrian soils (Lüftenegger et al. 1988, p. 96); five small samples bulked (dry moss patch [Bryum sp.] beside snow water stream; wet moss patch [Bryum sp.] beside snow water stream; algae from flowing snow water stream; stream side, soil algal crust between stone; stream side, wet Bryum with algal growth) from Keble Valley (168°E 77°50'S), Ross Island, Cape Bird, Antarctic Region (Foissner 1996b, p. 99; identification uncertain); moss (Sarconeurum glaciale) from the Robertskollen area, western Dronning Maud Land, Antarctica (Ryan et al. 1989, p. 18). The record from the Manzanares River (La Pedriza, Madrid, Spain) by Fernandez-Leborans & Antonio-García (1988) is not substantiated by morphological data and thus should not be overinterpreted. Food not known; likely L. perisincirra ingests mainly bacteria because always translucent. Biomass of 106 specimens about 7 mg (Foissner 1987, p. 125).
Lamtostyla granulifera-group This group comprises Lamtostyla granulifera and L. decorata. It is characterised by an 18-cirri pattern with anteriorly displaced postoral ventral cirri and three dorsal kineties. In addition, both species have cortical granules.
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Lamtostyla granulifera Foissner, 1997 (Fig. 40a–n, Table 22) 1997 Lamtostyla granulifera n. sp.1 – Foissner, Biol. Fertil. Soils, 25: 332, Fig. 25–38, Table 6 (Fig. 40a–n; original description; the holotype slide [accession number 1998/116; Aescht 2003, p. 387] and two paratype slides [1998/117, 118] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria). 2001 Lamtostyla granulifera Foissner, 1997 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 45 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs).
Nomenclature: The species-group name granulifer·a, -a, -um (Latin adjective; [m; f; n]; bearing granules) is a composite of granulum (Latin noun; granule), the thematic vowel ·i- , and fero (Latin verb; bearing) and refers to the conspicuous cortical granules (Foissner 1997). Remarks: Lamtostyla granulifera has, like many oxytrichids, 18-frontal-ventraltransverse cirri. Only the three postoral ventral cirri are not arranged behind the adoral zone, but right of the proximal portion of the zone (Fig. 40n). Thus, Lamtostyla granulifera could be assigned to the oxytrichids according to the ventral cirral pattern. However, the dorsal ciliature consists, as in the amphisiellids, of three bipolar kineties, that is, dorsomarginal rows and dorsal kinety fragmentation – features characteristic for the Dorsomarginalia and the oxytrichids (Berger 1999, 2006) – are lacking. Since the amphisiellid median cirral row is short (less than 50% of body length), the classification in Lamtostyla proposed by Foissner (1997) is accepted. For a comparison of L. granulifera with L. decorata, see L. decorata. The ventral cirral pattern of L. granulifera is identical to that of L. longa (Fig. 43a), which, however, is somewhat smaller (85–130 µm in L. longa vs. 120–170 µm in L. granulifera), has an inconspicuous buccal cavity and adoral zone (vs. Cyrtohymena-like and prominent), possibly lacks cortical granules (vs. present), and has five dorsal kineties (vs. three). Lamtostyla raptans is larger than L. granulifera (prepared specimens 200 µm long), has more adoral membranelles (33), marginal cirri (about 60), and dorsal kineties (five), and, possibly, lacks cortical granules (Fig. 44a, b). For a grouping of L. granulifera, L. decorata, L. longa, and L. raptans, see genus section. Morphology: Foissner (1997) studied seven populations from life; however, usable protargol preparations were obtained only from the Venezuelan population, which he therefore fixed as type. Specimens from all populations burst when conventional fixatives were applied; and even in fixatives containing osmium tetroxide only few specimens maintained integrity. Only the rich population from a field near 1
Foissner (1997) provided the following diagnosis: Size in vivo about 150 × 35 µm. Slenderly elliptical and often slightly curved. Two macronuclear nodules. Cortical granules colourless, usually 1–2 µm across and arranged in conspicuous, narrowly spaced rows on ventral and dorsal side. Frontal cirral row about as long as adoral zone of membranelles, usually consisting of four cirri. Adoral zone continuous, proximal portion broadened spoon-like, consists of 24 membranelles on average. Buccal cavity deep and distinctly curved. On average 44 right and 45 left marginal cirri, 3 frontal cirri, 1 buccal cirrus, 2 pretransverse cirri, 5 transverse cirri, 3 cirri left of frontal row, and 3 dorsal kineties.
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Puerto Ayacucho provided – after fixation in Stieve’s solution mixed with 2% osmium tetroxide (3:1) – sufficient specimens for a detailed analysis. The reason for the extreme fragility of L. granulifera under various fixatives is unknown and surprising because live specimens are rather robust and even withstand slight cover glass pressure (Foissner 1997). The detailed description provided by Foissner (1997) is based mainly on the type population from Venezuela. The life observations largely agree for all populations and are thus combined where appropriate. However, the South African population from Cape Town might be a distinct (sub)species because it is markedly more slender (200 × 30 µm) and has the cortical granules (0.5–2.0 µm, colourless) arranged only around the cirral and bristle bases (Fig. 40k). The infraciliature matches, according to the live observations, that of the type population. Body size in life 120–170 × 20–55 µm, usually about 150 × 36 µm (n = 10). Body length:width ratio highly variable, namely 3.5:1 to 6:1; prepared specimens usually broader (from 2:1 to 4:1) due to poor fixation. Body outline slenderly to rather broadly elliptical, often almost parallel-sided and slightly curved, widest usually in or near mid-body, both ends evenly rounded (Fig. 40a–c). Thailand population and one Australian population slightly wedge-shaped because somewhat narrowed posteriorly (Fig. 40f). Body dorsoventrally flattened up to 2:1, very flexible. Specimens yellowish to brownish at low magnification (≤100), possibly due to dense cortical granulation and/or cytoplasmic inclusions. Two macronuclear nodules in central body portion slightly left of midline, distinctly ellipsoidal (from 2:1 to 3:1), with many tiny chromatin bodies. Micronuclei in small indentation of macronuclear nodules, conspicuous because large, compact, and distinctly ellipsoidal (5–8 × 3–4 µm, mean = 5.9 × 3.3 µm, length:width ratio from 1.5:1 to 3:1; Fig. 40a, l). Contractile vacuole near left cell margin distinctly ahead of mid-body, during diastole with two lacunar collecting canals (Fig. 40d). Cortical granules (Fig. 40d, h–k) colourless, rather hyaline and bright, conspicuous because 1–4 µm (usually 2 µm) across and in narrowly spaced rows leaving blank only cirri and dorsal bristles (except South African population, see above and Fig. 40k); found not only in somatic, but also in buccal cortex and around cytopharynx; usually disappear (burst? extruded?) when cells are slightly squeezed between slide and cover glass; do not stain with methyl green-pyronin, in protargol preparations occasionally appearing as dark, tiny (<0.3 µm) dots. Cytoplasm usually packed with 1–15 µm-sized globular and irregularly shaped, greasily shining inclusions and food vacuoles (Fig. 40a). Movement of L. granulifera conspicuous because glides rapidly to and fro, frequently changing direction. Adoral zone of membranelles make cells with look bushy because occupying only about 28% of body length (Table 22) and spoon-like broadening of proximal half, with bases of largest membranelles about 12 µm wide and of conventional fine structure (Fig. 40l, n). Proximal quarter of zone covered by buccal lip and undulating membranes (Fig. 40a, n). Buccal cavity deep, but rather narrow, distinctly curved anteriorly, posterior half covered by “bay-like” projecting buccal lip (Fig.
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40a). Undulating membranes distinctly curved, both very likely composed of narrowly spaced dikinetids and optically intersecting in anterior third. Paroral distinctly shorter than endoral, which terminates near proximal end of adoral zone. Paroral cilia in life about 15 µm long, endoral cilia about 40 µm (!) long, beating wavelike into conspicuously large, conical pharynx supported by fine fibres (Fig. 40a, l, n). Note that this species has, like many oxytrichids and hypotrichs in general, 18 frontal-ventral-transverse cirri (for discussion see remarks; Fig. 40l, n). Frontal cirri distinctly enlarged, composed of 4–5 kineties each, shape rather variable, right cirrus occasionally difficult to identify because close to distal end of adoral zone. Buccal cirrus close to summit of curve formed by paroral, composed of four kineties with about five cilia each (Fig. 40n). Three cirri left of amphisiellid median cirral row (for homologisation with cirri of 18-cirri hypotrichs, see Fig. 40n), each cirrus composed of three kineties with 4–6 cilia each. Amphisiellid median cirral row short because composed of four cirri, extends slightly obliquely from near right frontal cirrus to almost level of proximal end of adoral zone; anterior pair consists, like marginal cirri, of two kineties with about five cilia each, posterior two cirri composed of three kineties with about five cilia each. Usually two small pretransverse ventral cirri close to two rightmost transverse cirri. Transverse cirri distinctly enlarged, in life about 20 µm long and projecting beyond posterior body margin, associated with long fibres extending to mid-body, forming conspicuous, wedge-shaped bundle. Marginal cirri in life about 12 µm long, composed of two kineties with 5–7 cilia each (Fig. 40n). Left marginal row conspicuous because extending around posterior body margin, its rearmost cirri thus easily mistaken for caudal cirri; commences somewhat ahead of level of proximal end of adoral zone. Right marginal row begins at level of right frontal cirrus and terminates near transverse cirri, thus separated from left marginal row by small but distinct break. Dorsal bristles in life 3–4 µm long, arranged in three almost bipolar kineties. Kinety 1 commences near midline of cell and extends in short, sharp bow to left body margin, where it continues as straight line to posterior body end. Rows 2 and 3 extend in slightly oblique bows from anterior right half to posterior left half of body. Thus, a large elliptical, barren area is formed between kineties 1 and 2. Caudal cirri lacking (Fig. 40m). Occurrence and ecology: Lamtostyla granulifera is very likely confined to terrestrial habitats (Foissner 1998). The type locality is a field (Mahada) soil from the farm of Don Pedro Cortez, El Sapo village, about 50 km north of Puerto Ayacucho, Venezuela (about 68°W 5°N; Foissner 1997). The Mahada was three years old and
b
Fig. 40a–k Lamtostyla granulifera (from Foissner 1997. From life). a: Ventral view of a representative specimen (150 µm) with a large food vacuole containing a very small hypotrichous ciliate. b, c: Ventral and right lateral view of saccular shape variant. d–g: Shape variants (d also shows arrangement of cortical granules) from several populations. h–k: Arrangement of cortical granules (1–4 µm) in ventral (upper series; arrow in k marks cirrus) and dorsal side (lower series) of populations 1, 3–6. CG = cortical granules, CV = contractile vacuole with collecting canals, DB = dorsal bristle. Page 206.
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planted with bananas and manioc and the soil was spongy, strongly enriched with organic material (mainly cow dung), light brown, contained many fine roots from the grass cover, and had a pH of 6.5. Foissner (1997) reported that he has found this species in terrestrial habitats from widely distant regions, indicating that it has a broad ecological and geographical range. However, he has never found it in Europe, and most populations had a low abundance, appearing about one week after wetting the sample and disappearing in the second week, indicating that Lamtostyla granulifera is more r- than K-selected (Foissner 1997). Population 1: thick bark (pH 4.6) from a rain forest tree in the vicinity of Cairns, Australia. Population 2: litter and soil (0–5 cm) from a young Eucalyptus forest (pH 4.9) at the entrance to Foog Dam near Darwin, Australia. Population 3: litter, roots, and brown soil (0–5 cm) from mixed forest with bamboo and Pinus sp. at base of „female” Tsukuba mountain near Tokyo, Japan. Population 4: deciduous litter and red-brown soil (pH 6.4) from a coastal rain forest near Kata Karon town, Phuket peninsula, Thailand. Population 5: litter with much fungal hyphae and upper (0–5 cm), very sandy soil layer from secondary mixed (rain) forest (Pinus sp., deciduous trees) near the bank of Skeleton River at base of Table Mountain, Botanical Garden of Cape Town (Kirstenbosch), Republic of South Africa. Population 6: type locality (see above). Population 7: material from a termite (Anoplotermes sp.) hill from Ilha de Marchantaria, an island in the Rio Solimões near Manaus, Brazil. Not found in 73 soil samples from Namibia (Foissner et al. 2002). Lamtostyla granulifera feeds on naked amoebae, heterotrophic flagellates, and ciliates such as Cyclidium muscorum, Leptopharynx costatus, Drepanomonas sp., and Oxytricha setigera, which are ingested whole, and remain moving for some time in the food vacuoles (Fig. 40a; Foissner 1997). Biomass of 106 specimens about 90 mg (Foissner 1998, p. 205).
b
Fig. 40l–n Lamtostyla granulifera (from Foissner 1997. Protargol impregnation). l, m: Infraciliature of ventral and dorsal side and nuclear apparatus of a specimen from the type population, 150 µm. Note that the left marginal row curves around the posterior body end. Arrow marks anterior end of amphisiellid median cirral row. Cirri left of amphisiellid median cirral row circled. n: Infraciliature of anterior ventral portion at higher magnification. Arrow marks buccal cirrus; cirri which very likely originate from the same anlage are connected by a broken line. The cirri of this species can be easily homologised with the cirri of the 18-cirri hypotrichs (see general section or Fig. 6a in Berger 1999). “Frontoventral cirri” (cirri III/2, IV/3, VI/3, VI/4) circled by compact line; “postoral ventral cirri” (cirri IV/2, V/3, V/4) circled by dotted line. AZM = distal end of adoral zone, E = endoral, FC = frontal cirri (connected by dotted line), LMR = left marginal row, MA = macronuclear nodule, MI = micronucleus, P = paroral, PT = pretransverse ventral cirri, RMR = right marginal row, I–VI = frontal-ventral-transverse cirri anlagen, 1–3 = dorsal kineties. Page 206.
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Lamtostyla decorata Foissner, Agatha & Berger, 2002 (Fig. 41a–n, 42a–j, Table 22) 2002 Lamtostyla decorata nov. spec.1 – Foissner, Agatha & Berger, Denisia, 5: 712, Fig. 158a–w, 381l, Table 41 (Fig. 41a–n, 42a–j; original description, see remarks; the holotype slide [accession number 2002/289] and three paratype slides [2002/290–292] as well as four voucher slides [2002/293–296] from Namibian site 1 population and three voucher slides [2002/297–299] from Australian population are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; Foissner et al. 2002, p. 40 and Aescht 2003, p. 384).
Nomenclature: The species-group name decorat·us, -a, -um (Latin adjective [m; f; n]; ornamented) refers to the granule plaques (Foissner et al. 2002). Remarks: We classified this species in Lamtostyla because stomatogenesis commences near the transverse cirri, as in L. australis (Foissner et al. 2002). See genus section for a more detailed foundation of the generic assignment. Lamtostyla decorata is likely closely related to L. granulifera, L. longa, and L. raptans because they have the same number and arrangement of frontoventral cirri and pretransverse and transverse cirri (see L. granulifera and L. longa groups in remarks of genus section). It differs from (i) L. granulifera by body shape (elongate and often twisted about main body axis vs. slenderly to rather broadly elliptical), cortical granules (0.3 µm across and packed around dorsal bristles vs. 1–2 µm across forming many closely spaced rows), number of right and left marginal cirri (36 and 32 vs. 44 and 45), and the width of the adoral membranelles (6 µm vs. 12 µm); (ii) L. longa by body shape (elongate and often twisted about main body axis vs. slenderly to rather broadly elliptical), number of right and left marginal cirri (36 and 32 vs. 23 and 21), number of dorsal kineties (three vs. five), and by the location of the buccal cirrus (about 8 µm behind anterior end of paroral vs. at front end); and (iii) L. raptans by size (100–170 × 20–40 µm vs. 200 × 40 µm), body shape (see L. longa above), number of adoral membranelles (19–22 vs. 33–36), and number of dorsal kineties (three vs. five). Possibly, Lamtostyla longa and L. raptans differ from L. decorata also by the lack of cortical granules. However, such granules might have been overlooked by Hemberger (1985), who did not study live specimens in detail. In life, Lamtostyla decorata is characterised by the slender, usually twisted body, the cortical granules plaques around dorsal bristles, and the short amphisiellid median cirral row hardly extending above buccal vertex (Foissner et al. 2002). The three populations described by Foissner et al. (2002) – all studied independently because originally classified as distinct species – differ considerably in some morphometrics and morphological details, suggesting subspecies ranks. However, a 1
Foissner et al. (2002) provided the following diagnosis (includes only data from type material): Size about 140 × 25 µm in vivo. Oblong and usually twisted about main body axis. Cortical granules colourless, about 0.3 µm across, form conspicuous plaques around dorsal bristles. On average 2 macronuclear nodules, 32 cirri in left and 36 in right marginal row, 4 cirri in amphisiellid median cirral row (ACR), 3 cirri left of ACR, 1 buccal cirrus at summit of strongly curved undulating membranes, 7 transverse cirri, and 3 dorsal kineties. Adoral zone continuous, on average consists of 20 membranelles, with largest bases about 6 µm wide. Buccal cavity of ordinary width, deep.
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biogeographic pattern is hardly recognisable and the cirral pattern is very similar in all populations. Moreover, the number of adoral membranelles is rather similar, and the taxonomic value of some other features (e.g., shape of micronuclei) not known. Thus, Foissner et al. (2002) preliminarily submerged all populations in a single species. The descriptions of further populations should be awaited before splitting. Morphology: We studied three populations, namely two from Namibia (sites [1] and [31; type population]) and one from Australia. They agree in most, but not all features. Thus, data are kept separate and the following detailed description contains only data from the type population. For deviating data from the other two populations, see below. Body size 100–170 × 20–35 µm in life, body length:width ratio about 5–6:1 (range 4–9:1!) in life, on average 4.8:1 in protargol preparations, where cells are slightly inflated (Table 22). Body slender, often slightly sigmoidal, usually curved and/or even helical by half a turn about main body axis, distinctly flattened mainly in oral region (Fig. 41d, j, m); very flexible, but acontractile, fragile and thus rather distorted in the preparations. Two macronuclear nodules slightly left of midline, ellipsoidal, sometimes globular, with many medium-sized and small chromatin bodies; rarely specimens with four nodules occur. Micronuclei globular, attached to macronuclear nodules at variable positions. Contractile vacuole near left cell margin slightly ahead of mid-body, without distinct collecting canals. Cortical granules scattered around bases of cirri and in conspicuous plaques around dorsal bristles; individual granules inconspicuous because only about 0.3 µm across and colourless, neither impregnate with protargol nor stain or extrude when methyl green-pyronin is added (Fig. 41e, f, n). Cytoplasm colourless with many fat, refractive globules 1–10 µm across, specimens thus dark at low magnification. Movement conspicuous because slow and serpentine. Adoral zone occupies only 14–23%, on average 19% of body length, composed of about 20 membranelles, bases of largest membranelles about 6 µm wide in life. Buccal cavity deep and moderately wide, anterior cavity margin Cyrtohymena-like, that is, strongly curved; right margin forms prominent lip, covers cavity and proximal adoral membranelles. Undulating membranes distinctly curved, both likely composed of closely spaced dikinetids and optically intersecting near mid. Pharyngeal fibres, likely mixed with long endoral cilia, prominent in life and after protargol impregnation, of ordinary length and structure, extend obliquely backwards (Fig. 41a, b, d). Cirral pattern and number of cirri of usual variability (Fig. 41a–d, m; Table 22). Frontal cirri slightly to distinctly enlarged, right one, as is usual, behind distal end of adoral zone. Buccal cirrus close to summit of paroral. Usually three cirri left of amphisiellid median cirral row, including cirrus III/2, which is often slightly enlarged and somewhat dislocated to left and thus behind right frontal cirrus. Amphisiellid median cirral row usually composed of only four cirri, commences right of right frontal cirrus and terminates at 21% of body length on average; thus, row only slightly longer than adoral zone; distance between second and third cirrus usually
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Fig. 41n Lamtostyla decorata (from Foissner et al. 2002. Interference contrast). The colourless cortical granules are only 0.3 µm across, but form conspicuous plaques mainly around dorsal bristles. CG = cortical granules, RMR = right marginal row. Page 212.
slightly enlarged (this marks the margin of the two portions [2 + 2 cirri] of which the row is composed; Fig. 41d). Transverse cirri about 15 µm long in life, near rear body end and thus distinctly projecting; form Ushaped pattern with one (pretransverse) cirrus in U-cavity (Fig. 41b, m). Marginal cirri only about 8 µm long in life. Right marginal row commences dorsolaterally near front body end, ends subterminal; left row commences near proximal end of adoral zone and slightly shortened posteriorly, thus, marginal rows distinctly separated posteriorly. Dorsal bristles about 3 µm long in life, arranged in three kineties; kinety 1 distinctly shortened anteriorly. Caudal cirri absent (Fig. 41c, d, m).
b
Fig. 41a–m Lamtostyla decorata (type population; from Foissner et al. 2002. a, e–l, from life; b–d, m, protargol impregnation). a: Ventral view of representative specimen, 132 µm. b, c: Infraciliature of ventral and dorsal side and nuclear apparatus of holotype specimen, 138 µm. Short arrows mark anteriormost two cirri (= “frontoterminal cirri”) of amphisiellid median cirral row, long arrow denotes pretransverse ventral cirrus in U-cavity of transverse cirri. Arrowhead marks cirrus III/2. d, m: Infraciliature of ventral and dorsal side and nuclear apparatus of a strongly twisted (about 180°) specimen, 114 µm. Arrows mark left marginal row. Frontal cirri connected by dotted line; frontal and ventral cirri which very likely originate from the same frontal-ventral-transverse cirri anlage are connected by broken lines (has to be checked by ontogenetic data). e, f: Tiny (0.3 µm across), colourless granules occur around cirri and dorsal bristles, where they form conspicuous plaques. g, j–l: Shape variants. h, i: Same specimen in ventral and lateral view, showing that L. decorata is usually unflattened. ACR = amphisiellid median cirral row (composed of four cirri only), BC = buccal cirrus, FC = right frontal cirrus, RMR = right marginal row, TC = transverse cirri, II–VI = frontal-ventral-transverse cirri anlagen, 1–3 = dorsal kineties. Page 212.
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Observations on Namibian site (1) specimens (Fig. 42a–j; Table 22): The Namibian site (1) specimens are distinctly larger than those from the type locality (length 140–220 µm vs. 100–170 µm in life) and thus many morphometric features are considerably different, especially the almost doubled number of right and left marginal cirri. But there are also rather distinct differences in some morphological features: (i) the micronuclei are conspicuously ellipsoidal, sometimes almost cylindrical; (ii) the cortical granules around the bases of the cirri and dorsal bristles are a mixture of small (about 0.5 µm) and large (up to 2.0 µm) globules, as in the Australian specimens; and (iii) the distal 1–3 µm of all cilia/cirri and dorsal bristles are heavily argyrophilic, a curious property we never observed in any other hypotrich (Fig. 42d, j). Observations on an Australian population (Table 22): A morphometric analysis of the Australian specimens, which have a size of about 110 × 20 µm in life, reveals two rather distinct differences to the type population, namely, (i) 6–8, on average seven against 4–5, on average four cirri in the amphisiellid median cirral row; and (ii) 5–6, on average five against 6–9, on average seven transverse cirri. Furthermore, the cortical granules are larger (1.0–1.5 µm vs. 0.3 µm across) and the plaques (granules groups) not confined to the ciliature, but occur throughout the cortex. Cell division: Ontogenesis commences with the formation of a very narrow oral primordium extending from near the transverse cirri to the buccal vertex. Concomitantly, the posterior cirri of the amphisiellid median cirral row modify to anlagen. Occurrence and ecology: Very likely confined to terrestrial habitats. Type locality of L. decorata is the Welwitschia Drive (22°45'S 15°25'E; site 31 in Foissner et al. 2002) in the central Namib Desert, Namibia, where it was discovered in litter from Welwitschia mirabilis. Foissner et al. (2002) found a rather deviating population (Fig. 42a–j) at Namibian site (1), a wet, reddish, very sandy soil covered with a moist algal mat in a highland savannah at the west margin of the town of Windhoek. In addition, it occurred in dark grey mud from the Bambatsi Guest Farm between the towns of Khorixa and Outjo (site 49 in Foissner et al. 2002). The Australian population inhabited a highly saline soil from the shore of Lake Amadeus near the town of Alice Springs. In Austria, we found it in a soil from Salzburg (not studied in detail), indicating that L. decorata is a euryoecious, euryhaline cosmopolitan (Foissner et al. 2002). Abundance was low in all samples.
b
Fig. 42a–j Lamtostyla decorata (Namibian site 1 population; from Foissner et al. 2002. a, e, f, from life; b–d, g–j, protargol impregnation). a: Ventral view of a representative specimen, 167 µm. b, c: Infraciliature of ventral and dorsal side and nuclear apparatus showing the conspicuous micronuclei, 145 µm. Arrow marks pretransverse ventral cirrus in U-cavity formed by transverse cirri. d, j: The dorsal bristles (d) as well as cilia of cirri and paroral (j) have strongly argyrophilic distal end (arrows). e, f: The cortical granules, which are colourless and 0.5–2.0 µm across, form conspicuous plaques around bases of dorsal bristles (e). g, h: Ventral and dorsal view of posterior body region showing lack of caudal cirri and Ushaped arrangement of transverse cirri. i: The buccal cavity is covered by the very hyaline buccal lip, which is roofed by the paroral cilia. BL = buccal lip, F = fibres, LMR = left marginal row, MI = micronucleus, TC = transverse cirri, 1–3 = dorsal kineties. Page 212.
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The specimens of the type population feed on bacteria and Pseudocohnilembus sp., a ciliate, interestingly, also mainly ingested by the other populations. The Namibian site (1) specimens, however, ingested the flagellate Polytomella and mediumsized ciliates, such as Colpoda maupasi and Urosomoida agiliformis.
Lamtostyla longa-group Species of this group (L. longa, L. raptans) have the same 18-cirri pattern in the frontal area as the species of the L. granulifera-group which comprises L. granulifera and L. decorata. However, the species included in the L. longa-group have five dorsal kineties against three in the L. granulifera-group.
Lamtostyla longa (Hemberger, 1985) Berger & Foissner, 1988 (Fig. 43a–h, Table 22) 1982 Tachysoma longa n. spec.1 – Hemberger, Dissertation2, p. 249, Abb. 45a–c (Fig. 43a, b, e–h; revision of hypotrichs). 1985 Tachysoma longa n. spec. – Hemberger, Arch. Protistenk., 130: 412, Abb. 19 (Fig. 43a, b; original description; no formal diagnosis provided; type slides are deposited in the Institut für landwirtschaftliche Zoologie, University of Bonn, Germany). 1988 Lamtostyla longa (Hemberger, 1985) nov. comb. – Berger & Foissner, Zool. Anz., 220: 116, Fig. 1, Table 1 (Fig. 43a, b; combination with Lamtostyla; revision of Lamtostyla). 2001 Lamtostyla longa (Hemberger, 1985) Berger and Foissner, 1988 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 88 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs).
Nomenclature: No derivation of the name is given in the original description. The species-group name long·us, -a, -um (Latin adjective [m; f; n]; long, wide) likely alludes to elongated body shape. Fig. 43a–h Lamtostyla longa (a, b, after Hemberger 1985; e–h, from Hemberger 1982. Protargol impregnation). a, b: Infraciliature of ventral side and nuclear apparatus, 103 µm. Cirri left of amphisiellid median cirral row circled. c, d: Anterior and posterior portion of specimen shown in (a). Lamtostyla longa has 18 frontal-ventral-transverse cirri, which very likely originate (as in the 18-cirri hypotrichs) from six anlagen (has to be confirmed by ontogenetic data; designation according to the numbering system by Wallengren 1900. For details see general section and Berger 1999, p. 16). Arrows mark rearmost basal body pairs of dorsal kineties. Cirrus II/2 is the buccal cirrus. e, f: Infraciliature of ventral side and nuclear apparatus of variant, which lacks the elongated dorsal bristles at rear end of kineties; size not indicated. g, h: Infraciliature of ventral side and nuclear apparatus of an early to middle divider, size not indicated. ACR = amphisiellid median cirral row, AZM = adoral zone of membranelles, FC = frontal cirri (connected by dotted line), FT = frontoterminal cirri, MA = macronuclear nodules, MI = micronucleus, PVC = postoral ventral cirri, PTVC = pretransverse ventral cirri, TC = transverse cirri, I–VI = frontal-ventral-transverse cirri streaks (anlagen, primordia), 1–4 = cirri within an anlage (from posterior to anterior). Page 218. 1 2
This name is disclaimed for nomenclatural purposes (ICZN 1999, Article 8.3). See next footnote. See same footnote at Uroleptoides binucleatus.
d
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Remarks: The original description lacks detailed live data. Thus, some important features (e.g., presence/absence of cortical granules) are lacking. Hemberger (1982, 1985) assigned the present species to Tachysoma, an oxytrichid genus, because it lacks distinct caudal cirri. However, the rearmost dorsal bristles are longer than the remaining bristles so that a classification in Oxytricha (with caudal cirri) would also have been possible (for review of oxytrichids, see Berger 1999). Hemberger (1985) found specimens which lack the elongated dorsal bristles, indicating that similar species exist (Fig. 43e, f). Berger & Foissner (1988) transferred the present species to Lamtostyla because the arrangement of the cirri in the frontal area closely resembles that of L. perisincirra, L. islandica, and the type species L. lamottei. Most Lamtostyla species have a low number (2–4, usually three) of dorsal kineties and – more important – lack a dorsomarginal kinety or fragmentation of a kinety. By contrast, the present species has “4 plus 1/2” kineties, indicating that at least one dorsomarginal kinety or a fragmentation occurs. The presence of 18 frontal-ventral-transverse cirri is no indicator that it is an oxytrichid because this pattern very likely occurred in the last common ancestor of the Hypotricha for the first time. The postoral ventral cirri are displaced anteriad so that the highly characteristic 18-cirri pattern is difficult to recognise (Fig. 43a, c, d). However, ontogenetic data are needed for a more detailed discussion and a proper classification. Thus, I preliminarily retain the classification in Lamtostyla. Lamtostyla raptans, which occurred in the same locality, is very similar to the present species, except that it is larger (200 × 40 µm) and therefore has more adoral membranelles and marginal cirri (Table 22). Further, they differ in the undulating membranes and transverse cirri pattern (compare Fig. 43a, 44a). Lamtostyla granulifera, which has basically the same cirral pattern as L. longa, has only three dorsal kineties. Unfortunately, the cortical granules of L. granulifera cannot be used as distinguishing feature because the presence/absence of these organelles was not checked in L. longa (see above). For a characterisation of the various Lamtostyla-groups, see remarks at genus section. Morphology: Body size 85–130 × 35–50 µm in life(?), body length:width ratio about 3:1. Body margins almost in parallel, both ends rounded. Body very sensitive, and likely flexible. Two macronuclear nodules likely in common position, that is, slightly left of midline about in body centre. Each macronuclear nodule with one micronucleus. Contractile vacuole in “standard position”, that is, near left cell margin about in mid-body (or slightly ahead of it). Nothing known about cortical granules (present or absent), cytoplasm (colour, inclusions ...), and movement. Adoral zone occupies about 20–25% of body length, composed of 18–19 membranelles of ordinary fine structure, always conspicuously bent. Buccal field likely of ordinary size. Undulating membranes rather short, slightly curved and arranged almost in parallel (Fig. 43a). Cirral pattern as shown in Fig. 43a. The 18-frontal-ventral-transverse cirri pattern is difficult to recognise due to the anteriorly dislocated postoral ventral cirri (see Fig. 43c, d for interpretation and homologisation). Three slightly enlarged fron-
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tal cirri near distal portion of adoral zone. Buccal cirrus right of anterior end of undulating membranes. Three cirri (including cirrus III/2) left of amphisiellid median cirral row, which is composed of four cirri. Two pretransverse ventral cirri ahead of five transverse cirri, which are slightly enlarged and about 15 µm long. Marginal cirri about 12 µm long; right marginal row commences about at level of buccal cirrus, terminates – like left row – about at level of transverse cirri. Left marginal row commences close to proximal end of adoral zone; marginal rows distinctly separated posteriorly; however, gap occupied by rearmost, elongated dorsal bristles (see below). Dorsal bristles 3–4 µm long, arranged in four bipolar(?) kineties and a half kinety (exact arrangement and ontogenesis not known, preventing a more proper classification; see remarks). Caudal cirri lacking, both basal bodies of rearmost bristle complex, however, with 13 µm long cilia (vestigial caudal cirri?). Cell division: Hemberger (1982) found an early to middle divider, showing that the 18 frontal-ventral-transverse cirri originate from the ordinary six (I–VI) anlagen (Fig. 43g, h). Occurrence and ecology: Very likely confined to terrestrial habitats (Foissner 1987, p. 127; 1998, p. 205). Type locality of L. longa is the area near the village of Puerto Maldonado (about 69°12'W 12°36'S), Department Madre de Dios, Peru, where Hemberger (1982, 1985) discovered it in forest soil. No further records published. Food not known. Biomass of 106 specimens about 135 mg (Foissner 1987, p. 127).
Lamtostyla raptans (Hemberger, 1985) Foissner, 1997 (Fig. 44a, b, Table 22) 1982 Tachysoma raptans n. spec.1 – Hemberger, Dissertation2, p. 253, Abb. 47 (Fig. 44a, b; revision of hypotrichs). 1985 Tachysoma raptans n. spec. – Hemberger, Arch. Protistenk., 130: 414, Abb. 21 (Fig. 44a, b; original description; no formal diagnosis provided; type slide is deposited in the Institut für landwirtschaftliche Zoologie, University of Bonn, Germany). 1997 Lamtostyla raptans (Hemberger, 1985) nov. comb. – Foissner, Biol. Fertil. Soils, 25: 334 (combination with Lamtostyla). 2001 Lamtostyla raptans (Hemberger, 1985) Foissner, 1997 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 88 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs).
Nomenclature: No derivation of the name is given in the original description. The species-group name raptans (Latin; predate) implies that the present species is (very likely) a predator. Remarks: The original description lacks detailed live data. Thus, some important features (e.g., presence/absence of cortical granules) are lacking. Hemberger 1
This name is disclaimed for nomenclatural purposes (ICZN 1999, Article 8.3). See footnote 2 at L. longa. 2 See same footnote at Uroleptoides binucleatus.
222
SYSTEMATIC SECTION Fig. 44a, b Lamtostyla raptans (after Hemberger 1985. Protargol impregnation). Infraciliature of ventral side and nuclear apparatus, 200 µm. Short arrow marks buccal cirrus, long arrow denotes fibres associated with pretransverse ventral cirrus. Frontal cirri connected by dotted line; cirri left of amphisiellid median cirral row circled; right frontal cirrus and cirrus III/2 connected by broken line. Note that the amphisiellid median cirral row is (very likely) composed of two portions, namely, the frontoterminal cirri (anterior two cirri) and the cirri originating from anlage V (details on terminology see general section). The ventral cirral pattern of this species resembles that of L. longa, L. granulifera, and L. decorata (see remarks at genus section for grouping of these four species). ACR = amphisiellid median cirral row, AZM = adoral zone of membranelles (note spoon-like broadening of proximal portion; also present in L. granulifera), CV = contractile vacuole, FC = right frontal cirrus, LMR = left marginal row, MA = macronuclear nodule, MI = micronucleus, RMR = right marginal row, TC = transverse cirri. Page 221.
(1982, 1985) assigned the species provisionally to Tachysoma because the cirral pattern is reminiscent of many 18-cirri oxytrichids (except for the anteriorly dislocated postoral ventral cirri, a feature also occurring in Gonostomum) and caudal cirri are lacking (for review on oxytrichids, see Berger 1999). He noted that the classification in Tachysoma is unsatisfactory because the marginal rows of the present species are only indistinctly separated posteriorly. However, not the distance between the rear end of the marginal rows is so important, but the lack of the caudal cirri which is characteristic for Tachysoma. Hemberger also stated that a new genus could be established if similar species will be found. Lamtostyla longa, also described by Hemberger (1985), is such a similar species. Thus, they are placed in the L. longa-group (see genus section). However, the establishment of a new (sub)genus is avoided because some important data (exact arrangement of dorsal kineties and their formation) are not known. Foissner (1997) transferred the present species to Lamtostyla because it highly resembles Lamtostyla granulifera described by himself, especially in the cirral pattern, the nuclear apparatus, and the spoon-like broadening of the adoral zone of membranelles. These two species differ in body size (200 µm long vs. 143 µm; Table 22), number of adoral membranelles (33–36 vs. 23–27), number of marginal cirri
Lamtostyla
223
(around 60 in both rows in L. raptans vs. around 44 in L. granulifera), and dorsal kineties (five vs. three), and, possibly, by the cortical granules (lacking vs. present). However, such granules might have been overlooked by Hemberger, who did not study live specimens in detail. Interestingly, the present species, L. longa, and L. granulifera occur in terrestrial habitats of South America, indicating that this is the major area of distribution of this group. Most Lamtostyla species have 2–4, usually three, dorsal kineties. By contrast, Lamtostyla raptans has, like the related L. longa, five dorsal kineties. Ontogenetic data are needed to show whether or not dorsomarginal rows and/or fragmenting kineties are present in L. raptans. If such specific rows are present then the classification in Lamtostyla, respectively, the amphisiellids is incorrect (see Amphisiellidae). However, so far the classification in Lamtostyla seems to be the most parsimonious solution. Morphology: Body size 200 × 40 µm in life(?), body length:width ratio about 5:1. Body margins slightly converging posteriorly, that is, body widest anteriorly, both ends rounded. Body very flexible. Two macronuclear nodules likely in common position, that is, slightly left of midline about in body centre. Each macronuclear nodule with one micronucleus. Contractile vacuole relatively large, near left cell margin about at 33% of body length. Nothing known about cortical granules (present or absent), cytoplasm (colour, inclusions ...), and movement. Adoral zone occupies about 25% of body length, composed of 33–36 membranelles of ordinary fine structure, proximal portion broadened spoon-like. Buccal field likely relatively large. Undulating membranes of about equal length, curved, optically slightly intersecting, and rather long (Fig. 44a). Cirral pattern as shown in Fig. 44a. It is basically an 18-frontal-ventraltransverse cirri pattern with two specific features, namely, (i) the postoral ventral cirri are dislocated anteriorly (see L. longa for interpretation); and (ii) the left pretransverse ventral cirrus is lacking. Frontal cirri large, near distal portion of adoral zone with right cirrus, as is usual, behind distal end of zone. Buccal cirrus right of anterior portion of undulating membranes. Three cirri (including cirrus III/2) left of “amphisiellid median cirral row”, which is composed of four cirri; however, anterior two cirri (= frontoterminal cirri) not exactly in line with posterior two cirri, that is, amphisiellid median cirral row indistinct. One pretransverse ventral cirrus ahead of right transverse cirrus. Transverse cirri terminal, slightly enlarged, about 15 µm long. Marginal cirri about 12–15 µm long; right marginal row commences near distal end of adoral zone, ends about at level of transverse cirri. Left marginal row commences left of proximal portion of adoral zone, terminates at rear end in cellmidline; marginal rows thus only slightly separated posteriorly. Dorsal bristles 4–5 µm long, arranged in five bipolar(?) kineties (exact arrangement and ontogenesis not known, preventing a more proper classification; see remarks). Caudal cirri lacking. Occurrence and ecology: Very likely confined to terrestrial habitats (Foissner 1987, p. 127; 1998, p. 205). Type locality of L. raptans is the area near the village of
224
SYSTEMATIC SECTION
Puerto Maldonado (about 69°12'W 12°36'S), Department Madre de Dios, Peru, where Hemberger (1982, 1985) discovered it in a forest soil. No further records published. Food not known, likely predacious as indicated by the species-group name. Biomass of 106 specimens about 150 mg (Foissner 1987, p. 127).
Uroleptoides Wenzel, 1953 1953 Uroleptoides kihni nov. gen. nov. spec.1 – Wenzel, Arch. Protistenk., 99: 107 (original description). Type species (by original designation and monotypy): Uroleptoides kihni Wenzel, 1953. 1974 Uroleptoides Wenzel, 1953 – Stiller, Annls hist.-nat. Mus. natn. hung., 66: 130 (brief revision). 1974 Uroleptoides Wenzel – Stiller, Fauna Hung., 115: 61 (guide to Hungarian ciliates). 1977 Uroleptoides Wenzel – Corliss, Trans. Am. microsc. Soc., 96: 137 (classification of ciliates). 1979 Uroleptoides Wenzel, 1953 – Tuffrau, Trans. Am. microsc. Soc., 98: 525 (brief revision). 1979 Uroleptoides Wenzel, 1953 – Corliss, Ciliated protozoa, p. 310 (revision). 1982 Uroleptoides Wenzel, 19532 – Hemberger, Dissertation, p. 49 (revision of hypotrichs). 1983 Uroleptoides Wenzel, 1953 – Curds, Gates & Roberts, Synopses of the British fauna, 23: 418 (guide to British ciliates). 1985 Uroleptoides – Small & Lynn, Phylum Ciliophora, p. 456 (guide to ciliate genera). 1987 Uroleptoides Wenzel, 1953 – Tuffrau, Annls Sci. nat. (Zool.), 8: 115 (brief revision of hypotrichs). 1994 Uroleptoides Wenzel, 1953 – Tuffrau & Fleury, Traite de Zoologie, 2: 137 (revision of hypotrichs). 1999 Uroleptoides Wenzel, 1953 – Shi, Acta Zootax. sinica, 24: 253 (brief revision of hypotrichs). 1999 Uroleptoides Wenzel, 1953 – Shi, Song & Shi, Progress in Protozoology, p. 100 (brief revision of hypotrichs). 2001 Uroleptoides Wenzel 1953 – Aescht, Denisia, 1: 170 (catalogue of generic names of ciliates). 2001 Uroleptoides Wenzel, 1953 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 96 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Uroleptoides Wenzel, 1953 – Lynn & Small, Phylum Ciliophora, p. 453 (guide to ciliate genera). 2005 Uroleptoides – Berger, Int. Congr. Protozool., 12: 106 (brief note on amphisiellids). 2006 Uroleptoides Wenzel, 1953 – Berger, Monographiae Biol., 85: 1213 (brief note on exclusion from urostyloids).
Nomenclature: No derivation of the genus-group name is given in the original description. Uroleptoides is a combination of Uroleptus (a composite of the Greek noun he ura [tail] and the Greek adjective lept- [slender, small]) and the suffix -oides (meaning “like Uroleptus”). According to Article 30.1.4.4 of the ICZN (1999) a compound genus-group name ending with -oides is to be treated as masculine, unless its author, when establishing the name, stated that it had another gender or treated it as such by combining it with an adjectival species-group name in another gender form. Wenzel (1953) dedicated the type species (U. kihni) to Berthold Kihn 1
Wenzel (1953) provided the following diagnosis: Oxytrichidae mit langgestrecktem, caudal verjüngtem Körper, 2 ± randständigen, bis zum Körperende laufenden Marginalreihen, einer das Peristomfeld nach rechts begrenzenden, bis zum Körperende führenden, einfachen Ventralreihe. Auf dem Stirnrand des Frontalfeldes 3 starke Frontalcirren, dahinter Gruppen weiterer, schwächer ausgebildeter Cirren. 2 Hemberger (1982) provided the following diagnosis: Je 1 linke und rechte Maginalreihe; 1 Ventralreihe; keine oder unscheinbare Transversalcirren; deutlich differenzierte Frontalcirren, die sich aus 2 oder 3 longitudinalen Anlagen entwickeln; Köper langgestreckt, meist zahlreiche Makronuklei.
Uroleptoides
225
(male) so that Uroleptoides remained masculine. By contrast, Aescht (2001) assumed that Uroleptoides is feminine although referring to the same Article. Characterisation (A = supposed apomorphy): Amphisiellidae with continuous adoral zone of membranelles. Three more or less distinctly enlarged frontal cirri. Buccal cirrus present. Two or more cirri left of anterior portion of amphisiellid median cirral row which terminates usually behind mid-body. Postperistomial cirrus lacking. Transverse cirri present. One right and one left marginal row. Usually three dorsal kineties; dorsomarginal row, kinety fragmentation, and caudal cirri lacking. Terrestrial. Remarks: The characterisation above is rather vague because the cirral pattern of the type species is described only from life and shows a rather curious feature, namely, the presence of a second pseudorow of frontal cirri (Fig. 45a, arrows). Such a pattern, which has to be confirmed by protargol preparations, is difficult to explain without ontogenetic data and is perhaps due to a misinterpretation. For example, in Fig. 54e of Uroleptoides terricola a similar pseudorow1 is present, but shifted slightly rightwards. Anyhow, this feature is preliminarily not included in the characterisation and should not be overinterpreted at the present state of knowledge. On the other hand, the general appearance of U. kihni is reminiscent of slender, terrestrial amphisiellids because of the long frontoventral row which can be interpreted as amphisiellid median cirral row although we do not know whether it originates from two or three anlagen as in amphisiellids or from only a single anlage as in nonamphisiellids. Interestingly, Uroleptoides kihni was never found since its original description although many hundred Central European soil samples have been investigated (for review, see Foissner 1998). This can be explained at least in two ways: (i) Uroleptoides kihni is an extremely rare species not yet found again; or (ii) it is a species indeterminata, that is, the description and illustration provided by Wenzel (1953) are so misleading that U. kihni cannot be recognised, and perhaps it is identical with an other species. I prefer the first explanation and recommend searching some more years. Uroleptopsis citrina, the type species of Uroleptopsis, for example, had to wait 70 years until it was redescribed for the first time (Kahl 1932, Berger 2004a, 2006). Wenzel (1953) established Uroleptoides because the groups of weaker cirri on the frontal area of U. kihni prevented a classification in Uroleptus or Paruroleptus. However, in Uroleptus and its supposed synonym Paruroleptus the ventral cirri are zigzagging (that is, they originate from many anlagen), whereas the ventral cirri of Uroleptoides kihni obviously form a continuous row which very likely originates from one, two, or three anlagen. Thus, a close relationship of Uroleptoides and Uroleptus is unlikely. Stiller (1974, 1974a), Corliss (1977, 1979), Curds et al. (1983), and Foissner (1987a) retained this “holostichid” classification.
1
This pseudorow is formed by the buccal cirrus, cirrus III/2 (arrow in Fig. 54e), and the anteriormost cirrus of anlage IV.
226
SYSTEMATIC SECTION
Table 22 Morphometric data on Amphisiella australis sensu Foissner (1988; au4, au5, population I and II from Foissner 1988), Lamtostyla australis (au1, type population from Blatterer & Foissner 1988; au2, population II from Blatterer & Foissner 1988; au3, from Voß 1992), Lamtostyla decorata (de1, type population; de2, Australian population; de3, Namibian site 1 population; from Foissner et al. 2002), Lamtostyla elegans (el1–el3, type population; el1, specimens with four macronuclear nodules; el2, specimens with 5–8, usually six macronuclear nodules; el3, all specimens combined; from Foissner et al. 2002), Lamtostyla granulifera (gra, from Foissner 1997), Lamtostyla islandica (isl, from Berger & Foissner 1988), Lamtostyla longa (lon, from Hemberger 1985), Lamtostyla perisincirra (pe1, from Berger et al. 1984; pe2, from Hemberger 1982, 1985), Lamtostyla procera (pr1, type population; pr2, Namibian site 9 population; from Foissner et al. 2002), Lamtostyla quadrinucleata (qua, from Berger & Foissner 1989), Lamtostyla raptans (rap, from Hemberger 1985), Lamtostyla vitiphila (vit, from Foissner 1987a), Lamtostylides edaphoni (ed1, type population from Berger & Foissner 1987; ed2, raw culture of population I; ed3, pure culture of population I; ed4, raw culture of population II; ed2–ed4, from Petz & Foissner 1996), Lamtostylides halophilus (ha1, type population from site 18; ha2, population from Namibian site 69; from Foissner et al. 2002), Lamtostylides hyalinus (hya, from Berger et al. 1984), Lamtostylides kirkeniensis (kir, from Berger & Foissner 1988), Lamtostylides pori (por, from Wilbert & Kahan 1986), Uroleptoides binucleatus binucleatus (bi1, from Hemberger 1982, 1985; bi2, from Berger & Foissner 1989), Uroleptoides binucleatus multicirratus (bi3, Nambian site 5 population [= type population]; bi4, Namibian site 41 population; from Foissner et al. 2002), Uroleptoides longiseries (lo1, type population from Namibia; lo2, population from Israel; from Foissner et al. 2002), Uroleptoides magnigranulosus (mag, from Foissner 1988), Uroleptoides multinucleatus (mu1, type population; mu2, Namibian site 52 population; mu3, Kenyan population; from Foissner et al. 2002), Uroleptoides polycirratus (pol, from Berger & Foissner 1989), and Uroleptoides terricola (te1, from Hemberger 1982; te2, from Foissner 1984) Characteristics a Body, length
Species
mean
M
SD
SE
CV
au1 au2 au3 k au3 au4 au5 bi1p bi2 bi3 bi4 de1 de2 de3 ed1 ed2 ed3 ed4 el1 el2 el3 gra ha1 ha2 hya isl kir lo1
100.1 84.5 96.8 92.7 119.7 104.2 – 140.6 213.1 234.2 123.4 88.7 158.6 61.1 76.8 79.8 57.5 155.0 153.3 154.4 143.4 58.9 66.5 31.4 62.3 82.6 179.2
97.0 85.5 96.9 90.9 118.0 110.0 – 140.0 207.0 232.0 125.0 86.0 155.0 62.0 78.0 81.0 56.0 150.0 154.0 150.0 143.0 58.0 65.0 32.0 63.0 84.0 174.0
13.7 7.6 14.8 11.0 14.0 12.4 – 12.7 31.7 29.1 18.0 9.2 20.0 6.7 10.4 7.4 3.5 18.8 11.1 16.1 9.7 5.9 6.2 3.3 4.6 5.2 21.3
3.8 2.4 3.0 2.2 4.1 3.4 – 4.0 8.2 7.5 3.8 2.8 4.7 2.1 1.9 2.1 1.1 5.0 3.9 3.4 2.2 1.3 1.9 1.1 – – 5.9
13.7 9.0 15.3 11.9 11.8 11.2 – 9.0 14.9 12.4 14.6 10.4 12.6 11.0 13.5 9.3 6.1 12.1 7.3 10.5 6.8 10.0 9.3 10.6 7.4 6.3 11.9
Min
Max
n
77.0 72.0 78.7 74.5 98.0 87.0 220.0 126.0 150.0 186.0 90.0 72.0 130.0 49.0 55.0 62.0 52.0 126.0 140.0 126.0 125.0 50.0 60.0 26.0 56.0 72.0 152.0
127.0 94.0 126.7 111.8 145.0 122.0 260.0 168.0 265.0 290.0 154.0 102.0 200.0 69.0 109.0 92.0 63.0 190.0 170.0 190.0 160.0 70.0 77.0 35.0 70.0 90.0 224.0
13 10 25 25 12 13 10 10 15 15 23 11 18 10 31 13 11 14 8 22 19 21 11 10 12 11 13
Uroleptoides
227
Table 22 Continued Characteristics a Body, length
Body, width
Species
mean
M
SD
SE
CV
lo2 mag mu1 mu2 mu3 pe1 pol por k pr1 pr2 qua te1 te2 vit au1 au2 au3 k au3 au4 au5 bi1 p bi2 bi3 bi4 de1 de2 de3 ed1 ed2 ed3 ed4 el1 el2 el3 gra ha1 ha2 hya isl kir lo1 lo2 mag mu1 mu2 mu3 pe1 pol por k
160.0 104.2 259.4 173.1 207.8 42.9 150.0 95.5 129.3 138.9 86.3 – 92.6 89.1 28.8 27.1 35.8 32.4 40.5 32.6 50.0 40.9 51.5 61.4 26.2 23.3 41.2 15.9 27.4 30.8 17.3 29.6 35.4 31.7 64.5 25.6 25.2 13.7 18.3 20.6 40.3 43.0 46.6 34.0 26.7 64.3 15.2 46.7 16.1
162.4 104.5 281.0 168.0 210.0 42.0 150.0 95.0 126.0 145.0 84.0 – 91.0 88.0 29.0 27.0 37.3 29.8 40.5 32.0 – 41.0 52.0 60.0 26.0 23.5 40.0 16.0 27.0 30.0 17.0 29.0 35.0 30.0 65.0 25.0 25.0 14.0 18.0 21.0 38.0 42.0 45.5 32.0 26.0 63.0 15.0 46.0 17.0
15.8 10.5 43.7 19.3 20.5 5.1 10.0 7.7 16.7 25.3 10.1 – 8.9 8.2 2.3 1.9 6.7 6.0 4.9 6.9 – 6.0 8.7 9.8 3.7 2.8 5.5 1.5 5.0 4.0 1.9 5.2 5.6 6.0 9.0 2.9 5.6 2.2 18.0 1.2 9.3 7.7 4.9 6.3 3.3 5.1 1.7 5.0 1.6
5.6 3.0 19.5 4.2 5.7 1.0 5.8 2.2 4.3 9.5 2.6 – 4.0 – 0.6 0.6 1.3 1.2 1.4 1.9 – 1.9 2.2 2.5 0.8 1.4 1.3 0.5 0.9 1.1 0.6 1.4 2.0 1.3 2.1 0.6 1.7 0.7 – – 2.6 2.7 1.4 2.8 0.7 1.4 0.3 2.9 0.4
9.7 10.1 16.8 11.2 9.9 11.8 6.7 – 13.0 18.2 11.7 – 9.6 9.2 8.0 6.8 18.7 18.5 12.2 21.1 – 14.7 16.8 15.9 14.1 11.8 13.3 9.1 18.3 13.1 10.7 17.7 15.9 18.8 14.0 11.3 22.3 15.8 7.4 65.9 23.0 18.0 10.6 18.6 12.5 7.9 11.4 10.8 –
Min
Max
n
140.0 84.0 188.0 138.0 172.0 34.0 140.0 75.0 112.0 96.0 70.0 190.0 86.0 76.0 23.0 24.0 22.4 25.3 32.0 24.0 – 35.0 36.0 48.0 20.0 20.0 32.0 13.0 20.0 25.0 15.0 25.0 26.0 25.0 49.0 21.0 19.0 11.0 17.0 18.0 24.0 35.0 40.0 28.0 20.0 56.0 12.0 42.0 12.0
182.0 123.0 296.0 204.0 240.0 57.0 160.0 10.0 172.0 172.0 105.0 230.0 108.0 100 32.0 30.0 52.2 49.2 48.0 49.0 – 50.0 63.0 80.0 34.0 26.0 54.0 18.0 39.0 38.0 20.0 45.0 43.0 45.0 80.0 32.0 36.0 17.0 21.0 22.0 60.0 56.0 56.0 44.0 34.0 73.0 18.0 52.0 19.0
8 12 5 21 13 28 3 20 15 7 15 ? 5 7 13 10 25 25 12 13 10 10 15 15 23 4 18 10 31 13 11 14 8 22 19 21 11 10 12 11 13 8 12 5 21 13 28 3 20
228
SYSTEMATIC SECTION
Table 22 Continued Characteristics a Body, width
Body length:width, ratio
Adoral zone of membranelles, length
Species
mean
M
SD
SE
CV
Min
Max
n
pr1 pr2 qua te1 te2 vit b1 bi3 bi4 de1 de2 de3 el1 el2 el3 lo1 lo2 mu1 mu2 mu3 pr1 pr2 au1 au2 au3 au4 au5 bi2 bi3 bi4 de1 de2 de3 ed1 ed2 ed3 ed4 el1 el2 el3 gra ha1 ha2 hya isl kir lo1 lo2 mag
21.6 21.4 25.2 – 43.4 36.6 5.0 4.2 3.9 4.8 3.8 3.9 5.3 4.5 5.0 4.7 3.9 7.7 6.5 3.2 6.1 6.6 26.4 25.5 22.6 26.1 21.3 29.4 40.3 49.4 23.0 18.5 29.8 18.5 23.2 20.9 17.0 28.0 32.9 29.8 41.0 16.3 17.4 9.0 16.8 16.9 30.4 29.4 23.4
21.5 20.0 25.0 – 42.0 36.0 – 4.2 3.8 4.6 3.8 3.7 5.0 4.0 5.0 4.5 4.2 7.8 6.7 3.3 5.7 6.5 26.0 25.0 22.2 26.5 21.0 30.0 40.0 51.0 24.0 18.0 30.0 18.0 24.0 21.0 17.0 28.0 33.0 29.0 40.0 16.0 17.0 8.7 17.0 17.0 31.0 28.5 23.5
3.1 3.6 3.1 – 6.7 3.6 – 0.9 0.4 1.1 0.4 0.7 0.9 0.8 1.0 1.3 0.9 1.0 0.7 0.3 1.0 1.3 2.4 2.0 3.7 2.3 2.4 1.3 4.7 5.2 2.0 1.6 1.8 2.2 3.0 1.8 1.3 1.7 2.9 3.2 2.2 1.2 1.8 0.8 1.1 1.1 2.1 1.7 1.9
0.9 1.4 0.8 – 3.0 – – 0.2 0.1 0.2 0.2 0.2 0.2 0.3 0.2 0.4 0.3 0.4 0.2 0.1 0.3 0.5 0.6 0.6 0.7 0.7 0.7 0.4 1.2 1.3 0.4 0.5 0.4 0.7 0.6 0.4 0.4 0.5 1.0 0.7 0.5 0.3 0.5 0.3 – – 0.6 0.6 0.6
14.4 17.0 12.5 – 15.4 9.7 – 21.2 10.0 23.0 9.6 16.9 16.9 18.8 19.1 28.4 22.6 12.4 10.5 9.5 16.4 19.8 9.1 7.9 16.2 8.9 11.4 4.3 11.7 10.5 8.9 8.9 5.9 11.7 13.1 8.4 7.9 6.1 8.8 10.8 5.3 7.3 10.4 8.8 6.3 6.7 6.9 5.7 8.2
16.0 26.0 17.0 28.0 21.0 32.0 85.0 100.0 38.0 55.0 32.0 42.0 – – 2.9 6.0 2.9 4.6 3.0 7.4 3.4 4.3 3.1 5.8 3.4 7.6 3.6 6.0 3.4 7.6 3.2 8.1 2.5 5.2 6.7 8.9 5.4 7.8 2.8 3.8 5.0 7.7 4.8 8.5 22.0 30.0 23.0 29.0 13.0 29.6 22.0 29.0 16.0 25.0 28.0 31.0 33.0 48.0 40.0 56.0 19.0 26.0 15.0 21.0 27.0 34.0 15.0 22.0 16.0 29.0 18.0 25.0 16.0 20.0 25.0 31.0 29.0 37.0 25.0 37.0 38.0 46.0 15.0 19.0 14.0 20.0 8.0 10.0 15.0 18.0 15.0 18.0 27.0 33.0 28.0 32.0 21.0 28.0
12 7 15 ? 5 7 10 15 15 23 4 18 14 8 22 13 8 5 21 13 11 7 15 10 25 12 13 10 15 15 23 11 18 10 31 16 11 14 8 22 19 21 11 10 12 11 13 8 12
Uroleptoides
229
Table 22 Continued Characteristics a Adoral zone of membranelles, length
Species
mu1 mu2 mu3 pe1 pol por pr1 pr2 qua te2 vit Body length:length of adoral zone, ratio bi1 bi3 bi4 de1 de2 de3 el1 el2 el3 lo1 lo2 mu1 mu2 mu3 pr1 pr2 Paroral, length ed2 ed3 Endoral, length ed2 ed3 Distance 1 quag Anterior body end to amphisiellid median bi3 cirral row, distance bi4 lo1 Anterior body end to posterior end of au1 amphisiellid median cirral row, au2 distance au3 au4 au5 bi2 bi3 bi4 de1 de2 de3 ed1 ed2 ed3 ed4
mean
M
SD
SE
CV
Min
Max
n
38.0 39.0 33.1 32.0 38.8 39.0 12.8 13.0 52.7 56.0 27.0 28.0 20.1 20.0 22.3 22.0 18.3 19.0 27.4 27.0 22.6 22.0 5.0 – 5.3 5.4 4.8 4.5 5.4 5.2 4.8 4.8 5.3 5.3 5.5 6.0 4.7 5.0 5.2 5.0 5.9 6.1 5.5 5.4 6.8 7.1 5.2 5.3 5.4 5.4 6.5 6.4 6.3 6.1 10.3 10.0 9.0 9.0 9.2 9.0 7.8 8.0 11.7 11.0 8.3 7.0 10.7 10.0 7.3 7.5 38.4 38.0 38.1 37.5 36.7 38.1 59.2 58.0 53.3 53.0 83.7 82.0 175.5 180.0 218.3 219.0 25.5 25.0 22.7 24.0 20.2 20.0 19.2 18.5 29.2 30.0 28.0 27.5 19.5 19.0
3.9 2.3 3.6 0.8 5.8 1.7 1.5 3.3 2.5 0.6 1.5 – 0.6 0.5 0.8 0.3 0.6 0.6 0.5 0.7 0.9 0.7 0.8 0.4 0.7 0.9 1.3 1.8 0.9 1.1 0.9 2.3 3.3 2.7 1.9 4.1 3.4 3.0 9.7 8.5 8.2 28.5 29.2 2.6 2.8 2.6 2.6 6.0 3.4 1.6
1.8 0.5 1.0 0.2 3.3 0.5 0.4 1.2 0.6 0.2 – – 0.2 0.1 0.2 0.1 0.2 0.2 0.2 0.1 0.3 0.2 0.4 0.1 0.2 0.2 0.5 0.3 0.2 0.2 0.2 0.6 0.9 0.7 0.5 1.1 1.1 0.6 2.8 2.4 3.4 7.4 7.5 0.5 0.9 0.6 0.8 1.1 0.9 0.5
10.4 34.0 43.0 6.8 30.0 38.0 9.4 30.0 43.0 6.4 11.0 14.5 11.0 46.0 56.0 – 22.4 30.8 7.6 17.0 22.0 14.8 18.0 28.0 13.6 14.0 22.0 2.0 27.0 28.0 6.7 20.0 24.0 – – – 11.9 4.4 6.5 10.9 4.0 5.6 14.1 4.4 7.0 6.0 4.3 5.3 12.1 4.4 6.7 10.3 4.6 6.3 10.0 4.2 5.5 12.5 4.2 6.3 14.9 4.6 7.2 12.0 4.4 6.5 11.9 5.5 7.6 7.5 4.3 5.9 12.8 3.9 6.5 14.0 5.4 8.6 20.1 4.8 8.7 17.5 7.0 13.5 10.5 7.0 10.0 12.3 7.0 11.0 11.4 6.5 9.0 19.9 7.0 15.0 40.0 5.0 15.0 25.0 7.0 17.0 25.7 5.0 10.0 10.6 32.0 45.0 9.0 33.0 42.0 8.2 31.5 42.0 16.4 45.0 73.0 16.0 40.0 66.0 9.8 75.0 98.0 16.3 121.0 240.0 13.4 173.0 280.0 10.3 22.0 34.0 12.5 17.0 25.0 12.6 18.0 28.0 13.6 15.0 25.0 20.4 19.0 38.0 12.2 23.0 35.0 8.4 17.0 22.0
5 21 13 28 3 20 16 7 15 5 7 10 15 15 23 11 18 14 8 22 13 8 5 21 13 15 7 30 15 28 14 15 15 15 12 14 10 25 12 13 6 15 15 23 9 18 10 31 15 11
230
SYSTEMATIC SECTION
Table 22 Continued Characteristics a Anterior body end to posterior end of amphisiellid median cirral row, distance
Body length:length of amphisiellid median cirral row, ratio
Anterior body end to buccal cirrus, distance
Anterior body end to right marginal row, distance
Species
mean
M
SD
SE
CV
el1 el2 el3 gra ha1 ha2 hya isl kir lo1 lo2 mag mu1 mu2 mu3 pr1 pr2 qua te2 vit de1 de2 de3 el1 el2 el3 lo1 lo2 mu1 mu2 mu3 pr1 pr2 de1 de2 de3 el1 el2 el3 mu1 mu2 pr1 pr2 bi3 bi4 de1 de2 de3 el1 el2
47.6 49.5 48.3 38.5 12.0 12.7 8.0 16.8 18.6 123.9 109.5 49.3 183.6 90.0 153.5 39.4 44.4 32.5 66.3 37.7 4.9 3.9 7.7 3.3 3.1 3.2 1.4 1.5 1.4 1.9 1.4 3.3 3.2 12.4 10.3 15.8 10.7 14.6 12.1 17.2 15.8 7.4 11.0 7.9 8.9 6.9 7.8 7.8 8.0 11.4
47.0 49.0 47.0 38.0 12.0 13.0 8.1 17.0 18.0 123.0 107.5 50.0 165.0 87.0 145.0 40.0 41.0 32.0 66.0 39.0 4.8 4.0 7.8 3.0 3.0 3.0 1.4 1.5 1.4 1.9 1.4 3.3 3.1 12.0 10.0 16.0 11.0 15.0 11.0 17.0 16.0 8.0 10.0 9.0 9.0 7.0 8.0 8.0 7.0 11.0
9.5 8.1 8.9 3.4 1.2 2.1 1.1 1.3 2.2 13.0 8.9 9.4 36.9 15.6 16.0 5.0 10.2 3.9 – 8.4 0.7 0.5 1.1 0.5 0.5 0.5 0.1 0.2 0.3 0.2 0.1 0.2 0.5 1.3 2.7 1.4 0.9 2.5 2.5 3.2 1.1 1.1 1.8 3.6 3.4 1.8 0.8 2.6 3.0 4.3
2.5 2.9 1.9 0.8 0.3 0.6 0.3 – – 3.6 3.1 2.7 16.5 3.5 4.4 1.2 3.9 1.0 – – 0.1 0.2 0.3 0.1 0.2 0.1 0.0 0.1 0.1 0.0 0.1 0.1 0.2 0.3 1.0 0.3 0.2 0.9 0.5 1.4 0.3 0.3 0.7 0.9 0.9 0.4 0.3 0.6 0.8 1.5
20.0 16.3 18.3 8.7 10.1 16.1 13.8 7.5 11.8 10.5 8.1 19.0 20.1 17.3 10.4 12.6 23.0 12.1 – 22.2 13.9 12.2 14.9 14.1 16.9 14.9 6.3 10.1 21.7 9.6 6.9 6.8 16.0 10.2 26.2 8.6 8.5 16.7 20.6 18.6 6.7 14.7 16.6 44.8 38.3 25.7 9.6 33.8 38.0 37.9
Min
Max
n
34.0 39.0 34.0 33.0 10.0 9.0 6.2 14.0 17.0 106.0 100.0 28.0 152.0 64.0 135.0 30.0 33.0 25.0 63.0 25.0 3.9 3.3 6.1 2.3 2.3 2.3 1.4 1.3 1.2 1.5 1.2 3.0 2.4 10.0 6.0 14.0 10.0 11.0 10.0 13.0 14.0 6.0 9.0 0.0 0.0 4.0 7.0 2.0 4.0 5.0
73.0 62.0 73.0 48.0 15.0 16.0 10.0 18.0 24.0 152.0 124.0 63.0 237.0 124.0 180.0 49.0 60.0 39.0 70.0 46.0 6.3 4.7 10.1 4.0 3.7 4.0 1.6 1.8 2.0 2.4 1.5 3.7 3.8 14.0 14.0 19.0 13.0 19.0 19.0 22.0 18.0 9.0 14.0 14.0 14.0 11.0 9.0 12.0 14.0 18.0
14 8 22 19 21 11 10 12 11 13 8 12 5 20 13 16 7 15 3 6 23 9 18 14 8 22 13 8 5 20 13 15 7 23 7 18 14 8 22 5 12 16 7 15 15 23 6 18 14 8
Uroleptoides
231
Table 22 Continued Characteristics a
Species
mean
M
SD
SE
CV
Min
Max
n
el3 lo1 pr1 pr2 de1 de3 de1 de3 bi3 bi4 lo1 de1 de3 de1 de3 bi3 bi4 de1 de2 de3 ed1 lo1 bi3 bi4 de1 de2 de3 el1 el2 el3 lo1 au1 au2 au3 au4 au5 bi2 m bi3 bi4 de1 h de2 h de3 h ed1 h ed1 m ed2 h ed2 m ed3 h ed3 m ed4
9.2 6.9 6.5 4.9 6.1 5.9 3.7 2.0 9.6 9.1 7.5 7.6 7.9 5.7 4.6 70.1 67.3 33.5 25.1 44.5 17.4 49.8 65.7 81.1 52.1 37.1 53.4 72.5 76.4 73.9 65.0 17.8 15.2 18.2 20.2 16.0 25.2 26.5 25.9 14.7 13.0 18.2 9.9 9.3 16.4 17.4 18.1 18.5 11.8
9.0 6.5 6.0 5.0 7.0 6.0 3.0 1.0 11.0 9.0 7.0 8.0 8.0 6.0 4.0 70.0 70.0 34.0 25.0 42.0 17.0 48.0 63.0 78.0 53.0 39.0 53.5 70.0 80.0 73.0 64.0 16.0 14.5 18.5 21.0 15.0 25.0 26.0 26.0 15.0 13.0 18.5 10.0 9.5 16.0 16.0 17.8 18.8 12.0
3.8 1.9 1.3 1.8 2.5 1.9 2.0 1.8 2.9 1.9 1.5 0.9 2.2 1.3 0.9 15.8 9.8 4.7 3.4 8.4 2.5 8.9 11.7 12.9 11.6 6.5 6.5 9.3 8.4 8.9 5.6 3.7 2.0 3.9 1.9 3.0 3.8 2.5 3.8 3.4 2.1 2.4 1.3 1.6 3.6 3.8 2.2 3.9 1.2
0.8 0.5 0.3 0.7 0.5 0.5 0.5 0.4 0.7 0.5 0.4 0.2 0.5 0.3 0.2 4.1 2.5 1.0 1.0 2.0 0.8 2.5 3.0 3.3 2.4 1.8 1.5 2.5 3.0 1.9 1.7 1.0 0.6 0.8 0.5 0.8 1.2 0.6 1.0 0.7 0.6 0.6 0.4 0.5 0.7 0.7 0.6 1.0 0.4
41.5 27.2 20.3 36.5 40.6 32.4 54.8 87.5 29.9 21.0 20.2 11.7 27.8 22.2 18.8 22.5 14.5 14.1 13.4 18.9 14.4 17.9 17.7 15.9 22.3 17.6 12.1 12.8 10.9 12.1 8.7 20.9 13.4 21.2 9.2 18.7 15.2 9.2 14.6 22.9 16.3 13.1 12.8 17.6 22.2 21.7 12.2 21.0 9.9
4.0 18.0 4.0 10.0 5.0 9.0 2.0 7.0 1.0 11.0 2.0 10.0 1.0 8.0 0.0 7.0 5.0 15.0 5.0 12.0 6.0 10.0 6.0 9.0 5.0 14.0 3.0 8.0 3.0 6.0 39.0 97.0 52.0 84.0 23.0 42.0 20.0 31.0 33.0 65.0 14.0 22.0 38.0 66.0 52.0 90.0 63.0 110.0 33.0 70.0 22.0 49.0 38.0 63.0 60.0 86.0 62.0 85.0 60.0 86.0 54.0 75.0 14.0 25.0 13.0 19.0 11.1 25.8 17.0 22.0 11.0 20.0 20.0 31.0 22.0 30.0 20.0 33.0 7.0 20.0 10.0 17.0 11.0 21.0 8.0 12.0 6.0 11.0 11.0 25.0 12.0 26.0 13.0 22.5 11.5 26.0 9.0 13.0
22 12 16 7 21 18 20 18 15 15 13 23 18 21 18 15 15 23 11 18 10 13 15 15 23 13 18 14 8 22 11 13 10 25 12 13 10 15 15 23 13 18 10 10 31 31 14 14 11
Anterior body end to right marginal row, distance
Posterior body end to right marginal row, distance Posterior body end to left marginal row, distance Anterior body end to paroral, distance
Anterior end of paroral to buccal cirrus, distance Posterior body end to anteriormost transverse cirrus, distance Anterior body end to first macronuclear nodule, distance
Nuclear figure, length
Macronuclear nodule, length
232
SYSTEMATIC SECTION
Table 22 Continued Characteristics a Macronuclear nodule, length
Macronuclear nodule, width
Species el1 h el2 h el3 h gra ha1 m ha2 m hya h isl m kir m lo1 h lo2 h mag mu1 h mu2 h mu3 h pe1 h pe1 m pol c pr1 h pr2 h qua c te2 vit au1 au2 au3 au4 au5 bi2 m bi3 bi4 de1 h de2 h de3 h ed1 h ed1 m ed2 h ed2 m ed3 h ed3 m ed4 el1 h el2 h el3 h gra ha1m ha2m hya h isl m
mean
M
SD
SE
CV
Min
Max
n
12.3 13.3 12.6 24.5 15.3 17.4 5.4 10.2 13.2 16.5 15.4 19.3 11.4 7.5 9.3 8.0 8.3 19.3 24.3 24.9 8.1 13.5 12.6 6.6 7.8 8.6 10.0 7.6 8.0 10.3 10.4 7.4 6.5 8.1 4.7 4.8 7.4 7.5 9.2 9.4 4.8 7.5 6.8 7.2 9.1 7.8 6.9 3.5 5.3
13.0 14.0 13.0 25.0 15.0 18.0 5.6 10.0 14.0 16.0 16.0 20.0 12.0 8.0 9.0 8.0 8.0 21.0 24.0 26.0 8.0 12.5 13.0 6.0 7.7 7.4 10.0 8.0 8.0 10.0 10.0 7.0 6.0 7.5 4.0 5.0 7.5 7.0 9.0 9.0 5.0 8.0 7.0 7.0 9.0 7.0 7.0 3.5 5.0
1.5 2.7 2.0 2.2 1.8 3.0 0.7 1.8 1.2 1.7 2.3 2.8 2.4 2.5 1.9 1.4 1.5 3.8 4.1 2.5 1.0 1.7 1.6 0.9 0.8 1.9 0.7 0.7 0.9 0.7 1.9 1.2 1.0 2.5 0.5 0.6 1.3 1.8 0.9 1.8 0.8 – 1.2 0.9 0.9 1.1 1.4 0.5 0.5
0.4 0.9 0.4 0.5 0.4 0.9 0.2 – – 0.5 0.8 0.8 1.1 0.5 0.5 0.3 0.3 2.2 1.1 0.9 0.3 0.9 – 0.3 0.3 0.4 0.2 0.2 0.3 0.2 0.5 0.2 0.3 0.6 0.2 0.2 0.2 0.3 .2 0.5 0.2 – 0.4 0.2 0.2 0.2 0.4 0.2 –
12.5 20.1 15.9 9.1 11.5 17.3 12.5 17.2 8.9 10.1 15.1 14.3 21.1 32.9 20.5 18.2 17.9 19.6 16.9 10.0 12.2 12.8 12.9 13.9 10.8 22.1 7.4 8.5 11.8 6.9 18.5 16.2 15.0 31.3 10.3 13.4 16.8 24.6 9.7 18.7 15.6 – 17.3 12.0 10.1 14.1 19.9 15.7 8.6
10.0 9.0 9.0 21.0 12.0 13.0 4.2 8.0 11.0 14.0 11.0 15.0 8.0 4.0 5.6 6.0 6.0 15.0 14.0 20.0 7.0 11.0 11.0 5.3 6.0 6.7 9.0 7.0 7.0 9.0 8.0 6.0 5.0 7.0 4.0 4.0 5.0 4.5 8.0 7.0 4.0 7.0 5.0 5.0 7.0 6.0 5.0 2.8 5.0
15.0 17.0 17.0 28.0 19.0 22.0 6.2 13.0 14.0 19.0 18.0 24.0 14.0 13.0 13.0 11.0 11.5 22.0 29.0 27.0 10.0 15.0 15.0 7.6 9.0 13.0 11.0 9.0 10.0 12.0 16.0 10.0 8.0 18.0 5.5 6.0 10.0 12.0 11.0 13.0 6.0 8.0 9.0 9.0 10.0 10.0 9.0 4.8 6.0
14 8 22 19 21 11 10 12 11 13 8 12 5 21 13 28 28 3 15 7 15 4 7 13 10 25 12 13 10 15 15 23 13 18 10 10 31 31 14 14 11 14 8 22 19 21 11 10 12
Uroleptoides
233
Table 22 Continued Characteristics a
Species
Macronuclear nodule, width
Macronuclear nodules, length:width ratio
Macronuclear nodules, distance in between
Micronucleus, largest diameter
Micronucleus, length
kir m lo1 h lo2 mag mu1 h mu2 h mu3 h pe1 h pe1 m pol c pr1 h pr2 h qua c te2 vit de1 de2 de3 bi3 bi4 de1 de2 de3 ed1 el1 i el2 i el3 i ha1 ha2 hya isl kir pol quad quae au1 au2 au3 au4 au5 gra hya mag pe1 vit bi2 f bi3 q bi4 q de1 h
mean
M
SD
SE
CV
Min
Max
n
6.3 7.8 7.1 7.4 5.2 3.9 4.6 4.8 4.8 10.7 7.4 7.0 5.5 10.5 6.9 2.0 2.1 2.4 15.3 30.3 23.5 11.8 17.5 8.1 23.9 9.0 18.5 5.8 4.8 0.9 4.3 13.4 25.7 3.7 7.5 1.7 1.7 2.0 3.2 3.0 5.9 1.2 3.1 2.0 1.4 5.6 4.8 4.8 2.7
6.0 8.0 7.0 7.0 6.0 4.0 4.2 5.0 5.0 11.0 6.5 7.0 6.0 10.5 7.0 2.2 2.0 2.5 13.0 28.0 23.0 12.5 16.5 8.0 21.0 8.0 20.0 6.0 4.0 1.1 4.0 14.0 25.0 3.0 7.0 1.7 1.7 2.0 3.0 3.0 6.0 1.2 3.0 2.0 1.4 6.0 5.0 5.0 2.5
0.7 0.7 – 1.0 1.1 0.6 – 0.5 0.5 0.6 1.7 1.2 0.6 0.6 0.7 0.5 0.5 0.5 8.9 9.7 10.4 3.5 5.4 2.0 6.1 2.9 8.9 1.7 4.2 0.7 1.4 3.2 2.1 1.9 2.5 0.2 0.2 0.2 0.3 0.2 0.9 0.2 0.3 0.3 0.1 1.1 0.7 0.6 –
– 0.2 – 0.3 0.5 0.1 – 0.1 0.1 0.3 0.4 0.4 0.2 0.3 – 0.1 0.1 0.1 2.3 2.5 2.2 1.0 1.3 0.6 1.6 1.0 1.9 0.4 1.3 0.2 – – 1.2 0.5 0.6 0.1 0.1 0.0 0.1 0.1 0.2 0.1 0.1 0.1 – 0.3 0.2 0.2 –
10.3 8.8 – 13.4 21.1 16.0 – 10.1 10.9 5.4 23.0 16.5 11.7 5.5 10.0 24.7 21.5 22.2 58.1 32.1 44.2 30.2 30.9 24.9 25.5 32.5 48.3 30.0 86.5 76.0 32.9 24.2 8.1 52.2 33.5 12.8 12.0 10.0 9.7 7.8 14.8 16.7 8.6 16.9 5.8 19.2 13.8 12.8 –
5.0 7.0 7.0 6.0 4.0 3.0 4.0 4.0 4.0 10.0 6.0 6.0 4.0 10.0 6.0 1.0 1.3 1.0 0.0 17.0 9.0 6.0 8.0 5.0 15.0 6.0 6.0 3.0 0.0 0.0 2.0 6.0 24.0 0.0 4.0 1.5 1.5 1.6 3.0 3.0 5.0 1.0 3.0 1.5 1.4 4.0 4.0 4.0 2.5
7.0 9.0 8.0 10.0 6.0 5.0 5.6 6.0 6.0 11.0 10.0 9.0 6.0 11.0 8.0 2.7 2.8 3.0 30.0 52.0 53.0 17.0 28.0 12.0 35.0 14.0 35.0 10.0 11.0 2.0 7.0 17.0 28.0 7.0 14.0 2.1 2.0 2.2 4.0 4.0 8.0 1.4 4.0 3.0 1.6 7.0 6.0 6.0 3.0
11 13 8 12 5 21 13 28 28 3 16 7 15 4 7 23 13 18 15 15 23 12 18 10 14 8 22 21 11 10 12 11 3 15 15 12 10 25 12 12 19 3 12 28 6 10 13 15 21
234
SYSTEMATIC SECTION
Table 22 Continued Characteristics a Micronucleus, length
Micronucleus, width
Macronuclear nodules, number
Species de2 h de3 h ed1 f ed2 ed3 ed4 el1 h el2 h el3 h ha1f ha2f isl f kir f lo1 h lo2 h mu1 h mu2 h mu3 h pol f pr1 h pr2 h qua f bi2 f bi3 bi4 de1 h de2 h de3 h ed1 f ed2 ed3 ed4 el1 h el2 h el3 h ha1 f ha2 f isl f kir f lo1 h lo2 h mu1 h mu2 h mu3 h pol f pr1 h pr2 h qua f au1
mean
M
SD
SE
CV
Min
Max
n
2.5 6.2 2.0 2.9 2.6 2.1 2.4 2.5 2.4 2.1 2.2 1.6 3.2 4.2 4.1 3.8 4.0 5.0 4.0 2.6 2.7 1.8 2.8 3.0 3.0 2.5 2.4 2.2 1.8 2.9 2.4 1.6 2.0 1.8 1.9 1.8 1.8 1.6 1.6 2.7 2.9 2.7 1.8 2.7 3.4 2.5 2.7 1.7 2.0
2.5 6.5 2.0 3.0 2.5 2.0 2.5 2.5 3.0 2.0 2.0 1.6 3.0 4.0 4.0 3.0 4.0 5.6 4.0 2.5 2.0 1.6 3.0 3.0 3.0 2.5 2.5 2.0 2.0 3.0 2.5 1.5 2.0 2.0 2.0 2.0 2.0 1.6 1.5 2.5 3.0 2.5 2.0 2.8 3.5 2.5 2.0 1.5 2.0
– 0.8 0.2 0.4 0.4 0.3 – – – – – 0.2 0.4 0.9 0.9 1.1 0.7 0.8 0.5 – – 0.6 0.5 – 0.2 – – – 0.4 0.4 0.4 0.2 – – – – – 0.1 0.2 – – – – – 0.6 – – 0.6 0.0
– 0.2 0.1 0.1 0.1 0.1 – – – – – – – 0.3 0.3 0.5 0.2 0.2 0.3 – – 0.2 0.2 – 0.1 – – – 0.1 0.1 0.1 0.1 – – – – – – – – – – – – 0.3 – – 0.2 0.0
– 12.4 8.1 12.9 14.5 15.1 – – – – – 10.1 12.7 22.5 21.7 28.8 17.8 16.4 12.5 – – 34.4 18.6 – 7.6 – – – 20.2 12.9 16.3 12.9 – – – – – 7.0 9.8 – – – – – 17.6 – – 33.9 0.0
2.4 5.0 1.5 2.0 2.0 2.0 2.0 2.0 2.0 2.0 2.0 1.4 3.0 3.0 3.0 3.0 3.0 3.0 3.5 2.4 2.0 1.4 2.0 3.0 2.5 2.0 2.0 2.0 1.0 2.0 2.0 1.5 1.5 1.5 1.5 1.5 1.5 1.4 1.5 2.5 2.5 2.5 1.5 2.2 2.8 1.5 2.0 1.4 2.0
3.0 7.0 2.0 4.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 2.0 4.0 6.0 5.6 5.0 5.0 5.0 4.5 3.0 4.0 3.5 4.0 4.0 3.5 3.0 2.5 3.0 2.0 4.0 3.0 2.0 2.0 2.0 2.0 2.0 2.0 1.8 2.0 3.0 3.0 3.0 2.0 3.0 4.0 3.0 4.0 3.0 2.0
9 18 10 31 16 11 14 8 22 21 11 12 11 12 8 5 15 13 3 6 3 14 10 13 15 21 9 18 10 31 16 11 14 8 22 21 3 12 11 12 8 5 15 13 3 6 3 14 14
Uroleptoides
235
Table 22 Continued Characteristics a Macronuclear nodules, number
Micronuclei, number
Species
mean
M
SD
SE
CV
au2 au3 au4 au5 bi1 bi2 bi3 bi4 de1 de2 de3 ed1 ed2 ed3 ed4 el1 el2 el3 gra ha1 ha2 hya lon lo1 lo2 mag mu1 mu2 pe1 pe2 pol pr1 pr2 qua rap te1 te2 vit au1 au2 au3 au4 au5 bi1 bi2 bi3 bi4 de1 de2
2.1 2.1 2.0 2.0 2.0 2.0 2.3 2.0 2.3 2.0 2.0 2.0 2.0 2.0 2.0 4.0 6.2 4.9 2.0 2.0 2.0 2.0 2.0 4.0 4.0 2.0 35.6 26.0 2.0 2.0 2.0 2.2 2.0 4.0 8.0 2.0 2.0 4.1 2.1 2.3 2.2 3.2 2.8 – 2.0 1.9 4.8 1.5 2.0
2.0 2.0 2.0 2.0 – 2.0 2.0 2.0 2.0 2.0 2.0 2.0 2.0 2.0 2.0 4.0 6.0 4.0 2.0 2.0 2.0 2.0 – 4.0 4.0 2.0 35.0 26.0 2.0 – 2.0 2.0 2.0 4.0 – – 2.0 4.0 2.0 2.0 2.0 3.0 3.0 2.0 2.0 2.0 4.0 2.0 2.0
0.3 0.3 0.0 0.0 – 0.0 – 0.0 0.6 0.0 0.0 0.0 0.2 0.0 0.0 0.0 0.8 1.2 0.0 0.0 0.0 0.0 – 0.0 0.0 0.0 1.8 4.0 0.0 – 0.0 – 0.0 0.0 – – 0.0 – 0.5 0.5 0.5 1.0 0.9 – 0.0 1.9 3.6 0.7 0.0
0.1 15.1 0.1 14.3 0.0 0.0 0.0 0. – – 0.0 0.0 – – 0.0 0.0 0.1 27.6 0.0 0.0 0.0 0.0 0.0 0.0 0.0 8.8 0.0 0.0 0.0 0.0 0.0 0.0 0.3 13.4 0.3 25.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 – – 0.0 0.0 0.0 0.0 0.0 0.0 0.8 5.1 1.0 15.3 0.0 0.0 – – 0.0 0.0 – – 0.0 0.0 0.0 0.0 – – – – 0.0 0.0 – – 0.1 24.7 0.2 21.0 0.1 22.7 0.3 52.5 0.3 33.5 – – 0.0 0.0 0.5 103.0 0.9 74.4 0.2 49.4 0.0 0.0
Min
Max
n
2.0 2.0 2.0 2.0 – 2.0 –2.0 2.0 2.0 2.0 2.0 2.0 2.0 2.0 2.0 4.0 5.0 4.0 2.0 2.0 2.0 2.0 – 4.0 4.0 2.0 34.0 17.0 2.0 – 2.0 2.0 2.0 4.0 – – 2.0 4.0 2.0 2.0 2.0 2.0 1.0 2.0 2.0 0.0 1.0 0.0 2.0
3.0 3.0 2.0 2.0 – 2.0 4.0 2.0 4.0 2.0 2.0 2.0 3.0 2.0 2.0 4.0 8.0 8.0 2.0 2.0 2.0 2.0 – 4.0 4.0 2.0 38.0 32.0 2.0 – 2.0 4.0 2.0 4.0 – – 2.0 5.0 3.0 3.0 4.0 5.0 4.0 3.0 2.0 5.0 15.0 3.0 2.0
10 25 13 12 10 9 15 15 23 13 18 10 31 18 11 14 8 22 19 21 11 10 30 13 7 12 5 17 28 10 3 16 7 15 ? ? 4 7 12 10 25 12 13 10 9 15 15 23 9
236
SYSTEMATIC SECTION
Table 22 Continued Characteristics a Micronuclei, number
Adoral membranelles, number
Species
mean
M
SD
SE
CV
Min
Max
n
de3 ed1 ed2 ed3 ed4 el1 el2 el3 gra ha1 ha2 isl kir lon lo1 lo2 mag mu1 mu2 mu3 pol pr1 pr2 qua rap te1 vit au1 au2 au3 au4 au5 bi1 bi2 bi3 bi4 de1 de2 de3 ed1 ed2 ed3 ed4 el1 el2 el3 gra ha1 ha2
2.1 2.2 2.0 1.9 2.3 2.7 7.5 4.5 2.1 2.0 2.0 1.8 2.0 2.0 1.9 1.9 3.3 6.6 2.9 8.9 3.7 1.7 1.3 2.3 4.0 2.0 3.5 22.4 21.2 21.6 24.0 21.8 25.0 24.5 32.1 39.9 20.4 19.6 24.6 16.7 16.8 17.4 16.1 21.7 25.4 23.1 24.6 17.0 18.3
2.0 2.0 2.0 2.0 2.0 3.0 7.0 3.0 2.0 2.0 2.0 2.0 2.0 – 2.0 2.0 3.0 7.0 3.0 8.0 3.0 2.0 1.0 2.0 – – 3.5 22.0 21.5 21.0 24.0 21.0 – 24.0 32.0 41.0 20.0 19.0 25.0 17.0 17.0 17.0 16.0 23.0 26.0 23.0 24.0 17.0 18.0
0.8 0.4 0.5 0.5 0.6 1.0 5.0 3.8 – 0.0 – 0.4 0.0 – – – 1.1 – 1.5 3.9 1.2 0.8 – 0.5 – – – 1.2 1.2 1.9 1.5 1.5 – 1.8 1.9 3.5 0.9 1.0 1.3 0.7 1.0 0.7 0.5 2.0 2.5 2.8 1.1 0.8 0.8
0.2 0.1 0.1 0.1 0.2 0.3 1.8 0.8 – 0.0 – – – – – – 0.3 – 0.4 1.1 0.7 0.3 – 0.1 – – – 0.3 0.4 0.4 0.4 0.4 – 0.6 0.5 0.9 0.2 0.3 0.3 0.2 0.2 0.1 0.2 0.5 0.9 0.6 0.2 0.2 0.2
35.9 19.2 25.8 25.8 28.5 36.6 66.1 85.1 – 0.0 – 21.2 0.0 – – – 32.2 – 50.9 44.0 31.5 44.1 – 20.5 – – – 5.4 5.8 8.8 6.4 6.7 – 7.2 6.1 8.8 4.4 4.9 5.4 4.1 6.0 4.1 3.4 9.3 9.6 12.1 4.3 4.8 4.3
1.0 1.0 1.0 1.0 1.0 0.0 2.0 0.0 1.0 2.0 2.0 1.0 2.0 – 1.0 1.0 2.0 6.0 1.0 5.0 3.0 1.0 1.0 2.0 – – 3.0 21.0 19.0 18.0 22.0 20.0 24.0 22.0 28.0 34.0 18.0 18.0 22.0 16.0 14.0 16.0 15.0 18.0 22.0 18.0 23.0 15.0 17.0
4.0 3.0 4.0 3.0 3.0 4.0 18.0 18.0 3.0 2.0 2.0 2.0 2.0 – 2.0 2.0 6.0 7.0 6.0 20.0 5.0 3.0 2.0 3.0 – – 4.0 26.0 23.0 25.0 27.0 24.0 28.0 28.0 36.0 45.0 21.0 21.0 27.0 18.0 19.0 19.0 17.0 24.0 29.0 29.0 27.0 18.0 19.0
18 10 31 17 11 14 8 22 19 21 3 12 11 30 12 8 12 5 14 13 3 7 3 14 ? ? 4 16 10 25 12 13 10 8 15 15 13 13 18 10 31 31 11 14 8 22 19 21 11
Uroleptoides
237
Table 22 Continued Characteristics a Adoral membranelles, number
Frontal cirri, number
Species
mean
M
SD
SE
CV
Min
Max
n
hya isl kir lon lo1 lo2 mag mu1 mu2 mu3 pe1 pe2 pol por pr1 pr2 qua rap te1 te2 vit au1 j au2 j au3 au4 au5 bi1 bi2 bi3 bi4 de1 de2 de3 ed1 ed2 ed3 ed4 el1 el2 el3 gra ha1 ha2 hya lo1 lo2 mag mu1 mu2 mu3
10.7 15.6 15.3 – 24.0 26.3 22.7 30.0 28.4 29.3 16.1 – 36.7 17.0 20.2 20.3 16.6 33.0 – 29.4 21.4 3.8 4.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 2.6 2.8 2.6 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0
11.0 16.0 16.0 – 23.0 26.0 22.5 30.0 28.0 29.0 16.0 – 37.0 17.0 20.0 20.0 17.0 – – 30.0 21.0 4.0 4.0 3.0 3.0 3.0 – 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0
0.5 0.5 1.0 – 2.0 0.9 1.2 2.2 2.4 1.2 0.8 – 1.5 0.7 1.7 1.6 1.6 – – 1.5 1.0 0.4 0.0 0.0 0.0 0.0 – 0.0 0.0 0.0 – .0 0.0 0.0 0.2 0.0 0.0 0.8 – 0.7 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0
0.2 – – – 0.5 0.3 0.4 1.0 0.6 0.3 0.1 – 0.9 0.2 0.4 0.6 0.4 – – 0.7 – 0.1 0.0 0.0 0.0 0.0 – 0.0 0.0 0.0 – 0.0 0.0 0.0 0.0 0.0 0.0 0.2 – 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0
4.5 3.3 6.6 – 8.2 3.4 5.4 7.5 8.4 4.0 4.9 – 4.2 – 8.6 7.9 9.6 – – 5.2 4.6 10.9 0.0 0.0 0.0 0.0 – 0.0 0.0 0.0 – 0.0 0.0 0.0 6.1 0.0 0.0 29.4 – 25.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0
10.0 15.0 13.0 18.0 22.0 25.0 21.0 27.0 25.0 28.0 15.0 15.0 35.0 16.0 17.0 18.0 14.0 33.0 26.0 27.0 20.0 3.0 4.0 3.0 3.0 3.0 – 3.0 3.0 3.0 2.0 3.0 3.0 3.0 2.0 3.0 3.0 1.0 2.0 1.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0
11.0 16.0 16.0 19.0 27.0 28.0 25 33.0 33.0 32.0 18.0 16.0 38.0 18.0 23.0 23.0 19.0 36.0 28.0 31.0 23.0 4.0 4.0 3.0 3.0 3.0 – 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0
10 12 11 30? 13 8 12 5 15 13 28 10 3 20 15 7 15 20 ? 5 7 15 10 25 12 13 10 5 15 15 23 10 18 10 31 18 11 14 8 22 19 21 11 9 11 8 12 5 6 13
238
SYSTEMATIC SECTION
Table 22 Continued Characteristics a Frontal cirri, number
Buccal cirri, number
Cirri left of anterior end of amphisiellid median cirral row, number
Species
mean
M
SD
SE
CV
Min
Max
n
pe1 pol pr1 pr2 qua te2 vit au1 au2 au3 au4 au5 bi1 bi2 bi3 bi4 de1 de2 de3 ed1 ed2 ed3 ed4 el1 el2 el3 gra ha1 ha2 lo1 lo2 hya mag mu1 mu2 mu3 pe1 pol pr1 pr2 qua te2 vit au1 au2 au3 b au4 b au5 b bi1 b bi2 b
3.0 3.0 3.0 3.1 3.0 3.0 3.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.1 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 0.8 0.9 1.0 1.0 1.4 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 5.3 1.0 1.0 2.7 1.0 1.1 5.1 3.9 4.3 4.6 3.8 3.0 2.8
3.0 3.0 3.0 3.0 3.0 3.0 3.0 1.0 1.0 1.0 1.0 1.0 – 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 6.0 1.0 1.0 3.0 1.0 1.0 5.0 3.0 4.0 5.0 3.0 – 3.0
0.0 0.0 0.0 – 0.0 – 0.0 0.0 0.0 0.0 0.0 0.0 – 0.0 – 0.0 0.0 0.0 0.0 0.0 0.2 0.0 0.0 0.0 – – 0.0 0.0 – 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1.2 0.0 0.0 0.5 0.0 – 2.0 2.1 0.7 1.4 1.2 – 0.8
0.0 0.0 0.0 – 0.0 – – 0.0 0.0 0.0 0.0 0.0 – 0.0 – 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 – – 0.0 0.0 – 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.7 0.0 0.0 0.1 0.0 – 0.4 0.6 0.1 0.4 0.3 – 0.4
0.0 0.0 0.0 – 0.0 – 0.0 0.0 0.0 0.0 0.0 0.0 – 0.0 – 0.0 0.0 0.0 0.0 0.0 17.4 0.0 0.0 0.0 – – 0.0 0.0 – 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 21.7 0.0 0.0 16.7 0.0 – 38.7 54.2 16.3 30.1 32.8 – 29.9
3.0 3.0 3.0 3.0 3.0 3.0 3.0 1.0 1.0 1.0 1.0 1.0 – 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 0.0 0.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 4.0 1.0 1.0 2.0 1.0 1.0 3.0 2.0 4.0 2.0 2.0 – 2.0
3.0 3.0 3.0 4.0 3.0 3.0 3.0 1.0 1.0 1.0 1.0 1.0 – 1.0 2.0 1.0 1.0 1.0 1.0 1.0 2.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 3.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 6.0 1.0 1.0 3.0 1.0 2.0 9.0 10.0 7.0 6.0 6.0 – 4.0
28 3 14 7 15 3 6 15 10 25 12 13 10 7 15 15 23 10 18 10 31 20 11 14 8 22 19 21 11 10 8 10 12 5 13 13 28 3 16 7 15 4 7 19 11 25 12 13 10 5
Uroleptoides
239
Table 22 Continued Characteristics a Cirri left of anterior end of amphisiellid median cirral row, number
Amphisiellid median cirral row, number of cirri
Species bi3 b bi4 b de1 b de2 b de3 b ed1 el1 b el2 b el3 b gra b ha1 ha2 isl kir lo1 b lo2 b mag b mu3 b pe1 b pol b pr1 pr2 qua te2 vit b au1 au2 au3 au4 au5 bi1 bi2 bi3 bi4 de1 de2 de3 ed1 ed2 ed3 ed4 el1 el2 el3 gra ha1 ha2 hya isl
mean
M
SD
SE
CV
Min
Max
n
3.4 3.3 3.1 3.0 3.3 1.0 3.5 2.6 3,2 2.9 1.2 1.7 3.1 1.0 3.0 3.0 4.4 3.9 3.0 3.0 3.2 3.3 3.2 5.7 3.3 12.5 13.3 13.5 19.0 17.3 – 22.3 66.7 80.7 4.1 7.2 4.1 8.0 9.5 9.9 7.8 12.8 11.1 12.2 3.9 3.6 4.2 4.0 5.7
3.0 3.0 3.0 3.0 3.0 1.0 3.0 2.0 3.0 3.0 1.0 2.0 3.0 1.0 3.0 3.0 4.5 4.0 3.0 3.0 3.0 4.0 3.0 5.0 3.0 13.0 13.5 14.0 19.0 17.0 – 23.0 65.0 77.0 4.0 7.0 4.0 8.0 9.0 10.0 7.0 13.0 12.0 13.0 4.0 4.0 4.0 4.0 6.0
0.9 1.0 – 0.0 0.6 0.0 1.1 1.7 1.4 – – 0.8 0.3 0.0 0.0 0.0 0.7 1.0 0.3 0.0 – 1.2 0.4 – – 2.1 1.4 1.0 3.6 2.2 – 1.5 8.5 10.3 – 0.6 – 0.7 1.3 1.2 1.1 2.7 5.6 3.9 – 0.6 0.9 0.0 0.5
0.2 0.3 – 0.0 0.1 0.0 0.3 0.6 0.3 – – 0.2 – – 0.0 0.0 0.2 0.3 0.1 0.0 – 0.5 0.1 – – 0.5 0.4 0.2 1.1 0.6 – 0.6 2.2 2.7 – 0.2 – 0.2 0.2 0.3 0.3 0.7 2.0 0.8 – 0.1 0.3 0.0 –
26.8 31.6 – 0.0 17.5 0.0 31.2 64.2 43.0 – – 45.5 9.4 0.0 0.0 0.0 15.1 24.5 9.1 0.0 – 36.3 12.9 – – 16.8 10.7 7.4 19.2 12.8 – 6.7 12.7 12.8 – 8.8 – 8.3 13.3 12.2 13.8 20.9 50.3 32.3 – 16.7 20.1 0.0 8.7
3.0 3.0 3.0 3.0 3.0 1.0 2.0 0.0 0.0 2.0 1.0 1.0 3.0 1.0 3.0 3.0 3.0 3.0 2.0 3.0 3.0 1.0 3.0 5.0 3.0 8.0 11.0 12.0 12.0 15.0 33.0 20.0 50.0 63.0 4.0 6.0 4.0 7.0 7.0 8.0 7.0 7.0 4.0 4.0 3.0 3.0 3.0 4.0 5.0
6.0 7.0 5.0 3.0 5.0 1.0 6.0 5.0 6.0 3.0 2.0 3.0 4.0 1.0 3.0 3.0 5.0 6.0 4.0 3.0 4.0 4.0 4.0 7.0 5.0 17.0 15.0 16.0 23.0 21.0 40.0 24.0 84.0 99.0 5.0 8.0 5.0 9.0 13.0 12.0 10.0 16.0 20.0 20.0 4.0 5.0 5.0 4.0 6.0
15 15 19 10 18 10 14 8 22 19 21 11 12 11 10 8 12 13 28 3 13 6 15 3 6 19 10 25 12 13 10 6 15 15 19 10 18 10 35 14 11 14 8 22 19 21 11 10 12
240
SYSTEMATIC SECTION
Table 22 Continued Characteristics a Amphisiellid median cirral row, number of cirri
Pretransverse cirri, number Transverse cirri, number
Species
mean
M
SD
SE
CV
Min
Max
n
kir lo1 lo2 mag mu1 mu2 mu3 pe1 pol pr1 pr2 qua te1 te2 vit gra au1 au2 au3 au4 au5 bi1 bi2 bi3 bi4 de1 l de2 l de3 l ed1 ed2 ed3 ed4 el1 el2 el3 gra ha1 ha2 hya l isl l kir l lon n lo1 lo2 mag mu3 pe1 pe2 pol
7.2 39.0 47.6 16.1 61.2 34.2 52.5 6.5 32.5 14.9 15.9 14.6 – 31.8 13.2 1.9 2.4 2.0 2.0 5.2 5.5 3.0 4.0 2.6 3.5 7.1 5.1 6.0 4.1 4.3 4.4 4.0 3.9 3.4 3.7 5.0 3.7 2.3 5.9 3.1 3.0 5.0 3.8 4.2 4.9 3.4 3.6 3.0 4.0
7.0 40.0 46.5 16.0 63.0 35.0 53.0 6.0 32.5 15.0 15.0 15.0 – 31.0 13.5 2.0 2.0 2.0 2.0 5.0 6.0 – 4.0 2.0 3.0 7.0 5.0 6.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 5.0 4.0 3.0 5.0 3.0 3.0 – 4.0 4.0 5.0 3.0 3.5 – 4.0
1.0 6.4 3.9 2.0 11.9 5.5 4.4 0.7 6.4 1.5 4.4 1.6 – 2.4 3.1 – 0.7 0.0 0.0 0.6 0.5 – 0.0 0.8 – 0.7 – 0.6 0.6 0.7 0.8 0.6 1.2 1.4 1.3 0.0 0.6 1.9 0.4 0.3 0.0 – 0.8 – 0.8 0.9 0.9 – 0.0
– 1.8 1.4 0.6 5.3 1.5 1.2 0.1 4.5 0.4 1.7 0.4 – 1.2 – – 0.2 0.0 0.0 0.2 0.1 – 0.0 0.2 – 0.2 – 0.1 0.2 0.1 0.2 0.2 0.3 0.5 0.3 0.0 0.1 0.6 0.1 – – – 0.2 – 0.2 0.2 0.2 – 0.0
13.7 16.3 8.2 12.3 19.5 16.2 8.4 10.7 19.6 10.1 27.8 11.2
6.0 24.0 43.0 12.0 43.0 22.0 46.0 6.0 28.0 12.0 11.0 12.0 26.0 30.0 8.0 1.0 2.0 2.0 2.0 4.0 5.0 – 4.0 2.0 3.0 6.0 5.0 5.0 3.0 3.0 3.0 3.0 0.0 1.0 0.0 5.0 2.0 0.0 5.0 3.0 3.0 – 2.0 4.0 3.0 2.0 2.0 2.0 4.0
9.0 50.0 54.0 19 73.0 43.0 61.0 8.0 37.0 17.0 23.0 18.0 30.0 35.0 16.0 2.0 4.0 2.0 2.0 6.0 6.0 – 4.0 4.0 4.0 9.0 6.0 7.0 5.0 6.0 6.0 5.0 5.0 5.0 5.0 5.0 4.0 4.0 6.0 4.0 3.0 – 5.0 5.0 6.0 5.0 5.0 4.0 4.0
11 13 8 12 5 13 13 28 2 15 7 15 ? 4 6 19 15 10 25 12 13 10 7 15 15 17 9 18 10 31 17 11 14 8 22 19 21 11 7 12 11 30 11 8 12 13 28 10 2
7.4 23.7 – 30.7 0.0 0.0 11.2 9.4 – 0.0 31.9 – 9.8 – 9.9 13.8 15.1 18.1 15.8 31.9 41.7 35.0 0.0 15.8 83.7 6.5 9.4 0.0 – 19.7 – 16.1 25.7 24.3 – 0.0
Uroleptoides
241
Table 22 Continued Characteristics a Transverse cirri, number
Left marginal cirri, number
Species
mean
M
SD
SE
CV
Min
Max
n
pr1 pr2 qua rap te1 te2 vit au1 au2 au3 au4 au5 bi1 bi2 bi3 bi4 de1 de2 de3 ed1 ed2 ed3 ed4 el1 el2 el3 gra ha1 ha2 hya isl kir lon lo1 lo2 mag mu1 mu2 mu3 pe1 pe2 pol por pr1 pr2 qua rap te1 te2 vit
4.2 3.5 3.0 5.0 – 6.3 1.9 31.9 28.7 27.4 47.2 42.5 55.0 46.0 73.7 77.9 31.5 34.2 57.7 17.6 17.7 25.4 15.3 48.6 45.9 47.6 45.8 15.9 22.6 11.6 14.1 41.1 – 55.8 61.5 44.8 88.5 68.6 80.3 14.7 – 41.0 22.0 54.6 54.9 36.6 – – 38.4 28.9
4.0 4.0 3.0 – – 6.0 2.0 33.0 29.0 28.0 47.0 41.0 – 45.0 75.0 76.0 32.0 34.0 58.0 18.0 18.0 24.5 15.0 50.0 49.0 49.0 45.0 16.0 22.0 11.5 14.0 41.0 – 56.0 62.5 42.5 90.5 67.5 79.0 15.0 – 44.0 22.0 53.5 51.0 35.0 – – 37.0 28.0
0.8 1.9 0.4 – – – – 3.1 2.3 2.9 4.9 5.2 – 5.1 8.8 14.2 2.7 2.6 7.2 1.8 2.9 3.6 1.9 5.1 7.8 6.1 2.5 2.2 6.0 1.1 0.8 3.8 – 3.8 5.7 7.1 5.9 8.3 10.1 2.1 – 9.9 3.4 6.3 11.8 6.1 – – 3.1 3.4
0.2 0.8 0.1 – – – – 0.8 0.7 0.6 1.4 1.4 – 1.8 2.3 3.7 0.6 0.7 1.7 0.6 0.5 1.1 0.6 1.4 2.7 1.3 0.6 0.5 1.8 0.3 – – – 1.1 2.0 2.1 3.0 2.2 2.8 0.4 – 5.7 1.0 1.8 4.5 1.6 – – 1.4 –
18.8 53.5 12.6 – – – – 9.8 8.1 10.6 10.3 12.1 – 11.0 12.0 18.2 8.7 7.6 12.5 10.1 16.4 14.3 12.5 10.4 16.9 12.9 5.4 13.6 26.8 9.3 5.6 9.3 – 6.9 9.2 15.9 6.7 12.1 12.6 14.2 – 24.0 – 11.6 21.6 16.7 – – 8.2 11.7
3.0 6.0 0.0 5.0 2.0 4.0 – – 6.0 8.0 6.0 7.0 1.0 2.0 26.0 37.0 24.0 31.0 22.0 35.0 37.0 53.0 35.0 52.0 – – 42.0 58.0 60.0 89.0 50.0 108.0 27.0 36.0 29.0 37.0 46.0 71.0 15.0 20.0 13.0 26.0 20.0 31.0 13.0 20.0 38.0 57.0 31.0 55.0 31.0 57.0 42.0 52.0 12.0 19.0 16.0 34.0 10.0 14.0 13.0 15.0 33.0 46.0 20.0 22.0 49.0 62.0 52.0 68.0 38.0 60.0 80.0 93.0 56.0 85.0 67.0 102.0 10.0 18.0 13.0 15.0 30.0 49.0 16.0 27.0 47.0 72.0 43.0 80.0 29.0 49.0 60.0 62.0 35.0 40.0 35.0 43.0 26.0 35.0
10 6 15 20 ? 3 7 14 10 25 13 13 10 8 15 15 21 12 18 10 31 12 11 14 8 22 19 21 11 10 12 11 30? 12 8 12 4 14 13 28 10 3 20 12 7 15 20 ? 5 7
242
SYSTEMATIC SECTION
Table 22 Continued Characteristics a Right marginal cirri, number
Dorsal kineties, number
Species
mean
M
SD
SE
CV
Min
Max
n
au1 au2 au3 au4 au5 bi1 bi2 bi3 bi4 de1 de2 de3 ed1 ed2 ed3 ed4 el1 el2 el3 gra ha1 ha2 hya isl kir lon lo1 lo2 mag mu1 mu2 mu3 pe1 pe2 pol por pr1 pr2 qua rap te1 te2 vit au1 au2 au4 au5 bi1 bi2 bi3
38.7 33.7 31.0 52.3 41.2 60.0 42.3 80.5 91.3 36.4 34.0 60.3 18.1 18.2 24.4 15.5 47.7 41.5 45.5 44.4 21.8 27.1 10.7 14.7 42.7 – 55.8 75.1 41.0 95.3 72.0 90.5 14.4 – 43.0 30.0 56.8 58.9 36.1 61.0 – 49.8 32.9 3.0 3.0 3.0 3.2 3.0 3.0 3.0
38.5 33.0 31.0 53.5 42.0 – 42.5 82.0 88.0 36.0 34.0 61.5 18.0 18.0 24.5 16.0 46.0 42.0 46.0 44.0 21.0 26.0 11.0 15.0 42.0 – 55.0 75.5 39.5 96.5 73.5 90.0 14.5 – 44.0 30.0 55.0 53.0 36.0 – – 48.0 33.0 3.0 3.0 3.0 3.0 – – 3.0
2.7 3.4 2.2 4.6 4.8 – 3.3 9.8 12.0 2.5 2.6 7.1 1.7 1.7 1.7 1.6 5.2 11.8 8.5 3.2 2.1 3.8 0.9 1.2 2.7 – 5.8 8.5 5.0 7.5 14.2 12.1 1.9 – 6.7 4.7 6.6 13.4 2.8 – – 4.3 3.4 0.0 0.0 0.0 – – – 0.0
0.7 1.1 0.4 1.3 1.3 – 1.7 2.5 3.1 0.5 0.7 1.7 0.5 0.3 0.5 0.5 1.4 4.2 1.8 0.7 0.5 1.2 0.3 – – – 1.7 3.0 1.4 3.8 3.1 3.3 0.4 – 3.8 1.1 1.8 5.1 0.7 – – 2.1 – 0.0 0.0 0.0 – – – 0.0
6.9 10.2 7.1 8.7 11.6 – 7.8 12.2 13.1 6.9 7.6 11.8 9.2 9.6 7.1 10.5 10.8 28.5 18.7 7.2 9.6 14.1 8.9 8.4 6.4 – 10.3 11.3 12.1 7.9 16.2 13.3 13.0 – 15.2 – 11.7 22.7 7.7 – – 8.6 10.4 0.0 0.0 0. – – – 0.0
35.0 44.0 27.0 39.0 24.0 34.0 41.0 56.0 33.0 49.0 – – 38.0 46.0 65.0 98.0 72.0 112.0 32.0 41.0 29.0 37.0 51.0 70.0 16.0 22.0 15.0 21.0 21.0 27.0 13.0 18.0 43.0 63.0 24.0 56.0 24.0 63.0 39.0 49.0 19.0 26.0 22.0 33.0 9.0 12.0 13.0 16.0 38.0 46.0 22.0 24.0 49.0 69.0 60.0 86.0 35.0 54.0 85.0 103.0 51.0 90.0 73.0 106.0 9.0 20.0 13.0 15.0 36.0 49.0 22.0 38.0 48.0 75.0 48.0 87.0 32.0 41.0 57.0 68.0 40.0 45.0 47.0 56.0 26.0 37.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 4.0 – – – – 3.0 3.0
14 10 25 12 13 10 4 15 15 22 12 18 10 31 14 11 14 8 22 19 21 11 10 12 11 30? 12 8 12 4 14 13 28 10 3 20 14 7 15 20 ? 4 7 14 10 12 13 10 1 15
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Table 22 Continued Characteristics a Dorsal kineties, number
Dorsal kinety 1, number basal body pairs Dorsal kinety 2, number basal body pairs Dorsal kinety 3, number basal body pairs
Species
mean
M
SD
SE
CV
Min
Max
n
bi4 de1 de2 de3 ed1 ed2 ed3 ed4 el1 el2 el3 gra ha1 ha2 hya isl kir lon o lo1 lo2 mag mu3 pe1 pe2 pol por pr1 pr2 qua rap te1 te2 vit ed1 ed1 ed1
3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 2.3 3.0 2.5 3.0 3.0 2.5 3.0 3.0 2.0 5.0 3.0 3.0 2.8 3.0 3.1 3.0 3.0 3.0 2.0 2.0 2.0 5.0 3.0 3.0 3.0 5.8 9.9 8.8
3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 2.0 3.0 3.0 3.0 3.0 2.0 3.0 3.0 2.0 – 3.0 3.0 3.0 3.0 3.0 – 3.0 – 2.0 2.0 2.0 – – 3.0 3.0 6.0 10.0 8.5
0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 – – – 0.0 0.0 – 0.0 0.0 0.0 – 0.0 0.0 0.5 0.0 0.3 – 0.0 – 0.0 0.0 0.0 – – – 0.0 1.0 1.3 0.9
0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 – – – 0.0 0.0 – 0.0 – – – 0.0 0.0 0.1 0.0 0.1 – 0.0 – 0.0 0.0 0.0 – – – – 0.4 0.4 0.3
0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 – – – 0.0 0.0 – 0.0 0.0 0.0 – 0.0 0.0 16.4 0.0 10.1 – 0.0 – 0.0 0.0 0.0 – – – 0.0 18.0 12.6 10.1
3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 2.0 3.0 2.0 3.0 3.0 2.0 3.0 3.0 2.0 – 3.0 3.0 2.0 3.0 3.0 – 3.0 – 2.0 2.0 2.0 – – 3.0 3.0 4.0 8.0 8.0
3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 2.0 – 3.0 3.0 3.0 3.0 4.0 – 3.0 – 2.0 2.0 2.0 – – 3.0 3.0 7.0 12.0 10.0
15 22 10 18 10 30 15 11 14 8 22 19 21 11 9 12 11 30 12 8 12 13 28 10 3 ? 12 7 15 20 ? 3 7 8 8 8
a All measurements in µm. ? = number of individuals investigated (= sample size) not indicated; if only one value is known it is listed in the mean column; if two values are available they are listed as Min and Max. CV = coefficient of variation in %, M = median, Max = maximum value, mean = arithmetic mean, Min = minimum value, n = number of individuals investigated, SD = standard deviation, SE = standard error of arithmetic mean. Data based on protargol-impregnated specimens, unless otherwise indicated. b
Cirrus III/2 (= cirrus behind right frontal cirrus) included.
c
Rearmost nodule.
d
Distance between first and second nodule.
e
Distance between second and third nodule.
f
Posteriormost micronucleus.
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SYSTEMATIC SECTION
Table 22 Continued g
Distance between anterior end of cell and rearmost cirrus (long arrow in Fig. 35d) of short row behind right frontal cirrus
h
Anteriormost macronuclear nodule, respectively, micronucleus.
i
Macronuclear groups, distance in between.
j
Cirrus III/2 (= cirrus behind right frontal cirrus) included.
k
From life.
l
Pretransverse ventral cirri included.
m
Posteriormost macronuclear nodule.
n
This species invariably has two pretransverse ventral cirri, which are not included.
o
For details see text.
p
Method (from life? protargol impregnation?) not indicated.
q
Inflated micronuclei, as shown in Fig. 51e, excluded.
Borror (1972, p. 9) synonymised Uroleptoides with Amphisiella because he considered the two type species (U. kihni and A. marioni) as synonyms. Synonymy of Amphisiella and Uroleptoides was accepted by Jankowski (1979, p. 68, 69), and Foissner & Blatterer (1990). The present revision shows that true Amphisiella species, that is, the type and some related species, are very likely confined to saline waters. In addition, they are probably characterised by more than three dorsal kineties, although for some species data are lacking or have to be rechecked. The many other species originally classified in, or transferred to, Amphisiella live in terrestrial habitats and largely have another habitus, although the cirral groups are the same as in true Amphisiella species. Interestingly, most (all?) these soil species have only three or two dorsal kineties and distinctly curved undulating membranes, whereas the number of dorsal kineties is usually increased and the membranes are more or less straight in Amphisiella species. Tuffrau (1979, 1987) and Tuffrau & Fleury (1994) classified Uroleptoides in the Kahliellidae, however, without providing a detailed explanation. Foissner & Foissner (1988, p. 83) followed this suggestion. Hemberger (1982) also accepted Uroleptoides and assigned it to the Amphisiellidae, basically because of the frontoventral row. Shi et al. (1999), Shi (1999), and Lynn & Small (2002) followed this proposal. I preliminarily also accept this classification, although we do not know whether the frontoventral row of U. kihni is formed by two or three anlagen (= supposed apomorphy of the amphisiellids) or by only one anlage, as, for example, in Orthoamphisiella. By contrast, Small & Lynn (1985) who also established a family Amphisiellidae, assigned Uroleptoides to the Cladotrichidae. Shi (1999) and Shi et al. (1999) put Hemiamphisiella Foissner, 1988 and Paramphisiella Foissner, 1988 into the synonymy of Uroleptoides. However, Hemiamphisiella has a postperistomial cirrus and caudal cirri (vs. absent in Uroleptoides) and lacks transverse cirri (vs. present). Paramphisiella has only one cirrus left of the
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245
anterior portion of the amphisiellid median cirral row (vs. more than one) and caudal cirri, but lacks transverse cirri. These differences strongly indicate that the synonymies proposed by the Chinese workers are not justified. Uroleptoides is reminiscent of Lamtostyla whose type species is also not described in every detail. For example, the origin of the frontoventral row (from one or more anlagen) is not known, that is, currently we do not know whether Lamtostyla and Uroleptoides are amphisiellids or not. However, Lamtostyla lamottei lacks the second pseudorow of frontal cirri so that it would be perhaps better to classify all Uroleptoides species (except U. kihni) in Lamtostyla. On the other hand, Lamtostyla lamottei has a rather short frontoventral row, whereas it is rather long in the Uroleptoides species. Since both type species are not described in detail I retain both Uroleptoides and Lamtostyla and separate these two taxa preliminary only by the length of the amphisiellid median cirral row, namely, more than 50% of body length in Uroleptoides (Fig. 45a) against less than 50% in Lamtostyla (Fig. 30a). Of course this is only a pragmatic solution, and (very likely) not a reflection of the phylogeny. Only the redescription of both type species will allow a more proper solution. Probably the species included in Lamtostyla and Uroleptoides in the present review do not form a monophyletic group. Previously I suggested classifying Uroleptoides as subgenus of Amphisiella to avoid the combination of terrestrial Amphisiella species with Uroleptoides (Berger 2005b). However, such a classification seems not to be useful and therefore Uroleptoides is accepted as valid genus in the present revision. Species included in Uroleptoides (alphabetically arranged basionyms are given): (1) Amphisiella longiseries Foissner, Agatha & Berger, 2002; (2) Amphisiella magnigranulosa Foissner; 1988; (3) Amphisiella multinucleata Foissner, Agatha & Berger, 2002; (4) Amphisiella polycirrata Berger & Foissner, 1989; (5) Amphisiella raptans Buitkamp & Wilbert, 1974; (6) Amphisiella terricola Gellért, 1955; (7) Uroleptoides binucleata Hemberger, 1985; (8) Uroleptoides kihni Wenzel, 1953. Species misplaced in Uroleptoides: The following species originally assigned to Uroleptoides are very likely misplaced in this genus. Uroleptoides atypicus Hemberger, 1985. Remarks: Now Amphisiellides atypicus (p. 654). Uroleptoides caudata Hemberger, 1985. Remarks: Now Paramphisiella caudata (p. 358). Uroleptoides quadrinucleata Foissner, 1984. Remarks: Now Hemiamphisiella quadrinucleata (p. 318). Uroleptoides qingdaoensis Song & Wilbert, 1989. Remarks: Now Hemiamphisiella terricola qingdaoensis (p. 310). Uroleptoides vitiphila Foissner, 1987. Remarks: Now Lamtostyla vitiphila (p. 187)
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SYSTEMATIC SECTION
Key to Uroleptoides species and similar species For the separation of Uroleptoides species, live data (presence/absence of cortical granules), the nuclear apparatus, and details of the cirral pattern (number of buccal cirri, length of amphisiellid median cirral row, presence/absence of caudal cirri) are needed. If you cannot identify your specimen/population with the key below, see also Lamtostyla and Lamtostylides. 1 2 3 4 5 6 7 8 9
-
Two macronuclear nodules. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Four or more macronuclear nodules. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Cortical granules lacking. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Cortical granules present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 One buccal cirrus (Fig. 54e). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4–6 buccal cirri (Fig. 55a, c). . . . . . . . . . . . . . Uroleptoides polycirratus (p. 285) Amphisiellid median cirral row extends distinctly beyond mid-body (Fig. 54a, b, e, f). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Uroleptoides terricola (p. 279) Amphisiellid median cirral row ends at about mid-body (Fig. 32f). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amphisiella australis sensu Foissner (1988) (p. 179) (2) Amphisiellid median cirral row terminates at 50% of body length on average (Fig. 52a, b). . . . . . . . . . . . . . . . . . . . . . . Uroleptoides magnigranulosus (p. 273) Amphisiellid median cirral row terminates at 60–82% of body length on average (Fig. 50a, h, 51a, e). . . . . . . . . . . . . . . . . . . . . . Uroleptoides binucleatus (p. 261) (1) Four macronuclear nodules (Fig. 46a, e, g). Uroleptoides longiseries (p. 249) More than four macronuclear nodules. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Eight macronuclear nodules; body length 300–500 µm (Fig. 47a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Uroleptoides raptans (p. 252) More than 8 macronuclear nodules; body length less than 350 µm. . . . . . . . . . . 8 12–15 macronuclear nodules arranged in two rows (Fig. 45a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Uroleptoides kihni (p. 246) 17–38, on average more than 20 macronuclear nodules. . . . . . . . . . . . . . . . . . . . 9 Caudal cirri lacking; amphisiellid median cirral row terminates at 71% of body length on average; 3 dorsal kineties (Fig. 48a–l). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Uroleptoides multinucleatus (p. 255) Caudal cirri present; frontoventral row terminates at 55% of body length on average; 4 dorsal kineties (Fig. 135a, b). . . . . . . . . . . Amphisiellides atypicus (p. 654)
Uroleptoides kihni Wenzel, 1953 (Fig. 45a) 1953 Uroleptoides kihni nov. gen. nov. spec. – Wenzel, Arch. Protistenk., 99: 107, Abb. 18 (Fig. 45a; original description; no formal diagnosis provided and very likely no type material available).
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247
1974 Uroleptoides kihni Wenzel – Stiller, Fauna Hung., 115: 61, 62 (Fig. 45a; guide to Hungarian ciliates). 1979 Amphisiella kihni comb. n. – Jankowski, Trudy zool. Inst., 86: 69 (combination with Amphisiella). 1982 Uroleptoides kihni Wenzel, 1953 – Hemberger, Dissertation, p. 49 (revision of hypotrichs). 1983 Uroleptoides – Curds, Gates & Roberts, Synopses of the British fauna, 23: 419, Fig. 248 (redrawing of Fig. 45a; guide to British ciliates). 2001 Uroleptoides kihni Wenzel, 1953 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 96 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs).
Nomenclature: Wenzel (1953) dedicated this species to Berthold Kihn, a doctor in the city of Erlangen, Germany. Type species of Uroleptoides. Borror (1972) synonymised Amphisiella marioni and Uroleptoides kihni. By contrast, Jankowski (1979) considered U. kihni as valid species within Amphisiella and therefore made the new combination. Remarks: Uroleptoides kihni is described only after live observation and fixation (Wenzel 1953). Thus, the exact cirral pattern is not known. Especially the three cirri behind the frontal cirri (Fig. 45a, arrows) are difficult to interpret (see genus section). The median row, extending from near the anterior body end to the rear cell end, and the cirri left of the anterior portion of this row indicate that U. kihni is an amphisiellid. Whether the elongated cirri at the rear body end are transverse cirri or caudal cirri is not known. The discussion in the genus section shows that the synonymy of Uroleptoides and Amphisiella, first suggested by Borror (1972, p. 9) because he synonymised the two type species, was likely overhasty. It is astonishing that U. kihni has not yet been rediscovered although European terrestrial habitats are comparatively well studied (e.g., Foissner 1998). The most important feature of U. kihni is likely the second pseudorow of frontal cirri (Fig. 45a, arrows; possibly this pseudorow is composed by the buccal cirrus, cirrus III/2, and the front cirrus of anlage IV). Together with the conspicuous nuclear apparatus it should make U. kihni unmistakable. Detailed redescription and neotypification, inter alia, because no type locality is fixed, are urgently needed. Uroleptoides kihni sensu Borror & Evans (1979) is probably identical with Afroamphisiella multinucleata (see there for details). Morphology: Body length 95–140 µm (method not indicated, likely after fixation with osmium), on average 103 µm (n = 6); body length:width ratio of specimen illustrated about 5.7:1 (Fig. 45a). Body outline slender, margins of posterior portion converging, rear end therefore very narrowly rounded. Body more or less distinctly flattened dorsoventrally. Nuclear apparatus difficult to recognise in life; about 12–15 macronuclear nodules arranged in two rows in postoral region; individual nodules about 8 × 4 µm. One micronucleus (about 2–3 µm across) each between anterior and posterior end of macronuclear rows (Fig. 45a). Contractile vacuole about in midbody near left cell margin, during diastole with distinct collecting canals extending from about proximal end of adoral zone to rear body fourth. Presence/absence of cortical granules and details of cytoplasm (colour, inclusions) not described. Movement slow and continuous, not very skilful.
248
SYSTEMATIC SECTION Fig. 45a Uroleptoides kihni (from Wenzel 1953. From life and after fixation and nucleus stain). Ventral view showing infraciliature (details must not be over-interpreted), nuclear apparatus, and contractile vacuole, 110 µm. Arrows mark second pseudorow of frontal cirri. ACR = amphisiellid median cirral row, AZM = distal end of adoral zone of membranelles, CC? = caudal or transverse cirri, CV = contractile vacuole with distinct collecting canals, FC = right frontal cirrus, MI = micronucleus. Page 246.
Adoral zone occupies about one fifth of body length (25% of body length in specimen illustrated), distal end at anterior body end. Buccal field obviously narrow, right border with high lip bearing triangular paroral. Cirral pattern basically amphisiellid. Three enlarged frontal cirri and immediately behind a second pseudorow of slightly smaller cirri (Fig. 45a, arrows). Such a double pseudorow (basically a short “bicorona”) is very uncommon and has to be confirmed; possibly it is due to a misinterpretation of the preparations (perhaps these are [from left to right] the buccal cirrus, cirrus III/2, and the front cirrus of anlage IV). Four somewhat weaker cirri form oblique row extending across frontal area (likely these are the “cirri left of the anterior portion of the amphisiellid median cirral row”). Amphisiellid median cirral row extends slightly sigmoidally from near distal end of adoral zone to near rear body end; composed of 27 cirri in specimen illustrated. Rear body end with four slightly moveable cirri, which project 8–10 µm beyond rear body end; whether these cirri are transverse or caudal cirri is not known. Right marginal row obviously commencing dorsolaterally, terminates – like left row – slightly ahead of rear body end; left row commences near proximal end of adoral zone. Dorsal ciliature, that is, length of bristles, number and arrangement of kineties, and presence/absence of caudal cirri, not described; thus, higher level classification very uncertain. Occurrence and ecology: Wenzel (1953) discovered Uroleptoides kihni in Bavaria, Germany. He studied samples from three areas, namely from the suburbs of the city of Erlangen, from the Franconian Jura (Güßweinstein), and from the suburbs of the village of Seeon (Lake Chiemsee area). Unfortunately, he forgot to mention in which of these areas he found the present species (see remarks). It occurred in dry mosses, leave litter, and in sterile exposed leave and Sphagnum samples. Sudzuki (1979, p. 124; without morphological data) found it in soil and moss samples from the Antarctic region. Likely par lapsus, Patterson et al. (1989, p. 211) listed U. kihni in the list of marine benthic ciliates. Food not known.
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249
Uroleptoides longiseries (Foissner, Agatha & Berger, 2002) comb. nov. (Fig. 46a–g, Table 22) 2002 Amphisiella longiseries nov. spec.1 – Foissner, Agatha & Berger, Denisia, 5: 680, Fig. 151a–g, Tables 135, 136 (Fig. 46a–g; original description; one holotype slide [accession number 2002/466], five paratype slides [2002/467–471], and three voucher slides [2002/472–474] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria).
Nomenclature: The species-group name longiseries is a composite of long·us, -a, -um (Latin adjective [m, f, n]; long), the thematic vowel ·i-, and series (Latin noun; row), referring to the long amphisiellid median cirral row (Foissner et al. 2002). Remarks: For a foundation of the exclusion from Amphisiella, see genus section. The present species has a long (70% of body length) amphisiellid median cirral row and is thus transferred to Uroleptoides (details see genus section). Moreover, many Uroleptoides species have – like U. longiseries – clusters of cortical granules around the cirri and dorsal bristles (see also genus section). However, they can be easily distinguished by the number of macronuclear nodules: two in U. binucleatus; four in U. longiseries; eight in U. raptans; and 34–38 in U. multinucleatus. Uroleptoides longiseries occurred together with U. magnigranulosus (Foissner et al. 2002). These two species can be easily distinguished by the length of the amphisiellid median cirral row (ending behind vs. near mid-body) and the number of macronuclear nodules (4 vs. 2). Morphology: We studied two populations (Foissner et al. 2002). However, since they are from rather distant locations, namely Namibia (Fig. 46a–e) and Israel (Fig. 46f, g), data are kept separate and the diagnosis and description contain only observations from the type population. The Israeli specimens, which were not studied in life, are highly similar to the Namibian type population, both in morphometric features and the rare characteristic that the cortical granules impregnate with protargol (Fig. 46f, g, Table 22). Body size 160–230 × 25–40 µm in life, usually about 190 × 30 µm; length:width ratio in life about 5–7:1, in protargol preparations 3.2–8.1:1, on average 4.7:1, that is, slightly stouter than in life because more or less distinctly inflated by the preparation procedures (Table 22). Body outline slenderly pisciform with anterior end slightly more broadly rounded than posterior; rear portion often almost tail-like and curved rightwards, a feature preserved even in most protargol-impregnated specimens; body very flexible but acontractile, often slightly to distinctly twisted about main axis (Fig. 46a, d). Macronuclear nodules serially in middle third of cell slightly left of midline, individual nodules ellipsoidal and with small to medium-sized chro1
Foissner et al. (2002) provided the following diagnosis: Size about 190 × 30 µm in vivo; very slenderly lanceolate. 4 macronuclear nodules. Cortical granules colourless, 0.5–1.5 µm across, in clusters around cirri and dorsal bristles. Amphisiellid median cirral row (ACR) extends beyond mid-body, composed of about 39 cirri. On average 24 adoral membranelles, about 56 cirri each in right and left marginal row, 3 cirri left of ACR, 1 buccal cirrus, 4 transverse cirri very near to posterior body end, and 3 dorsal kineties.
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SYSTEMATIC SECTION
Fig. 46a–e Uroleptoides longiseries (type population from Namibia; from Foissner et al. 2002. a–c, from life; d, e, protargol impregnation). a: Ventral view of a representative specimen (195 µm) having ingested a long bacterial rod and a small ciliate. Note the short adoral zone of membranelles and the deep buccal cavity. b, c: The cortical granules around cirri and dorsal bristles are colourless and 0.5–1.5 µm across, as in many other species of the group. d, e: Infraciliature of ventral and dorsal side and nuclear apparatus of holotype specimen (173 µm), which is rather strongly inflated due to the preparation procedures. Arrow marks the three cirri (cirrus behind right frontal cirrus included) left of amphisiellid median cirral row. Broken lines connect cirri which very likely originate from the same anlage; dotted line connects frontal
Uroleptoides matin bodies. Usually one micronucleus attached to anterior and to posterior macronuclear nodule, posterior micronucleus sometimes between third and fourth nodule; individual micronuclei ellipsoidal, occasionally pyriform, conspicuous because large, that is, about 5–6 × 3 µm in life. Contractile vacuole slightly ahead of midbody, during diastole with lacunar collecting canals. Cortical granules as in U. magnigranulosus, U. multinucleatus, and U. binucleatus multicirratus, that is, found only around bases of cirri and dorsal bristles, colourless and globular; granules around bristle rows 1 and 3 usually impregnate with protargol (Fig. 46d, e), while granules of row 2 rarely impregnate, possibly due to a preparation artifact because they impregnate in the Israeli specimens (Fig. 46g). Granules around cirri about 0.5 µm across, clusters around dorsal bristles composed of small (0.5 µm) and large (up to 1.5 µm) granules (Fig. 46b, c). Cytoplasm colourless, contains many lipid droplets up to 5 µm across mainly in posterior body portion, and some food vacuoles. Movement without peculiarities, that is, swims and glides rather rapidly on microscope slide and debris showing great flexibility. Adoral zone occupies 14–22%, on average 17% of body length, of usual shape and structure, except for proximal half, which widens slightly spoon-like (Fig. 46d); composed of an
b
251
Fig. 46f, g Uroleptoides longiseries (from Foissner et al. 2002. Protargol impregnation). Infraciliature of ventral and dorsal side and nuclear apparatus of an Israeli specimen, 183 µm. In this population, the cortical granules of all dorsal kineties impregnate. Page 249.
cirri. Note that the cortical granules are not impregnated in bristle kinety 2, which has a short break in this specimen. ACR = amphisiellid median cirral row, AZM = adoral zone of membranelles, CG = cortical granules, CV = contractile vacuole, FC = right frontal cirrus, LMR = left marginal row, MA = macronuclear nodule, MI = micronucleus, P = paroral, RMR = right marginal row, TC = transverse cirri, 1–3 = dorsal kineties. Page 249.
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average of 24 membranelles, bases of largest membranelles about 7 µm wide in life. Buccal cavity deep, as in Cyrtohymena (for review see Berger 1999), right margin forms curved lip covering proximal adoral membranelles and bearing paroral composed of about 10 µm long, closely spaced cilia. Undulating membranes distinctly curved, optically intersecting near mid buccal cavity. Pharyngeal fibres prominent in life and in protargol preparations. Cirral pattern and number of cirri of usual variability (Fig. 46a, d, f, Table 22). Frontal cirri form concave row; cirri distinctly enlarged and longer than those of marginal rows and amphisiellid median row. Buccal cirrus at summit of curve formed by undulating membranes. Usually three cirri left of anterior portion of amphisiellid median cirral row, anterior cirrus (likely this is cirrus III/2) behind right frontal cirrus, middle and posterior cirrus shifted slightly rightwards. Amphisiellid median cirral row conspicuous because terminating at 70% of body length on average, commences right of distal end of adoral zone and extends obliquely to midline in posterior body portion. Transverse cirri terminal and thus distinctly projecting, bases not larger than those of marginal cirri. Right marginal row extends onto dorsolateral surface anteriorly, terminates slightly ahead of right transverse cirrus; left row commences left of proximal end of adoral zone, ends at same level as right row or even on dorsal side of rear end. Cirri of amphisiellid median cirral row and of marginal rows short compared to size of cell, that is, 8–10 µm long in life, usually composed of two rows with four cilia each, closely spaced within rows, distance between individual cirri increase slightly in posterior portion of marginal rows. Dorsal bristles about 3 µm long in life, arranged in three bipolar kineties (Fig. 46e, g). Caudal cirri lacking. Occurrence and ecology: Uroleptoides longiseries is likely confined to terrestrial habitats (Foissner et al. 2002, p. 50). The type locality is the bank of the Bukaos River (25°40'S 18°10'E), about 80 km north of the town of Keetmanshoop, Namibia, where we found it in sieved litter. The Israeli population is from the Golan Heights, where it occurred in the upper layer of a moist, dark moder soil of an uncultivated grassland dominated by Poa sp. (collected on February 14, 1985 by Johannes Augustin, Salzburg). Uroleptoides longiseries, which is well-adapted to soil life by the slender body, was rare at both sites. Feeds on small ciliates (Colpoda maupasi) and filamentous bacteria (Foissner et al. 2002).
Uroleptoides raptans (Buitkamp & Wilbert, 1974) Hemberger, 1982 (Fig. 47a) 1974 Amphisiella raptans n. sp. – Buitkamp & Wilbert, Acta Protozool., 13: 205, Abb. 4 (Fig. 47a; original description; no formal diagnosis provided; the type slides are likely deposited in the Institut für landwirtschaftliche Zoologie der Universität Bonn, Germany).
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1976 Amphisiella raptans Buitkamp & Wilbert, 1974 – Bick & Buitkamp, Trans. Am. microsc. Soc., 95: 490, Fig. 3 (Fig. 47a; brief note on Matador Project). 1982 Uroleptoides raptans (Buitkamp & Wilbert, 1974) n. comb. – Hemberger, Dissertation, p. 60 (combination with Uroleptoides; see nomenclature). 2001 Amphisiella raptans Buitkamp and Wilbert, 1974 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 8 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Amphisiella raptans Buitkamp & Wilbert, 1974 – Foissner, Agatha & Berger, Denisia, 5: 691, Fig. 152h (Fig. 47a; comparison with Uroleptoides multinucleatus).
Nomenclature: No derivation of the name is given in the original description. The species-group name raptans (Latin; predate) refers to the fact that the present species is a voracious predator. Hemberger (1982) has transferred the present species to Uroleptoides. Whether or not this act is valid is not yet clarified (e.g., Foissner 1982, Hemberger 1985). According to the relevant code (ICZN 1964), dissertations obviously meet the criteria of publication because they are not mentioned in Article 9, which lists all acts that do not constitute a publication. Thus, I accept Hemberger (1982) as combining author. However, note that for the new species described by Hemberger (1982), the paper “Hemberger (1985)” is considered as authoritative (see U. binucleatus). Remarks: For the foundation of the classification in Uroleptoides see genus section. Redescription from life recommended. Uroleptoides raptans is a very large predator, whose body size (300–500 µm in protargol preparations?) separates it distinctly from other multinucleate Uroleptoides species, such as U. multinucleatus, which has a very similar body shape (Fig. 48a–l). This 175–300 µm long species has cortical granules and more (on average 26 vs. 8) macronuclear nodules. However, cortical granules might have been overlooked by Buitkamp & Wilbert (1974), who did not study live specimens in detail. If further investigations show that U. raptans has cortical granules, Uroleptoides multinucleatus should probably considered as subspecies of U. raptans with smaller size and distinctly more macronuclear nodules as distinguishing features (Foissner et al. 2002). Morphology: Body length 300–500 µm in protargol(?) preparations; body length:width ratio of specimen illustrated 5:1 (Fig. 47a). Body outline elongate with margins of anterior portion parallel, margins of posterior portion converging posteriorly, anterior end therefore broadly, posterior narrowly rounded; rear body half twisted about main body axis by half a turn. Rearmost fifth of cell translucent, remaining portion cloudily granulated. Body soft and flexible. Nuclear apparatus composed of eight ellipsoidal macronuclear nodules containing several small chromatin bodies, and four micronuclei. Contractile vacuole near left cell margin in first body third (in specimen illustrated, however, at 36% of body length; Fig. 47a), forms a vesicular elevation on dorsal surface. Presence/absence of cortical granules not mentioned. Movement rapidly searching; seemingly aggressive. Adoral zone occupies about 20% of body length, composed of ca. 33 membranelles. Buccal field wide (only in protargol preparations?), buccal lip distinctly curved leftwards anteriorly. No further details described or illustrated.
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SYSTEMATIC SECTION Fig. 47a Uroleptoides raptans (from Buitkamp & Wilbert 1974. Protargol impregnation). Infraciliature of ventral side and nuclear apparatus, about 400 µm. Arrow marks buccal cirrus, arrowhead denotes the two cirri left of the amphisiellid median cirral row. Asterisk marks cirrus III/2. Dotted line connects frontal cirri; broken lines connect cirri which very likely originate from same anlage (has to be confirmed by ontogenetic data). Hemberger (1982) supposed that the seven cirri in the frontal area develop from three anlagen. ACR = amphisiellid median cirral row, AZM = adoral zone of membranelles, CV = contractile vacuole, LMR = left marginal row, MA = macronuclear nodule, MI = micronucleus, P = paroral(?), PF = pharyngeal fibres, RMR = right marginal row, TC = transverse cirri. Page 252.
Frontal field with seven prominent cirri; according to my interpretation these are three frontal cirri along the distal portion of the adoral zone, one buccal cirrus at middle portion of buccal lip, one cirrus (= cirrus III/2) behind right frontal cirrus, and two cirri left of anterior portion of amphisiellid median cirral row. Amphisiellid median cirral row conspicuous because extending from distal end of adoral zone to near posterior cell end and composed of about 77 cirri in specimen illustrated (Fig. 47a). Three terminally arranged transverse cirri, not distinctly longer than marginal cirri, that is, about 10 to 12 µm. Each marginal row composed of about 80 cirri, not clearly separated posteriorly. Dorsal bristles 4–5 µm long; number and arrangement of kineties and presence/absence of caudal cirri not mentioned. Occurrence and ecology: Uroleptoides raptans is very likely confined to terrestrial habitats (Foissner 1987, 1998). The type locality is the Sceptre Association of the Brown Soil Zone (0–10 cm; Chernozemic brown soils of heavy clay texture, moderately calcareous) in the Prairie of southern Saskatchewan, Canada. Buitkamp & Wilbert (1974) discovered it during the course of the “Matador Project” (Matador is a village at 50°48'07"N 107°56'55"W). The vegetation of this area belongs to the Agropyron-Koeleria Faciation of the Mixed Prairie (StipaBouteloua) Association (Bick & Buitkamp 1976). Sudzuki (1979, p. 123) found U. raptans in Antarctic soil (no morphological data provided). Recently, we found it in a floodplain forest stand (Müllerboden) in Austria (Foissner et al. 2005, p. 624). Uroleptoides raptans is a very voracious predator, which, in a culture of Colpoda steinii, ingested twelve, about 30 µm-sized specimens per minute; feeds also
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on amoebas (Buitkamp & Wilbert 1974). Biomass of 106 specimens about 1280 mg (Foissner 1987, p. 121; 1998, p. 199).
Uroleptoides multinucleatus (Foissner, Agatha & Berger, 2002) comb. nov. (Fig. 48a–l, 49a, b, Table 22) 2002 Amphisiella multinucleata nov. spec.1 – Foissner, Agatha & Berger, Denisia, 5: 685, Fig. 152a–g, i–m, 387a–e, Table 137 (Fig. 48a–l, 49a, b; original description; one holotype slide [accession number 2002/391], two paratype slides [2002/392, 393], and six voucher slides [2002/394–399] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria).
Nomenclature: The species-group name multinucleatus is a composite of the Latin quantifier mult·us (many), the thematic vowel ·i-, and the Latin adjective nucleát·us, -a, -um ([m; f; n]; kernel-like), referring to the many macronuclear nodules (Foissner et al. 2002). Amphisiella is feminine, Uroleptoides is masculine. Thus, the ending of the species-group name has to be changed from multinucleat·a to multinucleat·us (ICZN 1999). Remarks: For a foundation of the exclusion from Amphisiella, see genus section. The present species is assigned to Uroleptoides because the median cirral row extends beyond mid-body (for details, see genus section). Moreover, most (all?) Uroleptoides species have clusters of cortical granules around the cirri and dorsal bristles. Uroleptoides multinucleatus is obviously most closely related to U. raptans (Fig. 47a). This 300–500 µm long species from a Canadian prairie soil has only eight macronuclear nodules (vs. more than 26 on average in U. multinucleatus) and lacks cortical granules. However, such granules might have been overlooked by Buitkamp & Wilbert (1974), who did not study live specimens in detail. If further investigations show that U. raptans has cortical granules, the present species should probably be considered as subspecies with smaller body size and distinctly more macronuclear nodules as distinguishing features (Foissner et al. 2002). As concerns the cortical granules, their pattern and size highly resembles that of U. magnigranulosus, which, however, has only two macronuclear nodules. Paramphisiella acuta (Fig. 70a–j) differs from U. multinucleatus by the generic features and the lack of cortical granules (checked in protargol slides), the shorter amphisiellid median cirral row (40% vs. 53–71% of body length on average), and the distinctly lower number of adoral mem1
Foissner et al. (2002) provided the following diagnosis (includes data from type population and Kenyan population): Size about 250–300 × 30–40 µm in vivo. Very slender and narrowed posteriorly. Cortical granules colourless, form clusters composed of small (about 0.5 µm) and large (up to 3 µm across) globules around cirri and dorsal bristles. Amphisiellid median cirral row (ACR) extends beyond mid-body, composed of an average of 53–61 cirri. On average 35–48 macronuclear nodules, 30 adoral membranelles, about 90 cirri each in right and left marginal row, 4 cirri left of ACR, 1 buccal cirrus, 3 transverse cirri, and 3 dorsal kineties.
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Fig. 48a–g Uroleptoides multinucleatus (from Foissner et al. 2002. a–e, from life; f, g, protargol impregnation). a: Ventral view of a representative specimen, 290 µm. b–d: Cortical granules around cirri (b, c) and dorsal bristles (d), granules 0.5–3.0 µm across. e: Strongly twisted specimen. f, g: Infraciliature of ventral and dorsal side and nuclear apparatus of holotype specimen, 285 µm. Arrow denotes a single cirrus left of anterior end of amphisiellid median cirral row (note variability of this feature). Dorsal bristle rows impregnated incompletely and are thus not shown. ACR = anterior end of amphisiellid median cirral row, BC = buccal cirrus, CV = contractile vacuole with collecting canals, E = endoral, FC = right frontal cirrus,
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branelles (about 15 vs. 28–30) and marginal cirri (45 left and 46 right vs. 69–88 and 72–95 on average). In life, Uroleptoides multinucleatus is characterised by the following combination of features (Foissner et al. 2002): large and slender body (about 175–300 × 25–40 µm; ratio of body length:width about 7–8:1); about 26–48 macronuclear nodules; cortical granules around cirri and dorsal bristles; amphisiellid median cirral row usually extending to posterior third of body. In eight protargol slides from the type locality, we found only six specimens, including one reorganiser (Foissner et al. 2002). Furthermore, the impregnation was of mediocre quality because the fixative had to be amended with osmium tetroxide for better preservation of this fragile organism. Thus, we could not clarify all details necessary for a perfect description and generic classification. However, we later found three further populations, whereby the one from Kenya matches the type material very well in almost all features, so that conspecificity is beyond reasonable doubt (Fig. 48h–l, Table 22). The specimens from Namibia (site 52; Table 22) and Venezuela (Fig. 49a, b) also agree with the type population in the main features, such as cirral pattern, oral apparatus including number of adoral membranelles, nuclear figure, and cortical granulation. However, Foissner et al. (2002) kept their data separate, while the Kenyan observations, reported below, are included in the diagnosis (see corresponding footnote). Morphology: As already mentioned in the last paragraph of the previous chapter, the descriptions/data of the four populations studied by Foissner et al. (2002) are kept separate. The type population from the Bambatsi Guest Farm in Namibia is described first. Body size of type population 250–350 × 30–50 µm in life, length:width ratio about 7:1 both in life and after protargol impregnation (Table 22). Body outline very elongate cuneate with anterior end more broadly rounded than posterior. Body very flexible and slightly to distinctly twisted about main axis (Fig. 48a, e). Macronuclear nodules in second to fifth sixth of cell, globular, ellipsoidal to elongate ellipsoidal, or even slightly dumb-bell-shaped, on average ellipsoidal (length:width ratio 2:1); chromatin bodies numerous and minute. Micronuclei about 4 µm across in life. Contractile vacuole distinctly ahead of mid-body at left cell margin, during diastole with conspicuous collecting canals. Cortical granules around cirri and dorsal bristles, colourless and globular, occasionally some impregnate with protargol; granules around cirri usually about 0.8 µm across, those around cirri of amphisiellid median row sometimes larger; clusters around dorsal bristles composed of small (about 0.5 µm) and large (up to 3 µm) granules (Fig. 48b–d). Cytoplasm colourless, contains many greasily shining globules 1–3 µm in diameter. Food vacuoles up to 50 µm across. Movement conspicuous because very slow and serpentine, showing great flexibility between soil particles.
b LMR = left marginal row, MA = macronuclear nodules, MI = micronucleus, P = paroral, RMR = right marginal row, TC = transverse cirri. Page 255.
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Fig. 48h–l Uroleptoides multinucleatus (Kenyan population from Foissner et al. 2002. h, i, from life; j–l, protargol impregnation). h: Ventral view of a representative specimen, 230 µm. i: Cortical granules around bristles occasionally impregnate with protargol. j, k: Infraciliature of ventral and dorsal side and nuclear apparatus of a specimen with four cirri (circled by dotted line) left of anterior end of amphisiellid median cirral row. Frontal cirri connected by dotted line; right frontal cirrus and cirrus III/2 connected by broken line. l: Ventral view of a specimen with three cirri (circled) left of anterior end of amphisiellid median cirral row. ACR = anterior (l) and posterior (j) end of amphisiellid median cirral row,
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Fig. 49a, b Uroleptoides multinucleatus (from Foissner et al. 2002. Scanning electron micrographs of Venezuelan specimens). Ventral views of oral region showing, inter alia, the three cirri (small arrows in a) left of the amphisiellid median cirral row, the buccal cirrus at the summit of the paroral, and the large bottom of the buccal lip (large arrows) separating the buccal cavity from the prominent paroral composed of a distinctly curved row of tightly spaced cilia. AZM = adoral zone of membranelles, BC = buccal cirrus, FC = right (a) and left (b) frontal cirrus, FS = crenelated frontal scutum, P = paroral. Page 255.
Adoral zone occupies only 13–18%, on average 15% of body length, of usual shape and structure, except for proximal half, which widens spoon-like (Fig. 48f, g; Table 22); composed of an average of 30 membranelles, bases of largest membranelles about 12 µm wide in life. Buccal cavity deep and large, as in Cyrtohymena, right posterior margin forms inconspicuous lip covering only few proximal adoral membranelles (but see observations on the Venezuelan population below). Undulating membranes distinctly curved and optically intersecting near mid, roughly as in Cyrtohymena (for review of this oxytrichid group see Berger 1999). Pharyngeal fibres very prominent in life and protargol preparations, extend obliquely backwards, likely mixed with long endoral cilia.
b LMR = left marginal cirral row, MI = micronuclei, PF = pharyngeal fibres, RMR = right marginal row, TC = transverse cirri, 1–3 = dorsal kineties. Page 255.
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Cirral and dorsal bristle pattern could not be recognised in every detail (but see Kenyan population described below). Frontal cirri about 15 µm long in life and distinctly enlarged, right one, as is usual, at distal end of adoral zone of membranelles (Fig. 48f). Buccal cirrus close to summit of paroral. Number of cirri left of anterior portion of amphisiellid median cirral row difficult to determine due to reasons mentioned in the remarks: one specimen unequivocally with only one cirrus, three specimens very likely with two cirri, and one likely with three. Amphisiellid median cirral row begins close to distal end of adoral zone and terminates at 71% of body length on average, composed of about 61 cirri. Transverse cirri terminal, only 15 µm long and therefore inconspicuous in life; prominent fibres extend from transverse cirri anteriad. Marginal cirri about 10 µm long in life, that is, short compared to size of cell; arranged in two rows which are not shortened posteriorly; right row extends onto dorsal surface anteriorly; left row commences left of proximal end of adoral zone. Dorsal bristles 3–4 µm long in life, likely arranged in three rows. Caudal cirri lacking (Fig. 48f, g). Observations on a Kenyan population (Fig. 48h–l; Table 22): As mentioned in the remarks and in the occurrence section, Uroleptoides multinucleatus was found also in a soil sample from Kenya by Foissner et al. (2002). The data match well, suggesting conspecificity. Some supplementary observations and a more detailed morphometric analysis define the species more exactly: (i) cells distinctly broadened in the protargol slides because some in vivo measurements show that they are as slender as those from Namibia, namely, 260 × 30 µm, 230 × 32 µm, 230 × 30 µm; (ii) morphometrics independent of the preparation procedures match very well, for instance, the number of adoral membranelles and marginal cirri; (iii) cortical granules, although rather large, easily overlooked because very hyaline, but occasionally impregnate with protargol; (iv) all in vivo observations match well; (v) cirri conspicuously short, namely, 8 µm, as in the Namibian specimens, except the transverse cirri, which are 20 µm long; (vi) there are 3–6, usually four cirri right of the amphisiellid median cirral row and 2–5, on average three transverse cirri in a short, oblique row; (vii) buccal field deep and almost semicircular; (viii) three bipolar dorsal kineties without caudal cirri. Observations on the Namibian site (52) population (Table 22): Size 140–210 × 20–35 µm in life and thus about one third smaller than type and Kenyan specimens; length:width ratio around 7:1 both in life and after protargol impregnation. Micronuclei ellipsoidal. Cytoplasm with many fat globules 1–10 µm across and 10 µm-sized food vacuoles containing heterotrophic flagellates. Cortical granulation as in type population. Almost invariably four transverse cirri. Three dorsal kineties easily recognizable in life due to the cortical granule clusters around the individual bristles. Paroral cilia up to 10 µm long and very closely spaced forming a conspicuous, almost semicircular, plate-like membrane. Observations on the oral apparatus of a Venezuelan population (Fig. 49a, b): The South American specimens resemble the Namibian site 52 population in most mor-
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phometrics, that is, cells are about one third smaller than those of the type population. The SEM micrographs show a peculiarity, namely, a large plate, likely the buccal lip, covering the semicircular, Cyrtohymena-like buccal field (Fig. 49a, b; see also Foissner & AL-Rasheid 2006). Obviously, this structure is very hyaline because it was not recognized in life. Furthermore, it is masked by the paroral, whose tightly spaced, up to 10 µm long cilia form a velum of similar size and shape. Further details, see figure explanations. Occurrence and ecology: Uroleptoides multinucleatus is very likely confined to terrestrial habitats (Foissner et al. 2002, p. 50). Type locality is the Bambatsi Guest Farm (20°10'S 15°25'E; 1150 m above sea-level) between the towns of Khorixas and Outjo in Namibia, where Foissner et al. (2002) discovered it with very low abundance in mud and soil of the Mopane savannah. In addition, it occurred with moderate abundance in the same area (= Namibian site 52) in a cow dung ball formed by a large Scarabaeus in the Mopane forest (pH 6.7). The Kenyan population was found in red soil (pH 6.6) under shrubs and trees in the Escarpment Mountains near the village of Limuru, surroundings of Nairobi; the abundance was high. The Venezuelan population occurred in mosses from a rain forest near the village of Pavoni, south of Puerto Ayacucho (05°40'N 67°38'W; Foissner et al. 2002). Uroleptoides multinucleatus is a voracious predator ingesting ciliates and flagellates like Polytomella (Foissner et al. 2002).
Uroleptoides binucleatus Hemberger, 1985 (Fig. 50a–h, 51a–i, Table 22) 1982 Uroleptoides binucleata n. spec.1 – Hemberger, Dissertation2, p. 52, Abb. 8a, b (Fig. 50a–d; see nomenclature). 1985 Uroleptoides binucleata n. spec. – Hemberger, Arch. Protistenk., 130: 401, Abb. 5 (Fig. 50a, b; original description; no formal diagnosis provided; type slides are deposited in the Institut für landwirtschaftliche Zoologie, University of Bonn, Germany). 1988 Amphisiella binucleata (Hemberger, 1985) nov. comb. – Foissner, Stapfia, 17: 113 (combination with Amphisiella).
1
This name is disclaimed for nomenclatural purposes (ICZN 1999, Article 8.3). See next footnote. According to the ICZN (1964), dissertations are not explicitly mentioned under Article 9, which describes all those acts that do not constitute a publication within the meaning of the Code. Thus, Hemberger’s (1982) thesis, for which the ICZN (1964) has to be applied, could possibly also be considered as original description of the present genus and many other taxa first described in this paper. Unfortunately, the situation is rather complicated and almost each thesis would need a detailed analysis whether or not it meets the requirements of publication (P. Tubbs, ICZN, Natural History Museum, London, pers. comm.). Thus, I do not accept Hemberger (1982), but Hemberger (1985) as original description of all the taxa discovered by him. However, I include the thesis in the list of synonyms because it is one of the most important papers on hypotrichs since Kahl (1932). To avoid nomenclatural problems each new name mentioned by Hemberger (1982) is individually disclaimed for nomenclatural purposes (see corresponding footnotes).
2
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1989 Uroleptoides binucleata Hemberger, 1985 – Berger & Foissner, Bull. Br. Mus. nat. Hist., 55: 32, Fig. 38–41, Tables 1, 8 (Fig. 50e–h; redescription; two voucher slides [reference numbers 1988:2:1:24, 1988:2:1:25] are deposited in the British Museum of Natural History in London; see nomenclature). 2001 Uroleptoides binucleata Hemberger, 1985 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 96 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Amphisiella binucleata (Hemberger, 1985) Foissner, 19881 – Foissner, Agatha & Berger, Denisia, 5: 663 (division in two subspecies; improved diagnosis). 2002 Amphisiella binucleata binucleata (Hemberger, 1985) Foissner, 1988 nov. stat.2 – Foissner, Agatha & Berger, Denisia, 5: 663 (establishment of subspecies). 2002 Amphisiella binucleata multicirrata nov. sspec.3 – Foissner, Agatha & Berger, Denisia, 5: 663, Fig. 148a–i, Tables 129–131 (Fig. 51a–i; original description; one holotype slide [accession number 2002/419], three paratype slides [2002/420, 423, 424], and three voucher slides [2002/483, 490, 491] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria).
Nomenclature: The species-group name binucleata (having two nuclei; Hemberger 1985) is a composite of the Latin numeral bi- (two) and the Latin adjective nucleát·us, -a, -um ([m; f; n]; kernel-like), referring to the two macronuclear nodules. Uroleptoides is masculine. Thus, the species-group name has to be changed from binucleata to binucleatus. Berger & Foissner (1989, p. 19) wrote “One slide of holotype specimens and 1 slide of paratype specimens of the new species and 1 neotype-slide of each other species described have been deposited in the British Museum (Natural History) in London”. Unfortunately, this sentence is incorrect as concerns the neotypes because it was not our intention to fix a neotype for each species redescribed, like, for example U. binucleatus, whose original type slides are deposited in the Bonn University. In that and some other cases “voucher slide” would have been the correct term. Remarks: For a foundation of the classification in Uroleptoides, see genus section. Hemberger (1982, 1985) described Uroleptoides binucleatus from a soil in Germany. Foissner (1988) transferred it to Amphisiella because he considered – like Borror (1972) and Jankowski (1979) – Uroleptoides as junior synonym of Amphisiella. However, Amphisiella species are confined to marine habitats (Berger 2004) so that the original classification in Uroleptoides seems appropriate. Berger & Foissner (1989) identified their German population with U. binucleatus because the body shape, the nuclear apparatus, the ventral cirral pattern, and the 1
Foissner et al. (2002) provided the following improved diagnosis: Size about 160–300 × 30–50 µm, usually > 200 µm long in vivo. Slenderly to very slenderly lanceolate. 2 macronuclear nodules and 3 dorsal kineties. Cortical granules colourless, inconspicuous and about 1 µm across or in conspicuous clusters composed of small (about 0.5 µm) and large (up to 3 µm across) globules around bases of cirri and dorsal bristles. Amphisiellid median cirral row extends beyond mid-body, composed of 22–67 cirri. On average 24–32 adoral membranelles, 42–81 cirri in right marginal row, 3 cirri left of amphisiellid median cirral row, 1 buccal cirrus, and 2–3 transverse cirri very near to posterior body end. 2 Foissner et al. (2002) provided the following diagnosis: Posterior body end bluntly pointed. Cortical granules inconspicuous, about 1 µm across. On average 24–25 adoral membranelles, 42–60 cirri in right marginal row, and 22–36 cirri in amphisiellid median cirral row. 3 Foissner et al. (2002) provided the following diagnosis: Posterior body portion narrowed tail-like. Cortical granules conspicuous, form clusters composed of small (about 0.5 µm) and large (up to 3 µm across) globules around bases of cirri and dorsal bristles. On average 32 adoral membranelles, 81 cirri in right marginal row, and 67 cirri in amphisiellid median cirral row.
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number of adoral membranelles agree very well with Hemberger’s type material. Differences exist in body size (type material 220–260 × 50 µm vs. 160–180 × 30 µm) and consequently in the number of right (60 vs. 38–46) and left (55 vs. 42–58) marginal cirri and cirri in the amphisiellid median cirral row (33–40 vs. 20–24). Hemberger observed three terminal cirri (transverse cirri?), whereas Berger & Foissner (1989) found four distinct transverse cirri. Hemberger (1985) used protargol impregnation (Table 22) and studied live specimens only very superficially, if at all. Thus, it is not known whether or not the type material has cortical granules. This uncertainty effectively makes the species indeterminable, because the group contains species with and without cortical granules. However, the granules are, in fact, rather difficult to discern and thus Hemberger may have overlooked them. The cortical granules of Berger & Foissner’s (1989) population, which otherwise matched the population studied by Hemberger rather well, are inconspicuous. Although the identification cannot be proven or falsified, the amended description by Berger & Foissner (1989) should be accepted because it is the most parsimonious solution of the problem. Uroleptoides magnigranulosus has a shorter amphisiellid median cirral row (Fig. 52b). Amphisiella australis sensu Foissner (1988) has a very similar general appearance, but lacks cortical granules so that synonymy with U. binucleatus can be excluded (Fig. 32a–i). Foissner et al. (2002) found, during a detailed survey of Namibian soils, populations showing distinct differences to the European material in the cortical granulation, some main morphometrics, and body shape. Thus, we split U. binucleatus into two subspecies. The improved diagnosis by Foissner et al. (2002; see corresponding footnote) is based on the data by Hemberger (1985), Berger & Foissner (1989), and Foissner et al. (2002). Subspecies included (alphabetically arranged basionyms are given): (1) Amphisiella binucleata multicirrata Foissner, Agatha & Berger, 2002; (2) Uroleptoides binucleatus binucleatus Hemberger, 1985.
Key to Uroleptoides binucleatus subspecies The separation of the two subspecies requires detailed live observation and protargol impregnation to check some main morphometrics. Amphisiella australis sensu Foissner (1988) is very similar, but lacks cortical granules (Fig. 32a–i). 1 Rear body end bluntly pointed; cortical granules inconspicuous, about 1 µm across; on average 24–25 adoral membranelles and 22–36 cirri in amphisiellid median cirral row (Fig. 50a, e, f, h). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Uroleptoides binucleatus binucleatus (p. 264) - Posterior body portion narrowed tail-like; cortical granules conspicuous, form clusters composed of small (0.5 µm) and large (3 µm) globules around bases of
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cirri and dorsal bristles; on average 32 adoral membranelles and 67 cirri in amphisiellid median cirral row (Fig. 51a, d, e). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Uroleptoides binucleatus multicirratus (p. 267)
Uroleptoides binucleatus binucleatus Hemberger, 1985 (Fig. 50a–h, Table 22) 1982 Uroleptoides binucleata n. spec.1 – Hemberger, Dissertation, p. 52, Abb. 8a, b (Fig. 50a–d; see nomenclature). 1985 Uroleptoides binucleata n. spec. – Hemberger, Arch. Protistenk., 130: 401, Abb. 5 (Fig. 50a, b; original description; no formal diagnosis provided; type slides are deposited in the Institut für landwirtschaftliche Zoologie und Bienenkunde, University of Bonn, Germany). 1988 Amphisiella binucleata (Hemberger, 1985) nov. comb. – Foissner, Stapfia, 17: 113 (combination with Amphisiella). 1989 Uroleptoides binucleata Hemberger, 1985 – Berger & Foissner, Bull. Br. Mus. nat. Hist., 55: 32, Fig. 38–41, Tables 1, 8 (Fig. 50e–h; redescription; two voucher slides [accession numbers 1988:2:1:24, 1988:2:1:25] are deposited in the British Museum of Natural History in London; see nomenclature of U. binucleatus and nomenclature below). 2001 Uroleptoides binucleata Hemberger, 1985 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 96 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Amphisiella binucleata binucleata (Hemberger, 1985) Foissner, 1988 nov. stat. – Foissner, Agatha & Berger, Denisia, 5: 663 (establishment of subspecies; diagnosis see species). 2002 Uroleptoides binucleata – Lynn & Small, Ciliophora, p. 454, Fig. 41 (Fig. 50h; guide to ciliate genera).
Nomenclature: For derivation of the species-group name see, Uroleptoides binucleatus. We established the two subspecies of U. binucleatus when the species was classified in Amphisiella (Foissner et al. 2002; see list of synonyms). Nevertheless I do not think that it is necessary to transfer the subspecies formally from Amphisiella to Uroleptoides, that is, I suppose that the name suggested in the heading is correct. Berger & Foissner (1989, p. 19) wrote “One slide of holotype specimens and 1 slide of paratype specimens of the new species and 1 neotype-slide of each other species described have been deposited in the British Museum (Natural History) in London”. Unfortunately, this sentence is incorrect as concerns the neotypes because it was not our intention to fix a neotype for each species redescribed, like, for example, Uroleptoides binucleatus, whose original type slides are deposited in the Institut für landwirtschaftliche Zoologie at the University of Bonn, Germany. In this and some other cases, “voucher slide” would have been the correct term. Remarks: The diagnosis of this subspecies by Foissner et al. (2002; see corresponding footnote at list of synonyms of U. binucleatus) is according to Hemberger (1985) and Berger & Foissner (1989). Further details, see U. binucleatus and U. binucleatus multicirratus. Morphology: This chapter is mainly based on the redescription by Berger & Foissner (1989), supplemented by Hemberger’s data. 1
See corresponding footnotes at U. binucleatus.
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Body size 160–180 × 30 µm in life; body length:width ratio of prepared specimens about 5.6:1 (Fig. 50a, e, g, Table 22); body size of type material 220–260 × 50 µm in life(?), length:width ratio 5:1 (Hemberger 1985). Body outline slender lanceolate, margins converging posteriorly, anterior end broadly rounded, posterior end bluntly pointed (Berger & Foissner 1989, Hemberger 1982, 1985). Body often slightly twisted, very flexible, inconspicuously flattened dorsoventrally. Macronuclear nodules lying almost in median of cell about in middle body third; individual nodules long ellipsoidal (length:width ratio 3.1:1; Table 22). Micronuclei about 7 × 4 µm in life, usually each one attached to a macronuclear nodule (Fig. 50b, e, h). Contractile vacuole slightly ahead of mid-body near left cell margin; collecting canals not observed (Fig. 50g); Hemberger (1982, 1985) did not describe a contractile vacuole. Cortical granules inconspicuous because less than 1 µm across and colourless, arranged around cirri and dorsal bristles and in buccal area; granules do not stain with methyl green-pyronin (reinvestigation of cortical granulation recommended; Fig. 50f); presence/absence of cortical granules not mentioned for type material. Cytoplasm colourless, densely granulated, in posterior portion many about 3 µm large yellowish greasy shining globules; food vacuoles 7–10 µm across. Rapid movement. Adoral zone occupies about 20% of body length (Berger & Foissner 1989, Hemberger 1982, 1985), composed of about 24 membranelles on average, bases of largest membranelles in life about 6 µm wide. Undulating membranes distinctly curved, optically slightly intersecting in mid-portion. Pharyngeal fibres conspicuous in life (Fig. 50a, e, h). Cirral pattern and number of cirri of usual variability (Fig. 50a, h, Table 22). Frontal cirri slightly enlarged, right one, as is usual, near distal end of adoral zone. Buccal cirrus inserted slightly behind middle of undulating membranes. One cirrus (= cirrus III/2) behind right frontal cirrus. Usually three, rarely two or four cirri (cirrus III/2 included) left of anterior portion of amphisiellid median cirral row, which extends on average to 60% of body length and is composed of an average of 23 cirri. Transverse cirri not enlarged, in life about 20 µm long (according to Hemberger 1982, 1985 about 18 µm), project distinctly beyond rear body end; Hemberger (1982, 1985) was uncertain whether the three terminal cirri were transverse cirri (Fig. 50a). Right marginal row commences at level of distal end of adoral zone, terminates about at level of transverse cirri; left row begins left of proximal end of adoral zone, extends along rear body end; marginal cirri in life about 10–12 µm long. Dorsal bristles in life about 3 µm (4 µm according to Hemberger) long, arranged in three kineties. Caudal cirri lacking. Cell division: Hemberger (1982, 1985) found a very early divider, showing that the proliferation of basal bodies for the anlagen of the opisthe commences left of the third quarter of the amphisiellid median cirral row (Fig. 50c, d). Occurrence and ecology: Uroleptoides binucleatus binucleatus is very likely confined to terrestrial habitats. So far only records from Europe have been published. The type location of U. binucleatus is not given by Hemberger (1985),
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who mentions only “suspension of mull rendzina soil”. However, this soil sample is not from Peru, the sole country mentioned under materials, but from Germany (see Hemberger 1982, p. 2 and Foissner 2000a). According to a personal communication by Norbert Wilbert (University of Bonn, Germany), the site is near the village of Mechernich (50°36'N 6°39'E), Germany. It is a beech forest on the “Kakushöhle Nordhang” and the outcrops are a middle devonian dolomite and a Pleistocene tufaceous limestone. We found U. binucleatus binucleatus in the city of Berlin, Germany, in a sample composed of litter, grass roots, and dark brown moder soil from the upper (0–4/5 cm) soil layer (pH 4.2) of a mixed coniferous/deciduous forest (with Pinus silvestris; Berger & Foissner 1989, p. 20; Foissner 2000a, p. 254). By contrast, Foissner (1998, p. 199), likely par lapsus, mentioned a paleotropic and neotropic distribution. The “neotropic” occurrence is likely due to the sparse and somewhat misleading description provided by Hemberger (1985; for explanation see above). The “paleotropic” record was, according to Foissner (1998, p. 212), provided by Foissner (1988). However, Uroleptoides binucleatus (or Amphisiella binucleata) is neither mentioned in this paper nor in the species list by Blatterer & Foissner (1988). Perhaps this record refers to A. australis sensu Foissner (1988) which is very similar to U. binucleatus binucleatus, but lacks cortical granules. Feeds very likely on heterotrophic flagellates (Berger & Foissner 1989). Biomass of 106 specimens of type population 412 mg (Foissner 1987, p. 128), of Berger & Foissner’s (1989) population only about 96 mg (Foissner 1998, p. 199).
Uroleptoides binucleatus multicirratus (Foissner, Agatha & Berger, 2002) comb. nov. (Fig. 51a–i, Table 22) 2002 Amphisiella binucleata multicirrata nov. sspec.1 – Foissner, Agatha & Berger, Denisia, 5: 663 (Fig. 51a–i; original description; one holotype slide [accession number 2002/419], three paratype slides [2002/420, 423, 424], and three voucher slides [2002/483, 490, 491] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria).
b Fig. 50a–h Uroleptoides binucleatus binucleatus (a–d, from Hemberger 1982, 1985; e–h, from Berger & Foissner 1989. a–d, h, protargol impregnation; e–g, from life). a–d: Infraciliature of ventral side and nuclear apparatus of non-divider and very early divider, 240 µm. Long arrow in (a) marks three terminal cirri, which are very likely transverse cirri; short arrow denotes buccal cirrus. Note that the two illustrations are identical except of the oral primordium. e: Ventral view of a representative specimen, 165 µm. f: Small (<1 µm) colourless cortical granules around cirri. g: Dorsal view. h: Infraciliature of ventral side. Arrow marks end of amphisiellid median cirral row. Broken line connects right frontal cirrus and cirrus III/2. Dotted line connects frontal cirri. AZM = adoral zone of membranelles, CV = contractile vacuole, MA = macronuclear nodule, MI = micronucleus, OP = oral primordium, RMR = right marginal row, TC = transverse cirri. Page 264. 1
Diagnosis see U. binucleatus section.
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Fig. 51a–d Uroleptoides binucleatus multicirratus (type population, from Foissner et al. 2002. a, from life; b–d, protargol impregnation). a: Ventral view of a representative specimen, 235 µm. This subspecies is usually longer than 200 µm and very flexible. It differs from the nominal subspecies mainly by the tail-like posterior body portion and the higher number (32 vs. 25 on average) of adoral membranelles, a rather constant and therefore important feature. b: Oral portion of the specimen shown in Fig. 51e at higher magnification (details on fibre system see text). Frontal cirri are connected by dotted line. Arrow marks buccal cirrus. c: Scheme of adoral zone, whose proximal portion is widened spoon-like. d: The bases of the cirri and dorsal bristles are surrounded by conspicuous granules up to 3 µm across. Most globules contain a darkly impregnated, tiny granule. ACR = amphisiellid median cirral row, AZM = adoral zone of membranelles, CV = contractile vacuole, DB = dorsal bristle, E = endoral, F = fibre, FC = right frontal cirrus (cirrus III/3), FL = fibre loop, FS = frontal scutum, LMR = left marginal row, P = paroral, PF = pharyngeal fibres, RMR = right marginal row, III/2 = cirrus behind right frontal cirrus. Page 267.
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Nomenclature: For derivation of the name binucleatus, see Uroleptoides binucleatus. The species-group name multicirratus (having many cirri) is a composite of the Latin quantifier mult·us (many), the thematic vowel ·i-, and the Latin adjective cirrat·us, -a, -um ([m; f; n]; having cirri) and refers to the high number of cirri and adoral membranelles characterising this subspecies (Foissner et al. 2002). The present subspecies was described when the species (U. binucleatus) was classified in Amphisiella. Thus, a formal transfer from Amphisiella to Uroleptoides is required. Remarks: Foissner et al. (2002) found, during a detailed study of Namibian soils, two populations, which were similar to U. binucleatus. The Namibian site 5 population (type) differed more distinctly from Hemberger’s population (= type of U. binucleatus) than the population studied by Berger & Foissner (1989). Thus, we classified the Namibian populations as subspecies, differing from the German populations (Hemberger 1985, Berger & Foissner 1989) mainly by the non-overlapping numbers of adoral membranelles and cirri in the amphisiellid median cirral row. Considering the tailed body, even species status would be appropriate. However, this group of hypotrichs has a profound variability, and thus we classified the Namibian population as a subspecies of U. binucleatus. This is emphasised by the population from Namibian site 41, whose specimens, which are well preserved, are even larger than those from site 5 (Table 22), but the cortical granules are inconspicuous, like those of U. binucleatus binucleatus, that is, pale (with a minute compact centre, however) and only 0.4–0.8 µm across. The Namibian site 5 population of U. binucleatus multicirratus is reminiscent of U. multinucleatus, differing from that species mainly by having only two macronuclear nodules. The present subspecies occurred together with U. magnigranulosus at both Namibian sites. Although the most deviating individuals of both species were difficult to separate in life, most “normal” specimens could be easily distinguished, especially in the protargol slides, by the different length of the amphisiellid median cirral row and the tail-like, respectively, comparatively widely rounded posterior body end (Table 22). Indeed, most U. magnigranulosus specimens were highly similar to those figured in the original description (Foissner 1988). The oral fibre system is very similar in both species. Morphology: Body size 180–300 × 35–65 µm in life, usually about 230 × 50 µm; length:width ratio 4.5–5.5:1 in life, in protargol preparations stouter, that is, about 4:1 on average because very soft and thus more or less inflated due to the preparation procedures (Fig. 51a, e, f; Table 22). Body outline elegantly pisciform with tail-like posterior body portion, a conspicuous feature preserved even in most protargol-impregnated cells, which are lanceolate or clavate. Body dorsoventrally only slightly flattened, very flexible, but acontractile; invariably rather distinctly twisted about main body axis, marginal rows thus never recognisable in full length if specimens are viewed ventrally. Macronuclear nodules usually in middle third of cell, ellipsoidal (2.2:1) to elongate ellipsoidal (3:1), on average 2.6:1; chromatin
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Fig. 51g–i Uroleptoides binucleatus multicirratus (Namibian site 41 population, from Foissner et al. 2002. From life). g, h: A representative specimen, 290 µm. The buccal lip (dotted) covers most of the buccal cavity and is itself covered by the closely spaced paroral cilia; together, they form a highly characteristic, crescentic pattern, impressively shown in the SEM-micrographs of U. multinucleatus (Fig. 49a, b). i: Most specimens are distinctly twisted about main body axis. ACR = amphisiellid median cirral row, BL = buccal lip, LMR = left marginal row, P = paroral. Page 267.
bodies numerous and minute. Micronuclei near or attached to macronuclear nodules, ellipsoidal to broadly ellipsoidal, conspicuous because about 6–7 × 4–5 µm in life; in some specimens inflated to up to 10 µmsized globules (Fig. 51e, f). Contractile vacuole slightly ahead of mid-body at left cell margin, likely with collecting canals during diastole. Cortical granules as described in U. multinucleatus, that is, found only around bases of cirri and dorsal bristles, colourless and globular, occasionally impregnate with protargol showing a minute, heavily argyrophilic inclusion (Fig. 51d); granules around cirri 0.8–1.0 µm across, clusters around dorsal bristles composed of small (about 0.5–1.0 µm) and large (up to 3 µm) granules. Cytoplasm colourless, contains many lipid droplets up to 10 µm across, especially in tail region, which is thus dark at low (<100) magnification. Movement conspicuous because slow and serpentine, showing great flexibility between soil particles; however, can also swim rather rapidly by rotation about main body axis. Adoral zone occupies only 15–23%, on average 19% of body length, of usual shape and structure, except for proximal half, which is widened spoon-like (Fig.
b Fig. 51e, f Uroleptoides binucleatus multicirratus (type population, from Foissner et al. 2002. Protargol impregnation). Infraciliature of ventral and dorsal side and nuclear apparatus of two different specimens because the holotype (e) is too opaque to see the dorsal kineties clearly. Specimens inflated due to preparation procedures. For details of oral and frontal ciliature see Fig. 51b. ACR = amphisiellid median cirral row, LMR = left marginal row, MI = micronuclei (inflated in holotype), RMR = right marginal row, TC = transverse cirri, 1–3 = dorsal kineties. Page 267.
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51a, b, e); composed of an average of 32 membranelles, bases of largest membranelles 10–12 µm wide in life. Buccal cavity deep and large as in Cyrtohymena (for review, see Berger 1999), bears complicated fibre system (Fig. 51b). The most conspicuous part of this system is associated with the undulating membranes, that is, a thick fibre originates from the anterior end of the endoral and splits into a short right branch extending into the frontal scutum and a long left branch extending backwards to merge into a fibrous plate originating from the rear end of the undulating membranes. This fibre loop is connected to the adoral fibre system via a fibre formed by horizontally extending fibres originating from the adoral membranelles. Further fibres, not shown in Fig. 51b, originate from the paroral and support the ventral and dorsal wall of the buccal cavity. Uroleptoides magnigranulosus has a very similar oral fibre system, although it is less conspicuous due to the smaller body size. Furthermore, the large buccal lip and the up to 20 µm long paroral cilia form a conspicuous, crescentic pattern, as shown in Fig. 51g, h and the SEMmicrographs of Uroleptoides multinucleatus (Fig. 49a, b). Undulating membranes distinctly curved, close together, optically intersecting near mid of buccal cavity. Pharyngeal fibres very prominent in life and protargol preparations, because mixed with the about 40 µm (!) long endoral cilia. Cirral pattern and number of cirri rather variable, as indicated by the high variability coefficients (Fig. 51a, b, e; Table 22). Frontal and transverse cirri about 20 µm, other cirri 12–15 µm long in life and closely spaced within rows, distances between individual cirri increase only in posterior portion of marginal rows. Frontal cirri distinctly enlarged, form concave row, right cirrus close to distal end of adoral zone. Buccal cirrus slightly enlarged, near summit of curve formed by undulating membranes. Usually three slightly enlarged cirri (cirrus III/2 included) left of anterior portion of amphisiellid median cirral row. Amphisiellid median cirral row conspicuous because composed of 67 cirri occupying 82% of body length on average, commences right of distal end of adoral zone and extends obliquely to subterminal left body margin. 2–4 transverse cirri at posterior end of cell, difficult to distinguish from marginal cirri. Marginal rows follow body curvature and thus usually distinctly spiralled, extend to near posterior body end. Dorsal bristles about 4 µm long in life, arranged in three rows leaving broad blank stripe in midline, that is, between rows 2 and 3 (Fig. 51f); rows l and 2 extend near left body margin and are slightly shortened anteriorly and posteriorly; row 3 extends along right body margin, anteriorly more distinctly shortened than rows l and 2. Caudal cirri lacking. Occurrence and ecology: Uroleptoides binucleatus multicirratus is known from two Namibian sites, namely, from the type locality (site 5), where it was rare, and from site 41, where it was rather abundant (Foissner et al. 2002). It is well-adapted to the soil environment by the slender and soft body. The type locality is near the Gariganus Guest Farm (26°30’S, 18°25’E), Namibia, where it was discovered in soil of an Aloe dichotoma forest. Site 41 is in the Spitzkoppe area (escarpment of the central Namib Desert) about 120 km north of the town of Swakopmund; the sample
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was composed of mud and grass (Aponogetum desertorum) roots from deep, dry granitic rock-pools (pH 5.8; Foissner et al. 2002). Uroleptoides binucleatus multicirratus is a voracious predator ingesting bacterial rods, coccal green algae, naked amoebae, heterotrophic flagellates (Polytomella), and medium-sized ciliates, such as Sathrophilus muscorum, Colpoda maupasi, and C. cucullus. The prey is ingested whole and rotates for some minutes in the food vacuoles (Foissner et al. 2002).
Uroleptoides magnigranulosus (Foissner, 1988) comb. nov. (Fig. 52a–l, 53a–e, Table 22) 1988 Amphisiella magnigranulosa nov. spec.1 – Foissner, Stapfia, 17: 115, Abb. 10a–l, 22, Tabelle 6 (Fig. 52a–l; original description; one holotype slide [accession number 1989/21] and one paratype slide [1989/22] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; Aescht 2003, p. 390). 2001 Amphisiella magnigranulosa Foissner, 1988 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 8 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Amphisiella magnigranulosa Foissner, 1988 – Foissner, Agatha & Berger, Denisia, 5: 693, Fig. 388a–e (Fig. 53a–e; detailed description of cortical granulation). 2007 Amphisiella magnigranulosa – Schmidt, J. Euk. Microbiol., 54: 203 (analysis of the nuclear small subunit RNA; GenBank accession number AM412774).
Nomenclature: The species-group name magnigranulosa (having large granules) is a composite of magn·us, -a, -um (Latin adjective [m; f; n]; large, strong), the thematic vowel ·i-, and granulos·us, -a, -um (Latin adjective; having many grains) and refers to the large cortical granules (Foissner 1988). Amphisiella is feminine, Uroleptoides is masculine. Thus, the ending of the species-group name has to be changed from magnigranulos·a to magnigranulos·us (ICZN 1999). Remarks: The amphisiellid median cirral row of the present species terminates at about 50% of body length. Thus, the pragmatic assignment to Lamtostyla (row length <50%) or Uroleptoides (>50%) is difficult. I preliminarily transfer it to Uroleptoides because almost all species so far assigned to Lamtostyla in the present book lack cortical granules, whereas such organelles are characteristic for the present species and several other Uroleptoides species. For a more general discussion of the separation of Amphisiella, Uroleptoides, and Lamtostyla, see genus section. Uroleptoides magnigranulosus is a common, binucleate species, which differs from other species with two macronuclear nodules by the presence of cortical granules (vs. lacking in Lamtostyla procera) or the length of the amphisiellid median cirral row (ending near mid-body vs. distinctly behind in Uroleptoides binucleatus). Morphology: Foissner (1988) studied six populations from various geographic regions. Partially, they differ distinctly in body size, body shape, and the cortical 1 Foissner (1988) provided the following diagnosis: In vivo etwa 120–200 × 30–60 µm große Amphisiella mit 1–2 µm durchmessenden, farblosen, subpelliculären Granula entlang der Infraciliatur. Durchschnittlich 23 adorale Membranellen und 3 Dorsalkineten. 2 Makronucleus-Teile.
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granules. By contrast, the cirral pattern of the various populations is rather similar. However, the slides were only of mediocre quality because the specimens were difficult to fix. The type population is described in detail first. Subsequently, deviating live data from the other populations are mentioned. By contrast, the diagnosis (see footnote) contains data (e.g., body size) from most/all populations studied by Foissner (1988). At the end of the morphology section is a paragraph about the cortical granulation studied in detail by Foissner et al. (2002). Type population (Fig. 52a–f): Body size about 110–140 × 40 µm. Body outline moderately wide elliptical to slightly sigmoidal, in protargol preparations always inflated because difficult to fix. Anterior and posterior body portion more or less distinctly narrowed. Body dorsoventrally flattened about 2:1, very flexible, and slightly contractile under cover glass pressure. Macronuclear nodules long ellipsoidal, that is, length:width ratio 2–3:1, arranged slightly left of midline in middle third of cell; contain many moderately-sized nucleoli. Usually three micronuclei about 4 × 3 µm in size. Contractile vacuole slightly ahead of mid-body near left cell margin, during diastole with two long collecting canals. Cortical granules colourless, conspicuous in life because about 1.5 µm in diameter, slightly vaulting pellicle, and not impregnating with protargol; arranged along infraciliature and in buccal field, where they are abundant and form a laminar structure. Cytoplasm colourless, usually with some large, globular to clod-shaped, greasy-shining inclusions and several food vacuoles. Movement moderately rapid, can attach closely to soil particles. Adoral zone occupies only about one fourth of body length, composed of 23 membranelles on average (Table 22). Bases of largest membranelles 7–10 µm wide. Buccal field deep and narrow, anterior margin hooks leftwards. Undulating membranes distinctly curved, of about same length, optically intersecting in rear portion. Pharyngeal fibres distinct in live and protargol preparations (Fig. 52a, b). Cirral pattern and number of cirri of usual variability (Fig. 52b, Table 22). Frontal cirri distinctly enlarged, arranged in bow-shaped pseudorow immediately behind distal portion of adoral zone. Buccal cirrus about at optical intersection of undulating membranes, that is, distinctly behind anterior end of paroral. Usually five cirri (cirrus III/2, that is, cirrus behind right frontal cirrus included) left of anterior portion of amphisiellid median cirral row; interestingly, they show a midventral pattern indicating that they originate from three anlagen (however, morphogenetic data are
b Fig. 52a–l Uroleptoides magnigranulosus (from Foissner 1988. a–f, type population from Kenya; g, k, l, Australian population; h–j, Yugoslavian population. a, d–l, from life; b, c, protargol impregnation). a: Ventral view of a representative specimen, 136 µm. b, c: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 110 µm. Short arrow denotes distal end of adoral zone, long arrow marks break in amphisiellid median cirral row. Frontal cirri connected by dotted line, cirri left of anterior end of amphisiellid median cirral row circled. Broken lines connect cirri which originate from anlagen II and III (corresponding transverse cirri not included). d, e: Shape variant in dorsal and left lateral view. f–h: Cortical granules of various populations. i–l: Shape variants in ventral and dorsal (l) view. ACR = anterior end of amphisiellid median cirral row, CG = cortical granules, CV = contractile vacuole, E = endoral, LMR = left marginal row, MA = macronuclear nodule, MI = micronucleus, PF = pharyngeal fibres, RMR = right marginal row, TC = transverse cirri, 1–3 = dorsal kineties. Page 273.
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Fig. 53a, b Uroleptoides magnigranulosus (from Foissner et al. 2002. Interference contrast. a, Australian specimen; b, Namibian site 63 specimen). Cells are heavily squeezed to show granules in large portions of cell. Fig. 53a is a ventrolateral view, (b) shows the dorsal anterior portion. Arrows in (b) mark granule rows; arrowheads in (a) denote individual granule clusters and white arrows mark small granules (large ones out of focus) in right marginal row. Note that the cortical granulation is very similar in Australian and Namibian specimens. DK = dorsal kinety, MR = right marginal row. Page 273.
needed for correct interpretation)1. Amphisiellid median cirral row likely composed of two portions because there is always a distinct break in the middle part; commences near distal end of adoral zone, terminates at 47% of body length on average (Table 22, see remarks for generic assignment). Cirri of amphisiellid median cirral row and marginal cirri about 10 µm long. Certainly no postperistomial ventral cirrus. 1
Two out of 12 specimens of the type population had six transverse cirri (Fig. 52b), most had five, and one specimen each had four, respectively, three transverse cirri. Accordingly, the specimens with six true transverse cirri (not five transverse cirri plus one pretransverse ventral cirrus) must have had seven frontal-ventral-transverse cirral anlagen.
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Distinct pretransverse ventral cirri lacking; however, it cannot be excluded that the rightmost cirrus of the specimen shown in Fig. 52b is the pretransverse ventral cirrus VI/2. Transverse cirri arranged in hook-shape, almost terminal, about 15 µm long, and therefore projecting beyond rear body end (Fig. 52a). Right marginal row commences about at same level as amphisiellid median cirral row, terminates about at level of transverse cirri. Left marginal row begins near proximal end of adoral zone, terminates at rear end of cell. Dorsal bristles about 3 µm long, arranged in three (rarely two) kineties; bristle rows 2 and 3 slightly shortened anteriorly (Fig. 52c). Caudal cirri lacking. Population from Australian sample 16 (Foissner 1988): Body size about 150 × 35 µm in life. Body outline slightly bow-shaped and somewhat twisted about main body axis. In about 50% of specimens the cortical granules along the dorsal kineties, but not that along the cirral rows, impregnated. Buccal field narrow and deep, however, anteriorly only slightly curved leftwards. Micronuclei very large, that is, about 5–7 × 5 µm in life. Population from Australian sample 19 (Foissner 1988): Size about 170 × 60 µm in life. Body shape very similar to type population, but posterior portion usually slightly more narrowed (Fig. 52k, l). Cortical granules right of cirri about 2 µm across, left of cirri about 3 µm (Fig. 52g), do not stain with methyl green-pyronin. Buccal field wide and deep, anteriorly not curved like a hook. Population from Ester Island (Foissner 1988): Very similar to type population. Population from Yugoslavia (Foissner 1988): Body size about 200 × 50 µm in life. Body outline usually slightly sigmoidal, posteriorly distinctly narrowed (Fig. 52i, j). Two types of colourless, globular cortical granules along infraciliature, namely, one type only 0.2 µm across and the other type 1–2 µm in diameter (Fig. 52h); on the dorsal side they are slightly argyrophil, on the ventral side they are not. Buccal field narrow, deep, and anteriorly curved leftwards. Dorsal cilia about 4 µm long, lively motile. Population from Salzburg, Austria (Foissner 1988): Very similar to Yugoslavian population, especially as concerns cortical granulation. Foissner et al. (2002) provided a detailed description of the cortical granulation, which is overlooked at superficial observation because the granules are colourless and rather hyaline (Fig. 53a–e). The granules are restricted to the ciliature, that is, found only around the cirri, the dorsal bristles, and in the wall of the buccal cavity. The clusters around the cirri are usually less distinct than those around the dorsal bristles. The buccal granules are conspicuous and of intermediate size (about 1 µm). Each granule cluster is composed of up to 15 minute (<1 µm) and 1–3 large (up to 3 µm, usually 1.5–2.5 µm) granules, which appear ring-shaped, indicating that they consist of two components of different refractivity. Although there is some variability in size and number of small and large granules, the granule clusters have a very similar pattern and appearance in populations from all main biogeographic regions. Thus, they are a highly constant and an important feature. The nature of the cortical granules of U. magnigranulosus is not known. On prolonged squeezing, they disap-
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Fig. 53c–e Uroleptoides magnigranulosus (from Foissner et al. 2002. Interference contrast. c, Australian specimen; d, e, Namibian site 51 specimen). Ventral views showing arrangement of cortical granules within cirral and bristle rows and in buccal cavity. Note that the granules, although being rather large, are not very conspicuous. ACR = amphisiellid median cirral row, AZM = adoral zone of membranelles, B = buccal cavity, DK = dorsal kinety, FV = food vacuole, MA = macronuclear nodule, PF = pharyngeal fibres, RMR = right marginal row, TC = transverse cirri. Page 273.
pear, that is, are extruded, fading away without forming a coat around the cell. They do not, or only faintly, stain with methyl green-pyronin, but occasionally impregnate lightly with protargol. Molecular data: The nuclear small subunit rRNA gene sequence is 1729 base pairs long (Schmidt et al. 2007, GenBank accession number AM412774). In the
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Bayesian analysis U. magnigranulosus is the sister to the urostyloid cluster Pseudokeronopsis rubra to Holosticha multistilata (Fig. 2 in Schmidt et al. 2007). With the Maximum-likelihood method no meaningful position could be estimated (Fig. 3 in Schmidt et al. 2007). Indeed, the cirral pattern of the specimen shown in Fig. 52b is somewhat reminiscent of a urostyloid because of the paired cirri left of the anterior portion of the amphisiellid median cirral row (for review of urostyloids see Berger 2006). However, without detailed ontogenetic data a serious discussion is not possible. Occurrence and ecology: Uroleptoides magnigranulosus is a rather common ciliate very likely confined to terrestrial habitats (Foissner 1998, p. 199). So far it is known from more than 100 soil samples world-wide (Foissner et al. 2002). Type locality is a coffee plantation between Riuru and Kalimoni near Nairobi, Kenya, where Foissner (1988) discovered it in the upper (0–5 cm) soil layer, which was russet, had a pH of 6.2, and contained much litter. Foissner (1988) found it also in terrestrial habitats from Salzburg (Austria), from the Ester Island (near airport), from former Yugoslavia (near sea coast), and from two sites in Australia, namely, in a soil (pH 4.3) with litter from near Lake Dobson (sample 16; Mt. Field National Park, Tasmania) and in bark (pH 5.6) grown with mosses and lichens (sample 19) from a secondary pine forest between Cairns and Innisfall (Blatterer & Foissner 1988, p. 4, 6). Recently, we found it in three of 12 Austrian forest preservation sites (Foissner et al. 2005, p. 624). In Namibia it occurred in about 23% of the soil samples investigated (Foissner et al. 2002, p. 58). Feeds on ciliates (Colpoda sp., Drepanomonas sp.); occasionally fungal spores and coccal green algae can be found in the food vacuoles (Foissner 1988). Biomass of 106 specimens about 90 mg (Foissner 1998, p. 199).
Uroleptoides terricola (Gellért, 1956) comb. nov. (Fig. 54a–h, Table 22) 1956 Amphisiella terricola n. sp. – Gellért, Acta biol. hung., 6: 95, Abb. 10 (Fig. 54a; original description; no formal diagnosis provided and likely no type material available). 1956 Amphisiella terricola Gellért – Gellért, Acta biol. hung., 6: 346 (supplementary data). 1972 Amphisiella terricola Gellért, 1955 – Borror, J. Protozool., 19: 9 (revision of hypotrichs). 1974 Amphisiella terricola Gellért – Stiller, Fauna Hung., 115: 96, Fig. 59 (redrawing of Fig. 54a; revision of hypotrichs). 1982 Amphisiella terricola Gellért, 1955 – Hemberger, Dissertation, p. 23, Abb. 3a, b (Fig. 54f–h; redescription after protargol impregnation; revision of hypotrichs; voucher slides are likely deposited in the Institut für landwirtschaftliche Zoologie und Bienenkunde, University of Bonn, Germany). 1984 Amphisiella terricola Gellért, 1955 – Foissner, Stapfia, 12: 114, Abb. 59a–d, Tabelle 28 (Fig. 54b–e; authoritative redescription; two voucher slides [accession numbers 1984/88, 1984/89] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; see nomenclature).
Nomenclature: No derivation of the species-group name terricola is given in the original description. It is a composite of the Latin noun terr·a (soil), the thematic
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vowel ·i-, and the Latin verb colere (to live in) and means living in soil. Usually, species-group names ending with -cola are considered as appositive substantives and are thus not changed when transferred to a genus of different gender (Werner 1972, p. 138). Foissner (1984, p. 8) wrote that he has deposited at least one slide of each redescribed species in the museum in Linz. According to Aescht (2003, p. 397), these are neotype slides, perhaps because the slides are labelled as neotypes. Anyhow, according to the ICZN (1999, Article 75.2) such neotype designations are invalid. Some authors cite the original description with the year 1955. I have a copy of the title page of volume 6 of the journal, showing the year 1956. However, it cannot be excluded that issues 1 and 2, which contain the original description, were already published in 1955. Remarks: For a foundation of the transfer from Amphisiella to Uroleptoides, see genus section. The original description of Uroleptoides terricola is rather detailed, although it is based only on fixed material stained with the opalblue method. The species was accepted by Borror (1972) and Stiller (1974) and redescribed after protargol impregnation by Hemberger (1982). Foissner (1984) provided data from live and protargol preparations. Foissner’s study largely confirmed the original description and the redescription, although the specimens investigated by Hemberger are almost twice as long as those of Gellért (1956a) and Foissner (1984). Interestingly, Hemberger’s specimens have fewer adoral membranelles and cirri in the amphisiellid median cirral row and marginal rows than Foissner’s population, indicating that Hemberger’s specimens were strongly inflated or his measurements overestimate the size. In spite of these differences, the conspecificity of the populations is beyond reasonable doubt. The redescription by Foissner (1984) consists of three paragraphs. Due to a printers error, the last paragraph, which treats the cirral pattern, is printed ahead of the heading “Genus Amphisiella ...”.
Fig. 54a–e Uroleptoides terricola (a, from Gellért 1956a; b–e, from Foissner 1984. a, opalblue staining after Bresslau; b, c, from life; d, e, protargol impregnation). a: Infraciliature of ventral side, nuclear apparatus, and contractile vacuole; body length according to text about 100 µm, according to figure legend around 160 µm. b: Ventral view of a representative specimen, 115 µm. c: Right lateral view. d, e: Infraciliature of dorsal and ventral side showing dorsal kineties, cirral pattern, and nuclear apparatus, 88 µm. Arrow in (d) marks a break in dorsal kinety 3; arrow in (e) denotes cirrus III/2. Arrowhead marks additional cirral row (remnant of previous generation?), probably a specific feature of this specimen like the break in dorsal kinety 3. Note that the amphisiellid median cirral row is composed of 26 cirri in this specimen; by contrast, the minimum value in the morphometry (Table 22) is 30, indicating that the seven cirri of the additional row are included. Broken lines connect cirri, which very likely originate from same anlage, dotted line connects frontal cirri. ACR = amphisiellid median cirral row, AZM = adoral zone of membranelles, CV = contractile vacuole, E = endoral, MA = macronuclear nodule with replication band, MI = micronucleus, P = paroral, RMR = right marginal row, TC = transverse cirri, 1–3 = dorsal kineties. Page 279.
d
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Morphology: The authoritative redescription by Foissner (1984) is provided first. In the last paragraph some additional and deviating data from the other populations are described. Body size in life about 100–130 × 40–60 µm. Body outline rather invariable orthogonal, converging anteriorly, right margin slightly concave, left slightly convex, broadly rounded anteriorly and posteriorly. Body about 2:1 dorsoventrally flattened. Macronuclear nodules in life about 20 × 13 µm, arranged behind proximal end of adoral zone, that is, slightly left (Foissner wrote, likely par lapsus, right) of midline, with many, very small chromatin bodies. Several micronuclei closely attached to macronuclear nodules, do not impregnate with protargol, in life about 5 × 4 µm. Contractile vacuole, as is usual, near left cell margin about in mid-body, without collecting canals during diastole. Pellicle colourless, conspicuous cortical granules lacking. Cytoplasm often packed with globular and clod-shaped, colourless, 2–15 µm-sized inclusions. Food vacuoles about 20 µm across. Movement slow, without peculiarities. Adoral zone of membranelles slightly contractile, thus somewhat shortened in protargol preparations (this feature is also recognisable in the fixed specimens of the other populations; Fig. 54a, b, e, f); zone occupies about 30% of body length in protargol-impregnated specimens (Table 22), composed of an average of 30 membranelles. Bases of largest membranelles about 11 µm wide. Buccal lip distinctly curved leftwards anteriorly. Undulating membranes of about same length, distinctly (almost semicircular) curved and optically intersecting in protargol preparations. Pharyngeal fibres conspicuous in protargol-impregnated specimens (Fig. 54e). Most cirri about 12 µm long in life. Frontal cirri slightly enlarged, in life about 15 µm long. Buccal cirrus about at level of cirrus III/2. Behind right frontal cirrus 5–7 zigzagging cirri (cirrus III/2 included), indicating that two cirri each are formed per anlage. Amphisiellid median cirral row commences near distal end of adoral zone, extends to midline of cell and terminates slightly ahead of transverse cirri (at 83% of body length in specimen illustrated; Fig. 54e). 6–7 transverse cirri arranged in subterminal, curved pseudorow; end of cirri extends to rear cell margin. Specimen illustrated with an additional row composed of seven cirri (Fig. 54e). Right marginal row commences dorsolaterally, terminates about in midline at rear end of cell; left row commences slightly ahead of proximal end of adoral zone; rows distinctly separated posteriorly; bases of rearmost marginal cirri rather small. Dorsal cilia about 5 µm long, arranged in three kineties; kinety 3 interrupted about in mid-body (Fig. 54d); it is unclear whether this is a specific feature of the specimen illustrated or of the species. If ontogenetic data show that this is an oxytrichid kinety fragmentation, then the classification in Uroleptoides, respectively, the amphisiellids is incorrect. Caudal cirri lacking. Additional observations from other populations (unless otherwise indicated, the data are from Gellért 1956a; for Hemberger’s data see mainly Table 22): body length about 100 µm, according to Gellért (1956b) 110–120 µm; length:width ratio 3:1; right cell margin concave, left convex, both ends rounded (Fig. 54a); one micro-
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Fig. 54f–h Uroleptoides terricola (from Hemberger 1982. Protargol impregnation). f, g: Infraciliature of ventral side and nuclear apparatus of an early divider, 213 µm. h: Infraciliature of ventral side of a middle divider. ACR = amphisiellid median cirral row, OP = oral primordium, I, VI = frontal-ventral-transverse cirri. Page 279.
nucleus attached to each macronuclear nodule; contractile vacuole empties every 16–18 seconds, respectively, every 22–25 seconds (Gellért 1956b); cytoplasm bright in anterior portion, somewhat darker in posterior portion; movement slow, often moving back; three frontal cirri; left of anterior portion of amphisiellid median cirral row two oblique rows with three cirri each (buccal cirrus and cirrus III/2 obviously included), according to Hemberger (1982), 8–10 cirri on frontal field (comprising frontal cirri, buccal cirrus, and cirri left of anterior portion of amphisiellid median cirral row); 5–8 transverse cirri, which project slightly beyond rear body end; transverse cirri about 20 µm long and marginal cirri about 15 µm (Hemberger 1982); buccal lip slightly curved; adoral zone occupies about 25% of body length, composed of 22 membranelles; paroral composed of six thick cilia (likely a misinterpretation of the preparation); three bipolar dorsal kineties; dorsal bristles about 5 µm long (Hemberger 1982).
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Cell division (Fig. 54f–h): Hemberger (1982) found two dividers. Stomatogenesis commences with the formation of an oral primordium near the left transverse cirrus (Fig. 54f). Later, six frontal-ventral-transverse cirri anlagen (I–VI) are formed. Furthermore, two anlagen – one for the proter and one for the opisthe – occur within the parental amphisiellid median cirral row (Fig. 54h). Later stages have to be studied to show whether the amphisiellid median cirral row is formed, as is usual, by two anlagen or only by one anlage. If it is formed from a single anlage only, Uroleptoides terricola has to be (very likely) removed from the amphisiellids. Occurrence and ecology: Uroleptoides terricola is likely confined to terrestrial habitats. It is recorded from all biogeographic regions, except for the Antarctica (Foissner 1998, p. 199). Type locality of U. terricola is the south-western side of the Magoska hill (Tokaj-Eperjes mountains) north-east of the village of Boldogkõváralja, administrative district of Abauj-Torna, Hungary, where Gellért (1956a) discovered it in the humus layer formed underneath the lichen Parmelia saxatilis. Gellért (1956b) found it in the same area in the humus layer formed under moss grown on a rock. Hemberger (1982) isolated his population from an infusion of faeces of Deroceras reticulatum, a common land snail (Kerney et al. 1983, p. 195) collected from the mouth of the river Sieg in Germany. Foissner (1984) recorded U. terricola from a xerothermic site without trees with shallow, brown, alluvial soil near the village of Vogelsang/Grafenwörth, Lower Austria (details see site 4 in Foissner et al. 1985, p. 87). Records not substantiated by morphological data and/or illustrations: soil of spruce forest near the village of Aigen-Schlägl, Upper Austria (Petz et al. 1988, p. 81); several forest stands in eastern Austria (Foissner et al. 2005, p. 624); soil of spruce forest (Ulm area, Ochsenhausen area) in southern Germany (Funke 1986, p. 72); salinised soils from the Hortobágy National Park, Hungary (Szabó 1999, p. 249); with moderate abundance in a chernozem soil from the centre of the Great Hungarian Plain, about 15 km south of the main road no. 33 between the villages of Hajdúszoboszló and Nagyhegyes (Szabó 2000, p. 14); litter from Fagus sylvatica and Quercus sp. in Slovakia (Tirjaková & Bartosová 2004, p. 16); microbiotic crusts containing cyanobacteria and bryophytes from desert soils of the Grand Canyon in northern Arizona, USA (Bamforth 2004, p. 417); blackwater inundation primary(?) rain forest from a small island in the region, where the yellow Rio Solimões units with the black Rio Negro, Janauari region, about 2 km east of Manaus, Brazil (Foissner 1997, p. 320). Uroleptoides terricola feeds on small ciliates, for example, Colpoda fastigata (Foissner 1984, Gellért 1956b). Biomass of 106 specimens about 100 mg (Foissner 1987, p. 121).
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Uroleptoides polycirratus (Berger & Foissner, 1989) comb. nov. (Fig. 55a–d, Table 22) 1989 Amphisiella polycirrata nov. spec.1 – Berger & Foissner, Bull. Br. Mus. nat. Hist., 55: 32, Fig. 31, 32, Table 8 (Fig. 55c, d; original description; the holotype slide [accession number 1988:2:1:1] and the paratype slide [1988:2:1:2] are deposited in the British Museum of Natural History, England). 2001 Amphisiella polycirrata Berger and Foissner, 1989 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 8 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Amphisiella polycirrata Berger & Foissner, 1989 – Foissner, Agatha & Berger, Denisia, 5: 691, Fig. 153a–d (Fig. 55a–d; description of live specimens).
Nomenclature: No derivation of the name is given in the original description. The species-group name polycirratus is a composite of the Greek quantifier polys (Greek; many) and cirrat·us, -a, -um (Latin adjective [m; f; n]; having cirri) and refers to the many buccal cirri. Amphisiella is feminine, Uroleptoides is masculine. Thus, the ending of the species-group name has to be changed from polycirrat·a to polycirrat·us (ICZN 1999). Remarks: For a foundation of the transfer from Amphisiella to Uroleptoides see genus section. The present species was originally described only after protargol preparations (Berger & Foissner 1989). It is a conspicuous hypotrich differing from Uroleptoides terricola (Fig. 54a–h) by body size (>150 µm vs. <150 µm), number of buccal cirri (4–8 vs. 1), and length of amphisiellid median cirral row (abutting on transverse cirri vs. more or less distinctly shortened). As these features are stable, Uroleptoides polycirratus is easily identified. Uroleptoides polycirratus is somewhat reminiscent of Bistichella namibiensis (Fig. 114a–g; see there for details). The original description lacks an illustration of a living specimen because “in vivo this species looks so similar to Uroleptoides terricola”, that we considered it unnecessary to draw the living aspect (Berger & Foissner 1989). This is confirmed by new observations (Foissner et al 2002). However, to emphasise several features, especially the lack of the cortical granules in all six populations studied, we provided a description and a figure of the live aspect of the specimens from a Namibian site (Foissner et al. 2002). Moreover, in the original description we were uncertain about the presence/absence of caudal cirri. New data show that it lacks such cirri (see morphology). Morphology: The live aspect is mainly from Foissner et al. (2002), whereas the data on the cirral pattern are largely from the type population (Berger & Foissner 1989). Body size about 170–200 × 40–60 µm in life. Body outline elongate elliptical (body length:width ratio about 4:1) with both ends broadly rounded in ordinary 1
Berger & Foissner (1989) provided the following diagnosis: After protargol impregnation about 150 × 46 µm, long ellipsoid. 2 macronuclear segments, 3 dorsal kineties. 6 buccal cirri, 4 transverse cirri, and 37 adoral membranelles on average. About 33 cirri in the right frontoventral row which terminates at the transverse cirri.
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Fig. 55a–d Uroleptoides polycirratus (a, b, from Foissner et al. 2002; c, d, from Berger & Foissner 1989. a, b, from life; c, d, protargol impregnation). a: Ventral view of a representative specimen from Namibia, 180 µm. b: Outline of a specimen with food inclusions from the Republic of South Africa. Such fat specimens occur also in Namibia. c, d: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen from type population, 150 µm. Long arrow in (c) marks cirrus III/2, short arrow denotes rearmost buccal cirrus. The two cirri left of the anterior portion of the amphisiellid median cirral row are circled. Arrow in (d) marks rearmost cirri of left marginal row (note that U. polycirratus lacks caudal cirri). ACR = amphisiellid median cirral row, AZM = distal end of adoral zone of membranelles, FC = right frontal cirrus, MA = macronuclear nodule, MI = micronucleus, P = paroral, RMR = right marginal row, TC = transverse cirri, 1–3 = dorsal kineties. Page 285.
specimens (Fig. 55a), bursiform when packed with large food inclusions (Fig. 55b). Body dorsoventrally flattened up to 3:1. Two ellipsoidal macronuclear nodules slightly left of midline; individual nodules about 30 × 15 µm in life and with many small chromatin bodies. Several conspicuously large (6 µm across) micronuclei. Contractile vacuole about in mid-body, with inconspicuous collecting canals during
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diastole. Cortex colourless, very flexible, without specific granules. Cytoplasm hyaline because free of refractive crystals, but frequently packed with lipid droplets up to 15 µm across postorally, making cells dark at low (≤ ×100) magnification. Moves rather rapidly on microscope slide and between organic debris, showing great flexibility. Adoral zone occupies about 35% of body length (Table 22), composed of an average of 37 membranelles in type population, while 45–50 membranelles each in Brazilian, South African, and Namibian specimens; however, the type population was weak and therefore such differences are to be expected. Membranelles of proximal half rather wide. Buccal cavity, although rather narrow, conspicuous because deep and extending to adoral zone of membranelles anteriorly. Buccal lip inconspicuous. Undulating membranes slightly bent, of about same length, optically intersecting about in mid-length (Fig. 55a, c). Pharyngeal fibres extend posteriorly. Cirral pattern of ordinary variability (Fig. 55c, Table 22); cirri thick, but not exceedingly long. Frontal cirri distinctly enlarged, arranged in oblique pseudorow with right cirrus at distal end of adoral zone. Buccal cirri extend right of paroral. One cirrus (= cirrus III/2) behind right frontal cirrus, and two cirri left of anterior portion of amphisiellid median cirral row, which is composed of an average of 32 cirri and extends from near distal end of adoral zone to transverse cirri; anteriormost three cirri of amphisiellid median cirral row slightly enlarged (possibly these are the cirri from anlage VI). Usually four transverse cirri subterminally arranged in oblique pseudorow, in life about 23 µm long, hardly projecting beyond rear body end. Marginal cirri about 17 µm long. Right marginal row commences dorsolaterally about at level of cirrus III/2, ends subterminally. Left row begins near proximal end of adoral zone, J-shaped and therefore ending in midline of cell at rear body end. In the original description we could not clarify whether the three cirri shown in Fig. 55d are caudal cirri or the rearmost cirri of the left marginal row (Berger & Foissner 1989). Later, Foissner (pers. communication) found that these are the rearmost cirri of the left marginal row, that is, the present species lacks caudal cirri. Dorsal cilia about 3 µm long, arranged in three more or less bipolar kineties (Fig. 55d). Occurrence and ecology: Uroleptoides polycirratus is very likely confined to terrestrial habitats (Foissner 1998, p. 199; Foissner et al. 2002, p. 50). Type locality is Garajan Kap, Madeira, Portugal, where Berger & Foissner (1989) found it in a slightly reddish-brown soil grown with Opuntia ficusindica and tufts of grass (0–5 cm; pH 4.8; about 150 m above sea-level). Later, it was found in Brazil, Republic of South Africa, Kenya, and Namibia, indicating that it is a cosmopolitan (Foissner et al. 2002). Uroleptoides polycirratus feeds on small (Cyrtolophosis mucicola) and mediumsized ciliates (Colpoda), and likely also on heterotrophic flagellates (Berger & Foissner 1989, Foissner et al. 2002). Biomass of 106 specimens about 120 mg (Foissner 1998, p. 199).
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Hemiamphisiella Foissner, 1988 1988 Hemiamphisiella nov. gen.1 – Foissner, Stapfia, 17: 121 (original description). Type species (by original designation): Hemiamphisiella terricola Foissner, 1988. 1989 Hemiamphisiella – Foissner & Blatterer, J. Protozool., 37: 9A, Abstract 53 (brief characterisation of Hemiamphisiella). 1994 Hemiamphisiella Foissner, 1988 2 – Eigner & Foissner, J. Euk. Microbiol., 41: 260 (redefinition). 1996 Hemiamphisiella Foissner, 1988 3 – Petz & Foissner, Acta Protozool., 35: 277 (redefinition). 2001 Hemiamphisiella Foissner 1988 – Aescht, Denisia, 1: 80 (catalogue of generic names of ciliates). 2001 Hemiamphisiella Foissner, 1988 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 30 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Hemiamphisiella Foissner, 1988 – Lynn & Small, Phylum Ciliophora, p. 452 (guide to ciliate genera).
Nomenclature: Hemiamphisiella is a composite of the Greek prefix hemi+ (half, partly) and the genus-group name Amphisiella (see there for derivation) indicating a similarity in parts with Amphisiella (Foissner 1988). Feminine gender because ending with the Latin suffix -ella (ICZN 1999, Article 30.1.3). Characterisation (A = supposed apomorphy): Amphisiellidae with elongate body, usually distinctly narrowed posteriorly and slightly twisted about main body axis. Adoral zone continuous. Three enlarged frontal cirri. One buccal cirrus. Only one cirrus (= cirrus III/2) left of anterior portion of amphisiellid median cirral row. Postperistomial cirrus4 present, originates from anlage IV. Amphisiellid median cirral row originates from anlage IV (forms middle portion of row), V (rear portion), and VI (anterior portion). Transverse cirri lacking (A?). One more or less long cirral row (= right ventral row) between rear portion of marginal rows (A). One left and one right marginal row. Three or four dorsal kineties. Caudal cirri present. Additional characters: Body flexible, about 2:1 flattened dorsoventrally; contractile vacuole at left cell margin, at mid-body or slightly ahead of it; dorsal bristles short, that is, less than 5 µm.
1
Foissner (1988) provided the following diagnosis: Amphisiellidae (?) mit 1 Cirrus links der Ventralreihe im Frontalfeld. Hinter dem Peristom 1 isolierter Cirrus zwischen der Ventralreihe und der linken Marginalreihe. Caudalcirren vorhanden. Meist eine zweite, stark verkürzte Ventralreihe in der posterioren Körperhälfte. 2 Eigner & Foissner (1994) provided the following improved diagnosis: The amphisiellid median cirral row originates from three rightmost anlagen. Usually one postperistomial cirrus develops from the third anlage from right. One cirrus left of amphisiellid median cirral row. Transverse cirri longitudinally arranged, originate from single anlage. Caudal cirri present. 3 Petz & Foissner (1996) provided the following improved diagnosis: The oral primordium originates in close contact with the amphisiellid median cirral row. The amphisiellid median cirral row commences anlagen formation within-row and originates from three rightmost anlagen. All dorsal kineties develop intrakinetally. Usually one postperistomial cirrus developing from third anlage from right. One cirrus left of amphisiellid median cirral row. Transverse cirri longitudinally arranged, originate from single anlage. Caudal cirri present. 4 This postperistomial ventral cirrus is certainly homologous to the postoral ventral cirrus IV/2 of the 18cirri hypotrichs (see cell division, general section, and Berger 1999, p. 16).
Hemiamphisiella
289
Remarks: Foissner (1988) established Hemiamphisiella for H. terricola Foissner, 1988 (= Strongylidium muscorum Kahl, 1932 sensu Foissner [1984]). Few years later, Eigner & Foissner (1994) studied the ontogenesis of this species and found highly interesting details indicating that the establishment of Hemiamphisiella was appropriate (however, see Pseudouroleptus paragraph below). As a consequence, they redefined Hemiamphisiella. Petz & Foissner (1996) slightly modified this improved diagnosis (see corresponding footnotes). Foissner (1988) transferred three further species to Hemiamphisiella, two of which (H. wilberti and H. granulifera) are indeed closely related to the type species. Hemiamphisiella quadrinucleata, the third species, however, differs distinctly from the other species, inter alia, in body shape (posterior end broadly rounded vs. more or less distinctly tailed), anterior end of left marginal row (straight vs. at least slightly curved rightwards), and dorsal kinety pattern (dorsomarginal row very likely present vs. absent1). Hemiamphisiella quadrinucleata resembles Nudiamphisiella interrupta as concerns the ventral and dorsal infraciliature, indicating that it is more closely related to this species than to the other Hemiamphisiella species. However, Hemiamphisiella quadrinucleata has a more or less isolated cirrus behind the proximal end of the adoral zone, a feature characterising Hemiamphisiella, but lacking in Nudiamphisiella. Unfortunately, ontogenetic data for H. quadrinucleata are lacking so that we do not know whether or not this cirrus is a true postperistomial cirrus as in H. terricola; ontogenetic data of H. terricola (Eigner & Foissner 1994) show that this cirrus is homologous to cirrus IV/2 of the 18-cirri hypotrichs (for review see Berger 1999, p. 16). However, it could also be possible that in H. quadrinucleata this cirrus originates not from anlage IV, but that it is the posteriormost cirrus of the anterior portion of the amphisiellid median cirral row (see Fig. 62d for explanation). Since ontogenetic data are lacking neither of the possibilities can be proved or disproved. Thus, I retain the classification in Hemiamphisiella, but list H. quadrinucleata as incertae sedis in Hemiamphisiella and include it also in the Nudiamphisiella key. Pseudouroleptus Hemberger, 1985 is very similar to Hemiamphisiella. I assigned the type species P. caudatus Hemberger, 1985 to the oxytrichids because of the presence of dorsal kinety fragmentation, that is, kinety 3 divides and the anterior portion forms kinety 3 and the posterior forms kinety 4 (Fig. 136c; Berger 1999, p. 888). The other four species so far assigned to Pseudouroleptus have, inter alia, a different dorsal kinety pattern so that the monophyly of this genus is very questionable. Thus I remove them from Pseudouroleptus and put them into the new genus Bistichella (present book). Unfortunately, the ontogenesis is not known for the type population of Hemiamphisiella terricola terricola, which has, like P. caudatus, four dorsal kineties (Fig. 56f). Eigner & Foissner (1994) studied the cell division of another H. terricola terricola population which has, however, only three dorsal kineties showing, as expected, no kinety fragmentation or dorsomarginal row (Fig. 57e, 58g, h). 1
Dorsal kinety 4 of the type population of the type species H. terricola (Fig. 56f) is very likely (almost certainly) not a dorsomarginal row.
290
SYSTEMATIC SECTION
Table 23 Morphometric data on Hemiamphisiella granulifera (gr1, type population from Foissner 1987a; gr2, population from Kenya, original data kindly supplied by W. Foissner), Hemiamphisiella quadrinucleata (qua, from Foissner 1984), Hemiamphisiella terricola qingdaoensis (qin, from Song & Wilbert 1989), Hemiamphisiella terricola terricola (te1, Austrian population from Foissner 1984; te2, Fiji island population from Foissner 1987a; te3, Australian population from Blatterer & Foissner 1988), and Hemiamphisiella wilberti (wil, from Foissner 1982) Characteristics a Body, length
Species mean
gr1 gr2 qin qua te1 te2 te3 wil Body, width gr1 gr2 qin qua te1 te2 te3 wil Adoral zone of membranelles, length gr1 gr2 qin qua te1 te2 te3 wil Anterior body end to anterior end of te1 amphisiellid median cirral row, distance wil d Anterior body end to rear end of amphigr1 siellid median cirral row, distance gr2 qua te2 Posterior body end to rear end of amphi- te1 siellid median cirral row, distance Anterior body end to anterior end of te1 right ventral row, distance te3 wil d Macronuclear nodule, length gr1 gr2 qua te1 te2 te3 wil Macronuclear nodule, width gr1 gr2
M
SD
SE
CV
Min
Max
n
136.9 118.4 161.0 68.1 152.3 135.0 245.0 171.1 25.0 38.3 34.7 24.8 34.6 30.4 61.1 33.8 32.8 35.9 34.2 20.7 40.5 31.7 53.3 43.4 10.9 15.0 108.0 91.5 46.3 101.5 29.9
130.0 118.0 153.0 67.0 148.0 132.0 240.5 175.0 24.0 38.5 36.0 25.0 34.0 30.0 59.5 32.5 34.0 36.5 34.0 21.0 41.0 32.0 54.0 43.0 11.0 15.0 109.0 89.5 46.0 100.0 31.0
14.9 9.6 21.8 7.0 16.9 18.8 29.4 18.5 4.2 5.0 4.5 2.7 5.0 3.7 14.4 5.4 4.3 4.1 2.6 1.1 4.4 2.6 3.7 3.2 2.0 1.4 12.3 6.8 4.4 12.2 6.3
– 3.4 10.9 1.8 4.7 – 9.3 5.9 – 1.8 3.2 0.7 1.4 – 4.6 1.7 – 1.4 1.8 0.3 1.2 – 1.2 1.0 0.6 0.4 – 2.4 1.1 – 1.8
10.9 8.1 6.8 10.3 11.1 13.9 12.0 10.8 16.7 12.9 12.8 10.9 14.6 12.1 23.6 16.1 13.1 11.4 7.5 5.1 10.9 8.4 7.0 7.3 18.1 9.4 11.4 7.4 9.4 12.0 21.2
120.0 105.0 135.0 56.0 137.0 115.0 197.0 150.0 18.0 31.0 31.0 21.0 28.0 25.0 40.0 25.0 25.0 29.0 32.0 18.0 35.0 28.0 47.0 40.0 8.0 13.0 91.0 83.0 38.0 82.0 21.0
160.0 132.0 180.0 83.0 202.0 175.0 291.0 206.0 32.0 45.0 38.0 28.0 45.0 39.0 92.0 43.0 40.0 41.0 37.0 22.0 50.0 35.0 59.0 49.0 14.0 17.0 120.0 105.0 56.0 133.0 42.0
9 8 16 15 13 11 10 10 9 8 16 15 13 11 10 10 9 8 10 15 13 11 10 10 13 10 5 8 15 11 13
106.8 148.0 72.0 16.0 15.9 8.3 7.3 8.1 10.9 18.6 7.6 8.0
106.0 151.0 73.0 15.0 16.0 8.4 7.0 8.0 11.5 19.0 7.0 8.0
11.3 20.8 8.1 1.9 3.5 1.4 1.6 2.2 2.5 2.1 0.7 1.1
3.1 6.6 2.6 – 1.2 0.4 0.4 – 0.8 0.7 – 0.4
10.6 91.0 140.0 14.0 124.0 185.0 11.2 53.0 84.0 12.1 14.0 20.0 21.9 10.0 21.0 17.2 5.6 10.0 21.4 5.6 10.0 26.7 5.0 11.0 23.2 6.0 14.0 11.3 15.0 21.0 9.6 7.0 9.0 13.4 7.0 10.0
13 10 10 9 8 15 13 11 10 10 9 8
Hemiamphisiella
291
Table 23 Continued Characteristics a Macronuclear nodule, width
Species mean
qua te1 te2 te3 wil Macronuclear nodules, distance in between wil Macronuclear nodules, number gr1 gr2 qin qua te1 te2 te3 wil Micronucleus, length gr1 gr2 qua te3 wil Micronucleus, width qua te3 wil Micronuclei, number gr1 gr2 qin qua te3 wil Adoral membranelles, number gr1 gr2 qin qua te1 te2 te3 wil Frontal cirri, number gr1 b gr2 b qin b qua te1 b te2 b te3 b wil b Buccal cirri, number gr1 gr2 qin qua te1
5.7 3.1 4.7 6.2 8.9 7.4 2.0 2.0 12.8 4.2 28.8 12.5 21.2 2.0 3.0 3.3 2.7 5.1 7.1 2.1 4.4 4.2 2.9 2.1 2.7 2.0 3.9 2.1 28.6 33.1 31.4 21.3 44.9 31.8 42.6 48.0 3.9 3.9 4.0 2.9 4.0 4.0 3.9 4.0 1.0 1.0 1.0 0.9 1.0
M
SD
SE
CV
Min
5.6 3.0 4.0 6.0 9.3 7.5 2.0 2.0 12.0 4.0 28.0 14.0 22.5 2.0 3.0 3.0 2.8 4.9 6.9 2.1 4.3 4.0 3.0 2.0 2.0 2.0 4.0 2.0 29.0 33.0 31.0 22.0 44.0 32.0 43.0 48.5 4.0 4.0 4.0 3.0 4.0 4.0 4.0 4.0 1.0 1.0 1.0 1.0 1.0
0.5 0.4 1.1 1.6 0.6 3.3 0.0 0.0 2.4 0.6 3.6 3.0 4.4 0.0 0.4 0.5 0.4 0.9 0.8 0.2 1.0 0.4 1.5 0.4 1.4 0.0 1.5 0.3 4.7 2.0 2.2 1.4 4.4 2.3 3.8 3.8 – 0.4 0.0 0.3 0.0 0.0 0.3 0.0 0.0 0.0 0.0 0.3 0.0
0.1 0.1 – 0.5 0.2 1.1 – 0.0 0.8 0.1 1.0 – 1.4 0.0 – 0.2 0.9 0.3 0.3 0.1 0.3 0.1 – 0.1 0.5 0.0 0.5 0.1 – 0.7 0.8 0.4 1.2 – 1.2 1.2 – 0.1 0.0 0.1 0.0 – 0.1 0.0 – 0.0 0.0 0.1 0.0
7.9 13.4 23.4 26.4 6.8 45.0 0.0 0.0 18.7 13.4 12.5 24.4 21.0 0.0 14.4 14.2 13.2 16.9 11.6 10.7 21.8 10.1 53.4 16.6 50.1 0.0 39.1 14.3 16.5 5.9 7.0 6.5 9.8 7.1 9.0 8.0 – 9.1 0.0 8.8 0.0 0.0 8.1 0.0 0.0 0.0 0.0 27.3 0.0
4.8 2.8 3.9 3.8 8.0 2.0 2.0 2.0 10.0 4.0 25.0 8.0 14.0 2.0 2.0 3.0 2.1 4.0 5.5 1.5 3.2 3.7 1.0 2.0 1.0 2.0 1.0 2.0 23.0 30.0 28.0 18.0 37.0 28.0 37.0 42.0 3.0 3.0 4.0 2.0 4.0 4.0 3.0 4.0 1.0 1.0 1.0 0.0 1.0
Max 7.0 4.2 6.0 9.0 9.5 12.0 2.0 2.0 16.0 6.0 35.0 16.0 26.0 2.0 4.0 4.0 3.5 6.2 8.0 2.5 6.1 5.3 5.0 3.0 5.0 2.0 6.0 3.0 37.0 36.0 35.0 23.0 52.0 35.0 49.0 53.0 4.0 4.0 4.0 3.0 4.0 4.0 4.0 4.0 1.0 1.0 1.0 1.0 1.0
n 15 13 11 10 10 10 9 8 16 15 13 11 10 10 9 8 15 10 10 15 10 10 9 8 16 15 10 10 9 8 14 15 13 11 10 10 9 8 16 15 13 11 10 10 9 8 16 15 13
292
SYSTEMATIC SECTION
Table 23 Continued Characteristics a Buccal cirri, number
Cirri left of anterior portion of amphisiellid median cirral row, number f Amphisiellid median cirral row, number of cirri
Species mean te2 te3 wil qua
gr1 c gr2 qin qua te1 te2 te3 wil Right ventral row, number of cirri gr2 qin h te1 te2 te3 wil Postperistomial ventral cirri, number gr1 gr2 qin qua te1 te2 te3 wil Left marginal cirri, number gr1 gr2 qin qua te1 te2 te3 wil i Right marginal cirri, number gr1 gr2 qin qua te1 te2 te3 wil Cirri ahead of right marginal row, number qin Dorsal kineties, number gr1 gr2 qin qua te1
M
SD
SE
CV
Min
Max
n
1.0 1.0 1.0 1.1
1.0 1.0 1.0 1.0
0.0 0.0 0.0 0.5
– 0.0 0.0 0.1
0.0 0.0 0.0 42.8
1.0 1.0 1.0 0.0
1.0 1.0 1.0 2.0
11 10 10 15
40.9 35.6 41.7 15.4 61.3 40.0 61.5 63.7 3.4 2.8 10.7 3.3 14.3 22.4 1.0 1.0 1.4 1.0 1.0 1.0 1.2 0.9 39.4 36.6 34.2 20.3 58.3 42.1 53.0 59.6 43.6 37.5 37.2 24.7 62.7 44.3 59.4 60.5 2.8 3.0 3.0 3.0 4.0 4.0
40.0 36.5 42.0 16.0 62.0 40.0 61.5 64.0 3.0 3.0 10.0 3.0 15.0 23.5 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 38.0 37.5 36.0 20.0 57.0 42.0 53.5 62.0 44.0 37.0 37.0 25.0 63.0 45.0 59.5 59.0 3.0 3.0 3.0 3.0 4.0 4.0
5.1 3.9 4.5 1.8 4.8 4.1 5.0 5.4 0.5 0.3 1.9 0.9 5.1 3.7 0.0 0.0 0.5 0.0 0.0 0.0 0.8 0.3 6.9 4.9 4.1 1.7 5.5 4.2 5.5 7.0 5.7 3.6 2.2 2.1 4.0 4.4 6.0 7.3 0.4 0.0 0.0 0.0 0.0 0.0
– 1.4 2.6 0.5 1.3 – 1.6 1.7 0.2 0.1 0.5 – 1.7 1.2 – 0.0 0.1 0.0 0.0 – 0.2 0.1 – 1.7 1.2 0.4 1.5 – 1.7 2.2 – 1.3 0.6 0.5 1.1 – 1.9 2.3 0.1 – 0.0 0.0 0.0 0.0
12.4 11.0 10.8 11.7 7.8 10.3 8.0 8.5 15.3 13.3 17.7 27.7 35.4 16.7 0.0 0.0 35.7 0.0 0.0 0.0 65.7 33.3 17.4 13.4 12.1 8.2 9.5 10.1 10.4 11.7 13.0 9.7 5.9 8.4 6.3 10.0 10.1 12.0 14.2 0.0 0.0 0.0 0.0 0.0
36.0 28.0 37.0 13.0 54.0 33.0 53.0 55.0 3.0 2.0 8.0 2.0 6.0 15.0 1.0 1.0 1.0 1.0 1.0 1.0 0.0 0.0 34.0 29.0 28.0 17.0 50.0 37.0 44.0 50.0 35.0 32.0 35.0 21.0 57.0 38.0 51.0 52.0 2.0 3.0 3.0 3.0 4.0 4.0
51.0 39.0 46.0 20.0 69.0 46.0 71.0 72.0 4.0 3.0 14.0 5.0 24.0 27.0 1.0 1.0 2.0 1.0 1.0 1.0 3.0 1.0 57.0 45.0 39.0 23.0 71.0 46.0 61.0 71.0 53.0 42.0 41.0 28.0 71.0 52.0 70.0 75.0 3.0 3.0 3.0 3.0 4.0 4.0
9 8 10 15 13 11 10 10 8 16 13 11 9 10 9 8 16 15 13 11 10 10 9 8 10 15 13 11 10 10 9 8 10 15 13 11 10 10 16 9 8 16 15 10
Hemiamphisiella
293
Table 23 Continued Characteristics a Dorsal kineties, number
Caudal cirri, number g
Species mean te2 te3 e wil qin qua
3.0 3.0 4.0 3.0 3.1
M
SD
SE
CV
Min
3.0 3.0 4.0 3.0 3.0
0.0 0.0 0.0 0.0 0.3
– 0.0 0.0 0.0 0.1
0.0 0.0 0.0 0.0 8.4
3.0 3.0 4.0 3.0 3.0
Max 3.0 3.0 4.0 3.0 4.0
n 11 10 4 16 15
a All measurements in µm. CV = coefficient of variation in %, M = median, Max = maximum value, mean = arithmetic mean, Min = minimum value, n = number of individuals investigated, SD = standard deviation, SE = standard error of arithmetic mean. Data based on protargol-impregnated specimens. b
Cirrus III/2 included.
c
Cirri (about three) of right ventral row included.
d
In this parameter, Foissner (1982) confused “left“ and “right ventral row”.
e
Blatterer & Foissner (1988) wrote 3.1 as arithmetic mean and 3 as minimum and maximum value. I checked the raw data and found that one specimen had 3 1/2 kineties (three kineties of body length and one distinctly shortened kinety). f
The other species usually have one cirrus (= cirrus III/2) left of the anterior portion of the amphisiellid median cirral row. However, in all morphometrics this cirrus was counted as fourth frontal cirrus.
g
Difficult to distinguish from left marginal row in H. terricola terricola; thus, the caudal cirri, which are very likely present (see cell division), are included in the number of left marginal cirri in this subspecies.
h
Designated as transverse cirri in original description.
i
Caudal cirri, if present, included.
Both in P. caudatus (Hemberger 1982, 1985) and H. terricola terricola (Eigner & Foissner 1994) the median cirral row is formed from three anlagen and each species has a conspicuous postoral (= postperistomial) ventral cirrus formed from anlage IV. The length of the right ventral row is likely insufficient to separate Pseudouroleptus and Hemiamphisiella because it is only a quantitative feature. Thus, at the present state of knowledge we cannot decide whether or not these two genera are synonymous. As a consequence, I preliminarily accept both the present genus (dorsal kinety fragmentation lacking) and Pseudouroleptus whose type species shows such a fragmentation (p. 658). The higher-level classification of Hemiamphisiella is difficult. Most often it was assigned to the amphisiellids (Foissner 1988, with doubt; Eigner & Foissner 1994, Petz & Foissner 1996, Lynn & Small 2002, Berger 2005b). According to Eigner (1997, p. 555; 1999, p. 46), Hemiamphisiella belongs to the Oxytrichidae because of the neokinetal 3 anlagen formation, that is, anlagen V and VI of both filial products originate from a V-shaped primordium. I am uncertain about the proper systematic position because, on the one hand, Hemiamphisiella forms a distinct cirral row which is reminiscent of the amphisiellid median cirral row, on the other hand H. terricola is obviously very similar to Pseudouroleptus caudatus (see above), a species
294
SYSTEMATIC SECTION
which I assigned to the oxytrichids because of the dorsal kinety fragmentation. However, since dorsal kinety fragmentation is not confirmed for Hemiamphisiella, I preliminarily retain it in the amphisiellids. The lack of transverse cirri is possibly an apomorphy because their presence is a feature of the ground pattern of the Hypotricha. The right cirral row between the rear portion of the marginal rows is likely a further apomorphy because in other amphisiellids and hypotrichs in general the rear portion of anlage VI is composed of two cirri only (VI/1 [rightmost transverse cirrus] and VI/2 [right pretransverse ventral cirrus]). Shi (1999, p. 253) and Shi et al. (1999, p. 100) put Hemiamphisiella and Paramphisiella into the synonymy of Uroleptoides. However, the type species U. kihni lacks a postperistomial cirrus strongly indicating that this synonymy is not justified. Species included in Hemiamphisiella (alphabetically arranged basionyms are given): (1) Hemiamphisiella terricola Foissner, 1988; (2) Strongylidium granuliferum Foissner, 1987a; (3) Strongylidium wilberti Foissner, 1982; (4) Uroleptoides qingdaoensis Song & Wilbert, 1989. Incertae sedis: (5) Uroleptoides quadrinucleata Foissner, 1984.
Key to Hemiamphisiella species Note the resemblance of Hemiamphisiella and Pseudouroleptus (p. 658)! 1 Two macronuclear nodules (e.g., Fig. 61a, e). . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 - More than 2 macronuclear nodules (e.g., Fig. 56a, f). . . . . . . . . . . . . . . . . . . . . . 3 2 Cortical granules colourless, 1–2 µm across, globular, densely arranged in longitudinal rows; few cirri between rear portion of marginal rows; 3 dorsal kineties (Fig. 61a–e). . . . . . . . . . . . . . . . . . . . . . . . . Hemiamphisiella granulifera (p. 315) - Cortical granules lacking; many cirri between rear portion of marginal rows; 4 dorsal kineties (Fig. 60a–g). . . . . . . . . . . . . . . . Hemiamphisiella wilberti (p. 311) 3 (1) Usually 4 (rarely up to 6) macronuclear nodules; posterior body end broadly rounded (Fig. 62a–e). . . . . . . . . . . . . . . Hemiamphisiella quadrinucleata (p. 318) - 8–35 macronuclear nodules; posterior body portion narrowed (Fig. 56a, 57a, 58a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemiamphisiella terricola (p. 294)
Hemiamphisiella terricola Foissner, 1988 (Fig. 56a–f, 57a–e, 58a–o, 59a–g, Table 23) 1984 Strongylidium muscorum Kahl, 1932 – Foissner, Stapfia, 12: 107, Abb. 55a–f, Tabelle 26 (Fig. 56a–f; misidentification [see remarks at H. terricola terricola]; for slide deposition, see nomenclature of H. terricola terricola). 1987 Strongylidium muscorum Kahl, 1932 – Foissner, Zool. Beitr. (N.F.), 31: 192, Abb. 2a–e, Tabelle 1 (Fig. 57a–e; misidentification; description of a Fiji Islands population; at least one voucher slide is
Hemiamphisiella
1988 1988
1989
1994 2001 2001
2002
295
deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; Foissner 1987a, p. 190). Hemiamphisiella terricola nov. spec. – Foissner, Stapfia, 17: 122 (original description; for slide deposition, see nomenclature of H. terricola terricola). Hemiamphisiella terricola Foissner, im Druck – Blatterer & Foissner, Stapfia, 17: 38, Tabelle 7 (morphometric characterisation of an Australian population; at least one voucher slide is deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; Blatterer & Foissner 1988, p. 5). Uroleptoides qingdaoensis sp. nov. – Song & Wilbert, Acta Zootax. sinica, 14: 391, Fig. 1-7, Table 1 (Fig. 59a–g; original description; in present paper classified as subspecies of H. terricola; slides of holotype specimens have been deposited in the Fishery College of the Ocean University of Qingdao, China). Hemiamphisiella terricola Foissner, 1988 – Eigner & Foissner, J. Euk. Microbiol., 41: 250, Fig. 40–55 (Fig. 58a–o; morphogenesis of population described by Blatterer & Foissner 1988). Hemiamphisiella terricola Foissner, 1988 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 30 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs and euplotids). Uroleptoides qingdaoensis Song and Wilbert, 1989 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 96 (nomenclator containing all basionyms, combinations, and higher taxa of euplotids and hypotrichs). Hemiamphisiella terricola – Lynn & Small, Phylum Ciliophora, p. 452, Fig. 35A, B (Fig. 56e; guide to ciliate genera).
Nomenclature: No derivation of the species-group name terricola is given in the original description. It is a composite of the Latin noun terr·a (soil), the thematic vowel ·i-, and the Latin verb colere (to live in) and means living in soil, and refers to the habitat where the species was discovered. Usually, species-group names ending with -cola are considered as appositive substantives and are thus not changed when transferred to a genus of different gender (Werner 1972, p. 138). Type species of Hemiamphisiella. For derivation of the name qingdaoensis, see subspecies H. terricola qingdaoensis. Remarks: For explanation of the misidentification by Foissner (1984, 1987a), see remarks at H. terricola terricola. According to Eigner & Foissner (1994, p. 250) and Foissner (1998, p. 210), Uroleptoides qingdaoensis Song & Wilbert, 1989 is possibly a junior synonym of H. terricola, which has cortical granules. By contrast, Uroleptoides qingdaoensis has no special cortical granules according to a hand-written note by Weibo Song in my reprint1. Thus, I do not synonymise U. qingdaoensis with H. terricola, but classify it as subspecies of this species. Further studies on Chinese populations will show whether or not the lack of the cortical granules is constant. Whether the different populations of H. terricola terricola are also distinct subspecies has to be checked by studying further populations from various regions.
1
I do not know whether the Chinese text of Song & Wilbert (1989) contains a note about the presence or absence of cortical granules; the English abstract does not.
296
SYSTEMATIC SECTION
Key to the subspecies of Hemiamphisiella terricola 1 Cortical granules colourless, globular, about 1.0–1.5 µm across (Fig. 56c, 57b, c). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemiamphisiella terricola terricola (p. 296) - Cortical granules lacking. . . . . . Hemiamphisiella terricola qingdaoensis (p. 310)
Hemiamphisiella terricola terricola Foissner, 1988 stat. nov. (Fig. 56a–f, 57a–e, 58a–o, Table 23) 1984 Strongylidium muscorum Kahl, 1932 – Foissner, Stapfia, 12: 107, Abb. 55a–f, Tabelle 26 (Fig. 56a–f; misidentification [see remarks]; for slide deposition, see nomenclature). 1987 Strongylidium muscorum Kahl, 1932 – Foissner, Zool. Beitr. (N.F.), 31: 192, Abb. 2a–e, Tabelle 1 (Fig. 57a–e; description of a Fiji Islands population; at least one voucher slide is deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; Foissner 1987a, p. 190). 1988 Hemiamphisiella terricola nov. spec.1 – Foissner, Stapfia, 17: 122 (original description; for slide deposition see nomenclature). 1988 Hemiamphisiella terricola Foissner, im Druck – Blatterer & Foissner, Stapfia, 17: 38, Tabelle 7 (morphometric characterisation of an Australian population; at least one voucher slide is deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; Blatterer & Foissner 1988, p. 5). 1994 Hemiamphisiella terricola Foissner, 1988 – Eigner & Foissner, J. Euk. Microbiol. 41: 250, Fig. 40–55 (Fig. 58a–o; morphogenesis of population described by Blatterer & Foissner 1988). 2001 Hemiamphisiella terricola Foissner, 1988 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 30 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs and euplotids). 2002 Hemiamphisiella terricola – Lynn & Small, Phylum Ciliophora, p. 452, Fig. 35A, B (Fig. 56e; guide to ciliate genera).
Nomenclature: For derivation of name, see H. terricola. The appendix stat. nov. (status novum, new status) in the heading means that this species is now classified as subspecies. Nominotypical subspecies, that is, type of H. terricola. Foissner (1984, p. 8) wrote that from all species redescribed in his paper at least one slide is deposited in the Oberösterreichische Landesmuseum in Linz (Upper Austria). Foissner (1988) recognised that the identification of the present species as S. muscorum was incorrect and thus established a new species for this population. On page 123 he wrote that the type material of H. terricola is deposited in Linz under the name Strongylidium muscorum. According to Aescht (2003, p. 391) the slide with the accession number “1984/83” is the neotype slide of S. muscorum. A neotypification of S. muscorum, however, would have made the establishment of H. terricola unnecessary, respectively, impossible. I checked the literature carefully and found that neither Foissner (1984) nor anybody else has ever neotypified Strongylidium muscorum validly according to the detailed instructions of the ICZN (1964, 1985, 1999). Thus, the description of S. muscorum sensu Foissner (1984) as Hemi1
Foissner (1988) provided the following diagnosis: In vivo etwa 170–240 × 25–45 µm große, posterior deutlich verschmälerte Hemiamphisiella mit farblosen, in Reihen angeordneten, kugeligen subpelliculären Granula. Viele (>10) Makronucleus-Teile. Rechte Ventralreihe stark verkürzt. Durchschnittlich 44 adorale Membranellen. 4 Dorsalkineten.
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amphisiella terricola by Foissner (1988) is valid and the slide 1984/83 in the museum in Linz is the holotype slide of H. terricola. When the subspecies status is accepted, the name has to be cited as follows: Hemiamphisiella terricola terricola Foissner, 1988. Remarks: Foissner (1984) identified this species as “Strongylidium muscorum Kahl, 1932” because of the nuclear apparatus, the sigmoidal body shape, the cortical granules, and the terrestrial habitat. He also discussed the differences between the original description and his description of an Austrian population, namely, the body length (100–110 µm vs. 170–240 µm) and the length of the right ventral row (not shortened vs. shortened). Foissner (1987a) described a population from the Fiji Islands whose specimens are distinctly smaller than those of the Austrian population, and therefore resemble the specimens described by Kahl (1932); however, since the extreme values overlap he considered the populations not as distinct species, but as ecoforms. Somewhat later, Foissner (1988) withdrew his identification as S. muscorum and considered the different body length and length of the right ventral row as main differences. Further, Blatterer & Foissner (1988) described an Australian population whose specimens are still larger than those of the Austrian population. Further populations have to be described to show whether these morphometric differences are stable or not. Stable differences would indicate the existence of further subspecies or even species. The description of the Azerbaijanian population by Alekperov (2005, p. 217) is in Russian and was not translated in detail. However, the illustration shows that the identification is very likely incorrect (Fig. 63a). Thus, this population is briefly characterised in the chapter on insufficient redescriptions. Strongylidium Sterki, 1878, respectively, its type species S. crassum is a little known taxon only briefly characterised in the original description. According to Kahl (1932, p. 551) it has a twisted body and two parallel ventral rows. Whether these two rows are a midventral complex composed of cirral pairs as in the urostyloids (for review, see Berger 2006) or two true frontoventral cirral rows is not known. In addition, Strongylidium likely lacks the conspicuous postoral ventral cirrus, making a synonymy of Strongylidium and Hemiamphisiella very unlikely. Fernandez-Leborans & Antonio-Garcia (1988, p. 147) recorded a Strongylidium muscorum. Since no identification literature is given it is not known whether the record refers to S. muscorum Kahl, 1932 or to Hemiamphisiella terricola Foissner, 1988 (= S. muscorum sensu Foissner 1984). Thus, this record from the Manzanares river (La Pedriza, Madrid, Spain) is not mentioned in the ecology section of the present species. Morphology: As discussed above, the three populations described so far differ very distinctly morphometrically so that one cannot exclude that they are subspecies or even species, inasmuch as they are from rather different locations (Austria, Australia, Fiji Islands). Thus, the descriptions are kept separate. Austrian population (= type population) described by Foissner (1984; Fig. 56a–f): Body size 170–240 × 25–45 µm in life. Body outline sigmoidal, anterior
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Fig. 56a–f Hemiamphisiella terricola terricola (from Foissner 1984. a, b, d, from life; c, e, f, protargol impregnation). a: Ventral view of representative specimen, 230 µm. b: Mitochondria. c: The colourless, about 1.5 µm-sized cortical granules became tiny dots in protargol preparations. d: Shape variant in dorsal view. e, f: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 163 µm. Long arrow marks postperistomial ventral cirrus, short arrow denotes right ventral row. Broken line connects cirri which originate from the same anlage (only shown for anlagen II and III), dotted line connects frontal cirri.
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portion slightly, posterior distinctly narrowed. Body flexible, about 2:1 flattened dorsoventrally, somewhat contractile, especially the rear portion, which becomes round soon after placing the cover slip on the specimen. 28 macronuclear nodules on average, arranged mainly in mid-body portion and left of midline; individual nodules about 14 × 6 µm in life, with many small chromatin bodies. Several, in life about 6.0 × 3.5 µm-sized, strongly refractive micronuclei (not observed in protargol preparations). Contractile vacuole near left body margin immediately behind proximal end of adoral zone. Cortical granules colourless (see Foissner 1987a, p. 194) and about 1.4 µm across in life make cells brownish; in protargol preparations granules are tiny and impregnated (Fig. 56c). Pellicle colourless. Cytoplasm, especially in posterior body portion, often packed with 1–3 µm-sized, indistinctly cuboidshaped, yellowish crystals which appear dark at low magnification. Food vacuoles 7–14 µm across. Movement without peculiarities, that is, slowly gliding; specimens often changing the direction. Adoral zone of membranelles occupies about 27% of body length on average (Table 23), formed like a question mark, composed of 45 membranelles on average (Fig. 56a, e, Table 23); proximal end about in midline, distal end extends slightly onto right body side. Bases of largest membranelles in life about 10 µm wide. Buccal field rather flat, undulating membranes slightly curved and only inconspicuously crossing optically (Fig. 56e). Pharyngeal fibres well recognisable in life and protargol preparations. Cirral pattern and number of cirri of usual variability (Fig. 56e, Table 23). Three frontal cirri distinctly enlarged. Cirrus III/2 somewhat smaller than frontal cirri, but larger than remaining cirri. Buccal cirrus slightly behind anterior end of paroral (Fig. 56e). One postperistomial ventral cirrus behind proximal end of adoral zone. Amphisiellid median cirral row continuous although composed of three portions (see cell division), commences right of right frontal cirrus, extends sigmoidally to 80% of body length on average (Table 23). Right ventral row commences at 70% of body length on average (Table 23), composed of widely spaced cirri. Transverse cirri lacking. All cirral rows distinctly twisted; cirri in life about 15 µm long, very narrowly spaced within rows, except in posterior portion where they are more widely spaced and finer. Right marginal row commences on dorsal side about at level of distal end of adoral zone, terminates on ventral side slightly ahead of rear cell end; left row distinctly curved rightwards anteriorly, extends onto dorsal surface posteriorly and somewhat on right body margin so that marginal rows overlap optically; however, rearmost three left marginal cirri are very likely caudal cirri (see cell division).
b
ACR = amphisiellid median cirral row, AZM = adoral zone of membranelles, CV = contractile vacuole, LMR = left marginal row, MA = macronuclear nodule, P = paroral, RMR = right marginal row, III/2 = cirrus behind right frontal cirrus, 1–4 = dorsal kineties. Page 296.
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Fig. 57a–e Hemiamphisiella terricola terricola (from Foissner 1987a. a, b, from life; c–e, protargol impregnation). a: Ventral view of representative specimen, 178 µm. b: Dorsal view showing arrangement of cortical granules. c: Part of pellicle with argyrophilic granules (= cortical granules). d, e: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 136 µm. Long arrow in (d) marks postperistomial ventral cirrus, short arrow marks right ventral row. Dotted line connects frontal cirri. ACR = amphisiellid median cirral row, AZM = adoral zone of membranelles, BC = buccal cirrus, LMR = left marginal row, MA = macronuclear nodule, RMR = right marginal row, III/3 = cirrus behind right frontal cirrus, 1–3 = dorsal kineties. Page 296.
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Fig. 58a–c Hemiamphisiella terricola terricola (from Eigner & Foissner 1994. Protargol impregnation). Infraciliature of ventral side of dividers of Australian population described by Blatterer & Foissner (1988). (details, see text). a: Very early divider, 186 µm. Arrow marks postperistomial ventral cirrus. b: Early divider, 159 µm. c: Middle divider, 150 µm. Arrow marks primordium for right marginal row of opisthe. OP = oral primordium, I–VI = frontal-ventral cirri anlagen. Page 296.
Dorsal bristles about 3 µm long according to Fig. 56f, arranged in four kineties; kineties 1–3 of body length, row 41 commences at 20% of body length. Very likely 1
Whether this is an ordinary bipolar kinety, a dorsomarginal kinety, or a kinety originating by fragmentation of kinety 3 is not known because the cell division was studied on a population with only three kineties (Fig. 58g, h). However, if the arrangement shown in Fig. 56f is constant, it can be excluded that ki-
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Fig. 58d, e Hemiamphisiella terricola terricola (from Eigner & Foissner 1994. Protargol impregnation; details, see text; parental structures white, new black). d: Middle divider (149 µm) showing six frontalventral cirri anlagen (I–VI) in each filial product. e: Late divider, 144 µm. Arrows mark migration of postperistomial ventral cirrus (homologous to cirrus IV/2 of the 18-cirri hypotrichs). Arrowheads denote bipartition of anlagen VI; the anterior portion, which is homologous to the frontoterminal cirri of the urostyloids or cirri VI/3 and VI/4 of the 18-cirri hypotrichs, forms the anterior portion of the amphisiellid median cirral row; the posterior part forms the right ventral row. Dotted line connects frontal cirri, broken lines connect cirri originating from same anlage (only shown for anlagen II and III). Page 296. nety 4 is a dorsomarginal kinety because such rows are usually not shortened anteriorly, but distinctly shortened posteriorly.
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inconspicuous caudal cirri present, only inconspicuously separated from rear end of left marginal row. Fiji Island population described by Foissner (1987a; Fig. 57a–e): the specimens of this population are distinctly smaller than those of the Austrian and Australian population (Table 23), but the cirral pattern agrees very well. Thus, Foissner (1987a) did not describe this population in detail, but provided (i) illustrations from live specimens (Fig. 57a, b); (ii) the infraciliature (Fig. 57d, c); (iii) a morphometric characterisation (Table 23); and (iv) some additional and deviating observations. Cortical granules globular, and colourless (Fig. 57b), after staining with methyl green-pyronin they form a red, slimy cover; after protargol impregnation they become tiny (Fig. 57c). On average only about 13 macronuclear nodules (vs. 28 in type population and 21 in Australian population). Right ventral row distinctly shorter than in type population, that is, does not overlap with rear end of amphisiellid median cirral row. Invariably three dorsal kineties (also three in Australian population, but four in type population). Australian population described by Blatterer & Foissner (1988; Fig. 58i–m): Body size in life 230–320 × 50–90 µm, that is, distinctly larger than the other populations (Table 23). The cell division of this population was described by Eigner & Foissner (1994; see below). Cell division (Fig. 58a–h, n, o): This part of the life cycle is described in detail by Eigner & Foissner (1994) for the Australian population characterised morphometrically by Blatterer & Foissner (1988). The oral primordium originates near the middle portion of the amphisiellid median cirral row (Fig. 58a). The postperistomial cirrus and the amphisiellid median row are obviously unchanged. Somewhat later, a narrow field of basal bodies is formed along the left margin of the amphisiellid median cirral row (Fig. 58b, n); it extends to the rear end of the undulating membranes. The postperistomial cirrus is obviously incorporated in the oral primordium and the buccal cirrus modifies to anlage II of the proter. Cirrus III/2 begins with the modification to anlage III of the proter. Next, the oral primordium divides into a large posterior portion, which forms the adoral zone, and small anterior portion, which forms the frontal-ventral anlagen Fig. 58f–h Hemiamphisiella terricola terricola (from Eigner & Foissner 1994. Protargol impregnation). Infraciliature of dividers (details, see text; parental structures white, new black). f: Ventral view of very late divider, 127 µm. Arrow marks migration of the rearmost cirrus (= postperistomial ventral cirrus) of anlage IV, broken lines connect cirri originating from same anlage (only shown for anlagen I–III, VI). g, h: Dorsal view of middle and late divider, g = 121 µm, h = 154 µm. MA = macronucleus, I–VI = frontalventral cirri anlagen, 1–3 = dorsal kineties. Page 296. Fig. 58i–k Hemiamphisiella terricola terricola (from Eigner & Foissner 1994. i, protargol impregnation; j, k, scanning electron micrographs). i: Infraciliature of anterior portion of ventral side, bar = 20 µm. Arrow marks postperistomial ventral cirrus. j, k: Oral region of interphasic specimens, bar = 20 µm (j), 10 µm (k). Arrow in (j) marks postperistomial ventral cirrus. Explanation of original labelling: ACR = amphisiellid median cirral row, AZM = adoral zone of membranelles, B = buccal cirrus, eM = endoral, LR = left marginal row, PL = buccal lip, pM = paroral, RR = right marginal row, 1–3 = frontal cirri (1 = left cirrus), 4 = cirrus III/2. Page 296.
d
d
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Legend on p. 303
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Hemiamphisiella
Legend on p. 303
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I–IV of the opisthe (Fig. 58c). The parental undulating membranes modify to proter’s anlage I. Anlage IV of the proter develops de novo or from migrating basal bodies of anlage II. The cirri in the middle segment of the amphisiellid median cirral row (formed from anlage IV) are modified to a V-shaped pattern producing proter’s and opisthe’s anlagen V and VI. However, Eigner & Foissner (1994) could not exclude that anlage VI develops de novo. The formation of the membranelles in the opisthe’s adoral zone proceeds from anterior to posterior (Fig. 58d, e). The cirral anlagen align and become longer. Six anlagen are formed per filial product (Fig. 58e, f). Anlage I splits, as is usual, longitudinally to form the undulating membranes and the left frontal cirrus (cirrus I/1). Anlage II forms the middle frontal cirrus and the buccal cirrus. Anlage III produces the right frontal cirrus and cirrus III/2 (= cirrus left of anterior portion of amphisiellid median cirral row). Anlage IV divides; the anterior portion (= all cirri of this anlage except the rearmost cirrus) forms the middle part of the amphisiellid median cirral row, and the rearmost cirrus forms the postperistomial cirrus, which can be easily homologised with the postoral ventral cirrus IV/2 of the oxytrichids (see Berger 1999, p. 16) because of the origin from anlage IV and the migration behind the buccal vertex (rarely, two, three, or no postperistomial cirrus are/is formed). Anlage V produces the posterior portion of the amphisiellid median cirral row (Fig. 58f). The anterior portion of anlage VI forms the anterior part of the amphisiellid median cirral row and the posterior portion of this anlage forms the right ventral row. Eigner & Foissner (1994) designated this row as transverse cirri. However, this designation is misleading because a transverse cirral row is, per definition, formed from the rearmost cirrus of different anlagen, that is, it is a pseudorow and not a true cirral row. In addition, the rearmost cirrus of anlage V, respectively, VI is not enlarged so that one can hardly say that Hemiamphisiella terricola has transverse cirri. The division of the nuclear apparatus proceeds in the usual way, that is, the macronuclear nodules fuse to a single mass and later divide into the species-specific number (Fig. 58g, h). The marginal rows divide by intrakinetal proliferation (Fig. 58c–f). The new dorsal kineties are formed according to the Gonostomum pattern (for explanation see Berger 1999, p. 73), that is, the anlagen are formed within the parental rows. Some basal bodies accumulate at the posterior end of the new rows to form the inconspicuous caudal cirri (Fig. 58g, h). In contrast to the Australian population studied by Eigner & Foissner (1994), the type population of H. terricola terricola has four dor-
b Fig. 58l–o Hemiamphisiella terricola terricola (from Eigner & Foissner 1994. l, protargol impregnation; m–o, scanning electron micrographs). l, m: Ventral views showing cirral pattern and nuclear apparatus, bar = 40 µm. Arrows mark postperistomial ventral cirrus. n: Early divider (bar = 40 µm) in ventral view showing the oral primordium (white arrows) from which a short anlage projects to the proter’s frontal area (black arrow). o: Late divider (bar = 40 µm) in ventral view. The postperistomial cirrus of the opisthe is already at its final position (white arrow), that of the proter not yet. Explanation of original labelling: ACR = amphisiellid median cirral row, LR = left marginal row, Ma = macronuclear nodule, RR = right marginal row, TC = right ventral row. Page 296.
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sal kineties (Fig. 56f, Table 23). Kinety 4 in Fig. 56f commences distinctly behind the anterior cell end, strongly indicating that it is not a dorsomarginal row. Perhaps it originates, as in Pseudouroleptus caudatus, by fragmentation of kinety 3 (Hemberger 1982, 1985; for review see Berger 1999, p. 888). However, if this assumption were correct, Hemiamphisiella would be the junior synonym of Pseudouroleptus (see remarks). Occurrence and ecology: Recorded from all continents except the Antarctica (Foissner 1998) and likely preferably in terrestrial habitats (Foissner 1987, 1998). Type locality is the floodplain on the left bank of the Danube river at the city of Zwentendorf (Lower Austria). Foissner (1984) found it there in the upper (0–2 cm) soil layer in a water meadow with willows; for a detailed description of this site see Foissner et al. (1985; site AA on p. 86). The Fiji Islands population occurred in the soil from the margin of a mangrove swamp near the campus of the University of the Fiji Island Viti Levu, South-Pacific (Foissner 1987a; sample collected by D.J. Patterson). Blatterer & Foissner (1988) found Hemiamphisiella terricola terricola at two sites in Australia, namely (i) in moss from soil of an autochthonous pine forest (Callitris sp.; site 8 in their paper); and (ii) in the rhizosphere of a highly saline soil grown with halophytes and grass from the bank of Lake Alexandrina (Point Pelican) near Adelaide. Further records: soil from four natural forest stands in eastern Austria (Foissner et al. 2005, p. 628); blackish compost soil from a municipal compost plant of Munich, Germany (Foissner 2000a, p. 258); soil from the Enns/Danube River floodplain, Upper Austria (Foissner 2004, p. 370); soil from near the Sourhope Research Station near Kelso in Southern Scotland (Finlay et al. 2001, p. 363; Esteban et al. 2006, p. 142); 0.2–0.5 specimens g-1 dry mass in dune soils from Norderney, a German North Sea island (Verhoeven 2002, p. 189); wet mosses, dry mosses and adhering sandy soil, and wet upper soil layer from and near the Sheldrick waterfalls, Shimba Hills Nature Reserve (4°25'S 39°20'E), Kenya (Foissner 1999, p. 323); soil samples from five sites (numbers 3, 29, 30, 49, 64) from Namibia, including Namib Desert (Foissner et al. 2002, p. 60a); upper 3 cm litter and soil layer from a tropical dry forest in the Santa Rosa National Park (Costa Rica), about 5 km east of the ranch house “La Casona” (Foissner 1995, p. 39).
Fig. 59a–g Hemiamphisiella terricola qingdaoensis (from Song & Wilbert 1989. a, b, from life; c–g, protargol impregnation). a: Ventral view of representative specimen, 192 µm. b: Left lateral view. c, d: Infraciliature of ventral and dorsal side and nuclear apparatus. Arrow in (c) marks cirrus III/2, long arrow in (d) denotes a single dorsal bristle (possibly a remnant of the previous generation), short arrow marks the three anteriormost cirri of the right marginal row, which are, according to Song & Wilbert (1989), possibly frontoterminal cirri. e: Infraciliature of oral region. f, g: Infraciliature of ventral and dorsal side of posterior body portion. Arrow in (f) marks right ventral row (= transverse cirri according to original description); according to the ontogenetic data from H. terricola terricola these are the rearmost cirri formed by anlage VI. AZM = adoral zone of membranelles, ACR = amphisiellid median cirral row, BC = buccal cirrus, CC = caudal cirri, CV = contractile vacuole, E = endoral, LMR = left marginal row, MA = macronuclear nodule, MI = micronucleus, P = paroral, RMR = right marginal row, III/2 = cirrus III/2 (= cirrus behind right frontal cirrus), 1–3 = dorsal kineties. Page 310.
d
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The Austrian population fed on hyphae and spores of fungi, flagellates, and ciliates (Cyclidium glaucoma; Foissner 1984), Fiji Islands specimens ingested heterotrophic flagellates, hyphae, diatoms, ciliates (Colpoda steinii), and in culture also rice starch (Foissner 1987a). Biomass of 106 specimens about 192 mg (Foissner 1987, p. 127; 1998, p. 204).
Hemiamphisiella terricola qingdaoensis (Song & Wilbert, 1989) comb. nov., stat. nov. (Fig. 59a–g, Table 23) 1989 Uroleptoides qingdaoensis sp. nov. – Song & Wilbert, Acta Zootax. sinica, 14: 391, Fig. 1–7, Table 1 (Fig. 59a–g; original description; slides of holotype specimens have been deposited in the Fishery College of the Ocean University of Qingdao, China). 2001 Uroleptoides qingdaoensis Song and Wilbert, 1989 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 96 (nomenclator containing all basionyms, combinations, and higher taxa of euplotids and hypotrichs).
Nomenclature: The species-group name qingdaoensis is obviously a composite of Qingdao, a city in China, and the Latin suffix -ens·is and refers to the area where the species was discovered. The appendices in the heading mean that the species U. qingdaoensis is first combined with Hemiamphisiella (H. qingdaoensis) and then classified as subspecies of H. terricola. In future, if the subspecies classification is accepted, the subspecies has to be cited as follows: Hemiamphisiella terricola qingdaoensis (Song & Wilbert, 1989) Berger, 2008. Remarks: Song & Wilbert (1989) assigned the present population to Uroleptoides. Although this genus is not very well defined (the type species is not described after silver impregnation!), a classification of the present subspecies in Uroleptoides does not seem justified because of the postoral (= postperistomial) ventral cirrus, which is very likely lacking in the type species of Uroleptoides, but is present in U. qingdaoensis. Thus, I transfer U. qingdaoensis to Hemiamphisiella whose type species has the same cirral pattern. For a foundation of the subspecies classification, see H. terricola. The original description of H. terricola qingdaoensis is in Chinese. Thus, the following description is based on the rather detailed English summary, the illustrations, and the morphometry provided by Song & Wilbert (1989). Morphology: Body length in life 150–200 µm; body length:width ratio about 4–5:1; specimen illustrated about 192 × 44 µm (Fig. 59a). Body dorsoventrally flattened (Fig. 59b). Body outline elongate-elliptical to spindle-shaped with posterior body end narrowly rounded (Fig. 59a). Macronucleus composed of 10–16 ellipsoidal nodules, mainly in left body portion (Fig. 59a, d). Contractile vacuole near left body margin, at about 40% of body length (Fig. 59a, b). Special cortical granules lacking
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(handwritten note by W. Song at abstract of original description). Cytoplasm colourless. Adoral zone, undulating membranes, and cirral pattern as in H. terricola terricola and as shown in Fig. 59c–g and given in Table 23. Rather often, not one, but two postperistomial ventral cirri. According to Song & Wilbert (1989), the present subspecies possibly has frontoterminal cirri. I suppose – like Song & Wilbert (1989, p. 395) themselves and like Eigner & Foissner (1994) – that these somewhat set off cirri are the anterior portion of the right marginal row (Fig. 59d). However, ontogenetic data are needed for a correct interpretation. Invariably three dorsal kineties, each with a single caudal cirrus at posterior end; consequently dorsomarginal row and kinety fragmentation lacking. Dorsal bristles likely of ordinary length, that is, around 3 µm (Fig. 59d, g). Occurrence and ecology: Very likely confined to terrestrial habitats. Type locality of H. terricola qingdaoensis is the hill Baguan in/near the city of Qingdao (36°08'N 120°43'E), Shandong, China. Song & Wilbert (1989) discovered it there in soil and cultured it according to the method by Buitkamp & Wilbert (1974). No further records published. Feeds on bacteria, flagellates, diatoms, and small ciliates (Song & Wilbert 1989).
Hemiamphisiella wilberti (Foissner, 1982) Foissner, 1988 (Fig. 60a–g, Table 23) 1982 Strongylidium wilberti nov. spec.1 – Foissner, Arch. Protistenk., 126: 32, Abb. 1a–g, 41, Tabelle 4 (Fig. 60a–g; original description; the holotype slide [accession number 1981/97; see Aescht 2003, p. 400] has been deposited in the Oberösterreichische Landesmuseum [LI], Upper Austria). 1988 Hemiamphisiella wilberti (Foissner, 1982) nov. comb. – Foissner, Stapfia, 17: 122 (combination with Hemiamphisiella). 2001 Hemiamphisiella wilberti (Foissner, 1982) Foissner, 1988 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 83 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs and euplotids).
Nomenclature: Foissner (1982) dedicated this species to Norbert Wilbert (University of Bonn, Germany), who contributed significantly to the alpha-taxonomy of the hypotrichs and euplotids (references, see Berger 2006a). Remarks: This species was originally assigned to Strongylidium Sterki, 1978, mainly because of the slight twisting of the cell and the prominent cirral row(s) between the marginal rows. Foissner (1988) transferred it to Hemiamphisiella because of the resemblance with the type species H. terricola terricola (= Strongylidium muscorum sensu Foissner 1984). It can be easily distinguished from H. terricola by 1
Foissner (1982) provided the following diagnosis: In vivo etwa 220–300 × 50–65 µm großes, leicht kontraktiles, sehr biegsames Strongylidium, dessen posteriorer Körperabschnitt häufig schwanzartig verschmälert ist. 2 Ventralreihen, die rechte beginnt etwa in Körpermitte und endet in der Nähe des posterioren Pols, die linke beginnt in der Höhe des Buccalcirrus und endet bei der Schwanzwurzel. Durchschnittlich 48 adorale Membranellen. 4 Dorsalkineten.
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the nuclear apparatus (8–35 macronuclear nodules in H. terricola vs. 2 in present species). By contrast, Hemiamphisiella granulifera also has only two macronuclear nodules. However, it differs from H. wilberti in several morphometric features (e.g., 23–37 adoral membranelles vs. 42–53; Table 23) and, more importantly, by the presence of cortical granules. Foissner et al. (2002, p. 654; 2005) found H. wilberti and could confirm the lack of cortical granules (unpublished notes kindly supplied by W. Foissner). However, they must not be confused with the sub-cortical mitochondria which are also rather distinct in H. wilberti (Fig. 60b, c). By contrast, Foissner & AL-Rasheid (2006, p. 2, 13, Fig. 38–40) described cortical granules for a population from near the city of Kefermarkt (Upper Austria), indicating that they confused it, likely par lapsus, with H. granulifera. This is also indicated by the relatively low number (likely below 40; see their Fig. 38) of adoral membranelles (vs. 42–53 in H. wilberti). Hemiamphisiella wilberti is very easily confused with Pseudouroleptus caudatus namibiensis Foissner, Agatha & Berger, 2002, which has a similar body shape, size, as well as nuclear pattern and infraciliature (p. 668). However, Pseudouroleptus caudatus namibiensis has distinct cortical granules (see genus section for preliminary, pragmatic separation of Pseudouroleptus and Hemiamphisiella). Morphology: Body size in life about 220–300 × 50–65 µm; body length:width ratio 5.5:1 in protargol preparations (Table 23). Body outline slightly sigmoidal, slender, posterior portion often narrowed tail-like (Fig. 60a, d); rarely pisciform specimens occur (Fig. 60f). Body flattened about 2:1 dorsoventrally; flexible, contracts to about two thirds of ordinary body length under cover-glass pressure. Two macronuclear nodules behind proximal portion of adoral zone, that is, left of midline; nodules about 20 × 11 µm in life, without distinct chromatin bodies. Micronuclei very conspicuously shiny in life, compact, about 10 × 6 µm (Fig. 60a, g). Contractile vacuole distinctly ahead of mid-body near left cell margin, without collecting canals (Fig. 60e). Cortical granules lacking. Pellicle shiny, close underneath numerous colourless, about 2 µm-sized, disk-shaped mitochondria arranged in indistinct rows; due to the close arrangement they become slightly polygonal in outline (Fig. 60b, c). Cytoplasm, especially in posterior body portion, packed with slightly yellowish, bulbous and cylindrical, 2–5 µm-sized crystals, so that cells appear dark at low magnification. Food vacuoles 4–25 µm across, filled with mainly unknown content. Often many about 10 µm-sized vacuoles with several bulbous crystals resembling those contained in the cytoplasm (Fig. 60a, c). Movement slow rather than fast, twitches hastily to and fro. Adoral zone occupies 25% of body length, shaped like a question mark, composed on average of 48 membranelles of ordinary fine structure (Table 23); distal end of zone extends distinctly (to 7.5% of body length in specimen shown in Fig. 60f) onto right body margin (Fig. 60a, f). Bases of largest membranelles about 7 µm wide. Buccal field shallow. Undulating membranes slightly curved, arranged in parallel and more or less of equal length, each likely composed of a double row (zigzag row) of basal bodies. Pharyngeal fibres extend obliquely backwards (Fig. 60a, f).
Hemiamphisiella
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Fig. 60a–g Hemiamphisiella wilberti (from Foissner 1982. a–e, from life; f, g, protargol impregnation). a: Ventral view of a representative specimen, 277 µm. b, c: Subcortical mitochondria in top view and lateral view. d: Left lateral view showing dorsoventral flattening, 243 µm. e: Dorsal view of shape variant. f, g: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 154 µm. Asterisk marks a short, supernumerary cirral row present in only few specimens. Short arrow denotes right ventral row; long arrow marks postperistomial ventral cirrus. Dotted line connects frontal cirri; broken lines connect cirri which originate from the same anlage (only shown for anlagen II and III). ACR = amphisiellid median cirral row, AZM = adoral zone of membranelles, CC = caudal cirri?, CV = contractile vacuole, LMR = left marginal row, MA = macronuclear nodule, MI = micronucleus, PF = pharyngeal fibres, RMR = right marginal row, III/2 = cirrus behind right frontal cirrus, 1–4 = dorsal kineties. Page 311.
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Cirral pattern and number of cirri of usual variability (Fig. 60a, f, Table 23). Cirri about 13 µm long, in posterior portion finer and more widely spaced than in anterior portion. Frontal cirri distinctly enlarged, about 16 µm long, arranged in oblique row with right cirrus behind distal end of adoral zone of membranelles. Buccal cirrus slightly behind anterior end of undulating membranes. Cirrus III/2 behind right frontal cirrus, about at level of buccal cirrus. One postperistomial ventral cirrus near proximal end of adoral zone (Fig. 60f). Amphisiellid median cirral row commences about at level of buccal cirrus, extends spirally to near base of tail. Right ventral row rather long, commences at 42% of body length (Table 23); sometimes an additional row right of this row (Fig. 60f, asterisk). Distinct transverse cirri lacking. Right marginal row commences on dorsal side about at level of right frontal cirrus, runs onto ventral side extending almost longitudinally to near rear end of cell. Left marginal row commences left of proximal portion of adoral zone, extends spirally onto dorsolateral surface posteriorly, ends likely at tip of tail. Dorsal bristles about 4 µm long in life, arranged in four sigmoidal kineties; kineties 1 and 2 almost of body length, kineties 3 and 4 commence subapical and terminate about at tail base (Fig. 60f, g). Caudal cirri likely present. Ontogenetic data are needed to know the exact pattern and origin of dorsal kineties (all kineties bipolar, or dorsomarginal row and/or kinety fragmentation present) and caudal cirri. Occurrence and ecology: Hemiamphisiella wilberti likely prefers terrestrial habitats, although at least two records from limnetic habitats have been published. Found in the Holarctis, the Paleotropis, and the Australis (Foissner 1987, 1998; Foissner et al. 2002, p. 53). Type locality is a floodplain near the village of Grafenwörth, Lower Austria (Foissner 1982). For detailed description of this site, see Foissner et al. (1985, p. 88; site AV, 48°23’N 15°45’E). They also found it in an agricultural soil in the same region, that is, near the village of Frauendorf an der Au (site FA in Foissner et al. 1985). Foissner et al. (2005, p. 628) found it in a floodplain soil (calcaric fluvisol; typical mull; recent clay) of a Pruno-Fraxinetum community in Eastern Austria (site Müllerboden; 48°00'N 16°42'E; altitude 160 m). Further terrestrial records: agricultural soils from the Ostrov region (48°38'N 17°46'E), Slovakia (Tirjaková 1988, p. 500); microbiotic crusts containing cyanobacteria and lichens from desert soils in the Grand Canyon in northern Arizona, USA (Bamforth 2004, p. 417). Limnetic records not substantiated by morphological data: sediment of the Guadarrama river in the region of Villalba near Madrid, Spain (Fernandez-Leborans et al. 1990, p. 513); mineral and thermo-mineral water in the Auvergne, France (Chaouite et al. 1990). Food not known in detail, occasionally testate amoebae (Trinema lineare) are ingested. Biomass of 106 specimens about 152 mg (Foissner 1987, p. 127; 1998, p. 204).
Hemiamphisiella
315
Hemiamphisiella granulifera (Foissner, 1987) Foissner, 1988 (Fig. 61a–e, Table 23) 1987 Strongylidium granuliferum nov. spec.1 – Foissner, Zool. Beitr., 31: 190, Abb. 1a–e, Tabelle 1 (Fig. 61a–e; original description; the holotype slide [accession number 188/130; Aescht 2003, p. 387] has been deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria). 1988 Hemiamphisiella granulifera (Foissner, 1987) nov. comb. – Foissner, Stapfia, 17: 122 (combination with Hemiamphisiella). 2001 Hemiamphisiella granulifera (Foissner, 1987) Foissner, 1988 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 82 (nomenclator containing all basionyms, combinations, and higher taxa of euplotids and hypotrichs).
Nomenclature: No derivation of the name is given in the original description. The species-group name granulifer·a, -a, -um (Latin adjective [m; f; n]; bearing granules) is a composite of granulum (Latin noun; granule), the thematic vowel ·i-, and fero (Latin verb; bear) and refers to the conspicuous cortical granules. Strongylidium has neuter gender, whereas Hemiamphisiella is feminine. Thus, the speciesgroup name granuliferum was correctly changed to granulifera by Foissner (1988). Remarks: This species is very similar to H. wilberti, which has the same nuclear apparatus. However, they differ in some morphometric features (Table 23) and, more importantly, in the cortical granulation, which is lacking in H. wilberti. Hemiamphisiella terricola (type species) also has cortical granules; however, it has not two, but many macronuclear nodules. Hemiamphisiella wilberti sensu Foissner & AL-Rasheid (2006, see their legend to Fig. 40) has cortical granules indicating that they confused H. granulifera and H. wilberti. Morphology: The following paragraphs are based on the original description (Foissner 1987a). Some unpublished data, kindly provided by W. Foissner, from populations from Kenya, USA, and Austria are added below. Body size in life about 140–200 × 25–40 µm, body length:width ratio of silverprepared specimens 5.5:1 on average (Table 23). Body outline slender, usually slightly sigmoidal, rear portion narrowed tail-like (Fig. 61a, d, e). Body somewhat flattened dorsoventrally, flexible. Macronuclear nodules slightly left of midline, contain many, moderately large chromatin bodies; distance between macronuclear nodules 4–14 µm, on average 8.3 µm (n = 11). Micronuclei globular, closely attached to macronuclear nodules. Contractile vacuole slightly ahead of mid-body near left cell margin. Cortical granules colourless and globular. Pellicle colourless, flexible; after protargol impregnation with many small granules, which are very likely the cortical granules (Fig. 61b, c). Movement not described, likely without peculiarities. Adoral zone occupies 24% of body length on average (Table 23), composed of around 29 membranelles. Buccal field rather deep, moderately wide. Undulating 1
Foissner (1987a) provided the following diagnosis: In vivo etwa 140–200 × 25–40 µm großes Strongylidium mit farblosen kugelförmigen subpelliculären Granula, 2 Makronucleus-Teilen und sehr stark verkürzter rechter Ventralreihe. Durchschnittlich 29 adorale Membranellen. 3 Dorsalkineten.
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membranes distinctly curved and arranged almost in parallel, about of equal length. Pharyngeal fibres extend obliquely backwards (Fig. 61a, d). Cirral pattern and number of cirri of usual variability (Fig. 61a, d, Table 23). Three distinctly enlarged frontal cirri. Buccal cirrus right of anterior portion of paroral. One cirrus (= cirrus III/2) behind right frontal cirrus. One postperistomial cirrus slightly behind proximal end of adoral zone. Amphisiellid median cirral row commences right of right frontal cirrus, extends sigmoidally to 79% of body length on average (Table 23). Right ventral row short, that is, composed of about three cirri almost in line and only indistinctly separated from rear end of amphisiellid median cirral row (Fig. 61d, arrow). Transverse cirri lacking. Right marginal row commences near anterior body end on dorsal side, extending along body margin from level of buccal vertex to end of cell. Left marginal row slightly curved rightwards anteriorly, that is, commences near proximal end of adoral zone, extends along left cell margin, terminates at rear body end. Dorsal bristles arranged in three sigmoidal kineties; all kineties rather distinctly shortened anteriorly: kinety 1 slightly more shortened anteriorly than kinety 2, and kinety 2 slightly shorter than kinety 3. Length of dorsal bristles not mentioned, according to Fig. 61e they are about 3 µm long. Probably two1 caudal cirri, which are, however, difficult to separate from rear end of left marginal row (thus they have been counted as left marginal cirri; Table 23). Additional and deviating data from the Kenyan population (unpublished, kindly supplied by W. Foissner; Table 23): Body size in life about 110 × 40 µm; body rather fragile, about 2:1 flattened dorsoventrally; macronuclear nodules about 22 × 13 µm in life; micronuclei about 4 µm across in life; cortical granules conspicuous, globular, about 1 µm across, arranged in more or less distinct longitudinal rows all over the cell; cytoplasm with few greasily shining globules about 2 µm across and, mainly in posterior portion, yellowish crystals 2–5 µm long; movement moderately fast; bases of adoral membranelles up to 6 µm wide; buccal field inconspicuous; very likely three caudal cirri. W. Foissner found H. granulifera also in eastern Austria (Burgenland) and in the USA (unpublished data kindly supplied). In the Austrian population, the colourless, globular cortical granules were about 1.5–2.0 µm across and densely arranged in longitudinal rows. In the population (body size in life 160–180 × 40–50 µm; dorsal bristles 3–4 µm long) from the USA the granules are 1.0–1.2 µm across. Occurrence and ecology: Hemiamphisiella granulifera is very likely confined to terrestrial habitats (Foissner 1987, p. 129; 1998, p. 203). Type locality is the Cape Garajan on the island of Madeira, Atlantic Ocean, where Foissner (1987a) discovered it with very low abundance in grassland soil grown with Opuntia (altitude 150 m; pH 4.8; coll. Wolfgang Petz). Further records: upper soil layer of a forest stand (Pruno-Fraxinetum) in eastern Austria (Foissner et al. 2005, p. 619); Burgenland, Austria (W. Foissner, unpublished); desert soil from near the city of Las Vegas, 1
This number must not be over-interpreted because the cirral pattern on the tip of the tail is very difficult to study (see next paragraph).
Hemiamphisiella
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Fig. 61a–e Hemiamphisiella granulifera (from Foissner 1987a. a, c, from life; b, d, e, protargol impregnation). a: Ventral view of representative specimen, 187 µm. b, c: Part of pellicle showing cirri and cortical granules. d, e: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 120 µm. Long arrow marks postperistomial ventral cirrus, short arrow denotes right ventral row. Broken lines connect cirri which originate from the same anlage (only shown for anlagen I–III); the cirrus behind the right frontal cirrus is cirrus III/2. Dotted line connects frontal cirri. ACR = amphisiellid median cirral row, AZM = adoral zone of membranelles, CC = caudal cirri (difficult to recognise because of tailed body end), CV = contractile vacuole, FC = right frontal cirrus, LMR = left marginal row (anterior portion more or less distinctly curved rightwards in Hemiamphisiella), MA = macronuclear nodule, MI = micronucleus, P = paroral, RMR = right marginal row, III/2 = cirrus III/2 (= cirrus behind right frontal cirrus = paramalar cirrus), 1–3 = dorsal kineties. Page 315.
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USA (W. Foissner, unpublished); upper soil layer (0–5 cm; pH 6.0; black, very wet and sandy) with few roots and litter from marshy area near a forest surrounding the Sheldrick waterfalls, Shimba Hills Nature Reserve, Kenya (Foissner 1999, p. 323; see description and Table 23 for some unpublished data of this population); in three out of 73 soil samples from Namibia (Foissner et al. 2002, p. 60). Feeds on heterotrophic flagellates, cysts of amoebas, and fungal spores (Foissner 1987a), but also on ciliates like Leptopharynx costatus (W. Foissner, unpublished; Kenya population) and Colpoda inflata (W. Foissner, unpublished; USA population). Biomass of 106 specimens about 52 mg (Foissner 1998, p. 203).
Incertae sedis in Hemiamphisiella Hemiamphisiella quadrinucleata (Foissner, 1984) Foissner, 1988 (Fig. 62a–e, Tables 23, 35) 1984 Uroleptoides quadrinucleata nov. spec.1 – Foissner, Stapfia, 12: 114, Abb. 60a–e, Tabelle 28 (Fig. 62a–e; original description; according to Aescht 2003, p. 395, two slides with syntypes [accession numbers 1982/59, 1984/90] have been deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; see nomenclature). 1988 Uroleptoides quadrinucleatus nom. corr. – Foissner, Stapfia, 17: 122 (see nomenclature). 1988 Hemiamphisiella quadrinucleata (Foissner, 1984) nov. comb. – Foissner, Stapfia, 17: 122 (combination with Hemiamphisiella). 2001 Hemiamphisiella quadrinucleata (Foissner, 1984) Foissner, 1988 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 97 (nomenclator containing all basionyms, combinations, and higher taxa of euplotids and hypotrichs).
Nomenclature: No derivation of the name is given in the original description. The species-group name quadrinucleata (Latin adjective) is a composite of the Latin Fig. 62a–e Hemiamphisiella quadrinucleata (from Foissner 1984. a–c, from life; d, e, protargol impregnation). a: Ventral view of a representative specimen, 90 µm. b: Dorsal view showing cortical granulation, contractile vacuole, and defecation. c: Right lateral view showing dorsoventral flattening. d, e: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 67 µm. Long arrow marks cirrus at proximal end of adoral zone. Whether or not this cirrus is homologous to the postperistomial ventral cirrus of the other Hemiamphisiella species is not yet known. It cannot be excluded that anlage IV is lacking in the present species, as, for example, in Nudiamphisiella interrupta. If this is the case, then the cirrus marked by the long arrow very likely originates from anlage VI. Short arrow marks indistinct break in amphisiellid median cirral row. Broken lines connect cirri which originate from same anlage; for the anlagen IV to VI this is only an assumption because ontogenetic data are needed to show the true origin of the various cirral groups. ACR = rear end of amphisiellid median cirral row, AZM = adoral zone of membranelles, CC = caudal cirri, CV = contractile vacuole, MA = macronuclear nodule, MI = micronucleus, P = paroral, RMR = anterior end of right marginal row, I–VI = frontal-ventral cirral anlagen, 1–4 = dorsal kineties (4 = dorsomarginal kinety?). Page 318. 1
Foissner (1984) provided the following diagnosis: In vivo 70–100 × 25–35 µm große, ellipsoide Uroleptoides mit meist 4 Makronucleus-Teilen und zahlreichen farblosen subpelliculären Granula, die in Längsreihen angeordnet sind. 3 Caudalcirren, 4 Dorsalkineten, von denen die rechte stark verkürzt ist.
d
Hemiamphisiella
319
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SYSTEMATIC SECTION
quantifier quadr·us (square, four) and the Latin adjective nucleat·us, -a, -um ([m; f; n]; kernel-like), and refers to the four macronuclear nodules. Foissner (1984) classified the present species in Uroleptoides, which is masculine. Thus, Foissner (1988) introduced the nomen corrigendum Uroleptoides quadrinucleatus. Foissner (1984, p. 8) deposited 1–3 holotype slides for each new species described. According to Aescht (2003), the two slides deposited (see list of synonyms) are syntypes, likely because no specific specimen of the type series was fixed as holotype specimen (ICZN 1999, Article 73.2). Remarks: Foissner (1984) assigned this species to the controversial genus Uroleptoides because it has caudal cirri, whereas such cirri are lacking in Amphisiella, the second genus taken into consideration by Foissner (1984). Later, when he established Hemiamphisiella with H. terricola as type species, he transferred it to this group, which is characterised, inter alia, by the presence of a postperistomial cirrus, a feature also present in U. quadrinucleatus. However, some characters – for example, body shape, anterior end of left marginal row, lack of right ventral row, presence of a dorsomarginal kinety (= kinety 4 in Fig. 62e) – indicate that the classification in Hemiamphisiella is incorrect. Possibly, it is related to Nudiamphisiella (further details see genus section). Ontogenetic studies and meaningful molecular data are needed for a more proper classification. Lamtostyla vitiphila has the same habitus including the nuclear apparatus (Fig. 34a–d). However, this species lacks caudal cirri, the postperistomial cirrus, the dorsomarginal kinety, and cortical granules. Morphology: Body size in life 70–100 × 25–35 µm; body length:width ratio 2.7:1 on average in protargol preparations (Fig. 62a, b, d, Table 23). Body outline rather constant, elliptical, right margin slightly concave, left slightly convex, anterior and posterior portion somewhat converging and rounded (Fig. 62a, b). Body flattened about 2:1 dorsoventrally, ventral side slightly, dorsal side moderately vaulted (Fig. 62c); slightly contractile under coverglass pressure. Usually four macronuclear nodules form two longitudinally arranged pairs in left body portion; nodules with many small chromatin bodies. One micronucleus attached to each pair of macronuclear nodules, either in between or laterally (Fig. 62a, e). Contractile vacuole about in mid-body at left cell margin, during diastole without collecting canals. Cytopyge terminal, right of body-midline; faecal balls loose (Fig. 62b). Pellicle colourless. Cortical granules (mucocysts?) arranged in longitudinal rows, colourless, about 0.5 µm across, stain red when methyl green-pyronin is added (Fig. 62b). Cytoplasm with some yellowish crystals and food vacuoles 5–10 µm across. Glides slowly among soil particles showing great flexibility. Adoral zone occupies 30% of body length on average (Fig. 62a, d, Table 23), composed of an average of 21 membranelles. Buccal field flat and small. Undulating membranes slightly curved, paroral commences distinctly ahead of endoral. Pharyngeal fibres extend obliquely backwards (Fig. 62a, d). Cirral pattern and number of cirri of usual variability (Fig. 62a, d, Table 23). Frontal cirri distinctly enlarged, arranged in slightly oblique row. Buccal cirrus right
Hemiamphisiella
321
Fig. 63a Hemiamphisiella terricola (from Alekperov 2005. Wet silver nitrat impregnation). Infraciliature of ventral side and nuclear apparatus, 195 µm. Short arrow marks buccal cirrus, long arrow denotes right ventral row. Details about misidentification see text. ACR = amphisiellid median cirral row. Page 322.
of anterior end of paroral, lacking in one out of 15 specimens. One (rarely two) cirrus (= cirrus III/2) behind right frontal cirrus. Amphisiellid median cirral row commences near right frontal cirrus, terminates at 68% of body length on average (Fig. 62d, Table 23); specimen illustrated with inconspicuous break about at level of buccal vertex, indicating that the row is formed at least from cirri of the anlagen V (posterior portion) and VI (anterior portion). Morphogenetic data are needed to know whether the middle portion is formed by the anlage IV, as in H. terricola, or in a different way. At proximal end of adoral zone a single cirrus, possibly homologous to the postperistomial cirrus (= cirrus IV/2) of the other Hemiamphisiella species. However, it cannot be excluded that it is the rearmost cirrus of the anterior portion of the amphisiellid median cirral row. Right ventral row and transverse cirri lacking. Right marginal row commences slightly behind level of frontal cirri. Anterior end of left marginal row straight, that is, not curved rightwards as in the other Hemiamphisiella species! Marginal rows only inconspicuously separated posteriorly; gap occupied by caudal cirri; bases of cirri become smaller in posteriad direction. Length of bristles not measured; according to illustration short, that is, about 2–4 µm (Fig. 62e). Dorsal bristles arranged in four kineties; kinety 1 slightly shortened anteriorly, kineties 2 and 3 more or less bipolar, kinety 4 terminates at 23% of body length in specimen illustrated, strongly indicating (basically proving) that it is a dorsomarginal row (such a row is also present in Nudiamphisiella [p. 560] and the other Dorsomarginalia Berger, 2006; see remarks). Usually three, rarely four fine caudal cirri (Fig. 62e); the presence of three caudal cirri also indicates that kinety 4 is a dorsomarginal row because such rows do not form a caudal cirrus. Occurrence and ecology: Hemiamphisiella quadrinucleata is likely confined to terrestrial habitats (Foissner 1987, 1998). Type locality is the Salesen Alm (altitude 1810 m; sample collected on 1982.08.11; Foissner 1984, p. 7), an alpine hut at the Stubnerkogel near the village of Bad Gastein, Austria, where Foissner discovered it in the upper soil layer (0–5 cm) of a pasture with young green alder (Alnus viridis). For autecological data on this population, see Foissner & Peer (1985, p. 44). Also
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recorded during experiments on soil compaction in an alpine pasture (Schlossalm area, Bad Hofgastein) near the type locality (Berger 1985, Berger et al. 1985, p. 107). Not recorded during a detailed study of Namibian soils (Foissner et al. 2002). Feeds on heterotrophic flagellates (Chilomonas sp.), green algae, and fungi (Foissner 1984). Biomass of 106 specimens about 42 mg (Foissner 1987, p. 128; 1998, p. 204).
Insufficient redescription Hemiamphisiella terricola Foissner, 1988 – Alekperov, 2005, Atlas of free-living ciliates, p. 217, Fig. 68.4 (Fig. 63a). Remarks: The description of this population is in Russian and was not translated in detail. However, the illustrations shows at least two significant differences to the type population and the other populations of H. terricola, namely, (i) the lack of the postperistomial ventral cirrus, a main feature of Hemiamphisiella; and (ii) the buccal cirrus is distinctly behind the anterior end of the undulating membranes whereas it is near the anterior end in H. terricola (Fig. 56e, 57d, 58a, 59c). These two features are reminiscent of Paramphisiella acuta (Fig. 70d). However, Alekperov (2005) also illustrated a right ventral row as is characteristic for Hemiamphisiella. Thus, it is very unlikely that the present population is P. acuta. Body size 190–250 × 50 µm (in silver preparations?); 42–45 adoral membranelles; 25–35 macronuclear nodules. Further details, see Fig. 63a. Found in soil(?) in Azerbaijan.
Group III: Terrestrial Amphisiellids which lack Frontal-ventraltransverse Cirri Anlage IV This group comprises two genera (Lamtostylides, Paramphisiella) which have obviously lost cirral anlage IV. Thus, they usually have only one cirrus (= cirrus III/2) left of the anterior portion of the amphisiellid median cirral row. Whether this loss is an apomorphy for a group comprising Lamtostylides and Paramphisiella, or occurred independently in both genera is not known. Thus, I do not introduce a name. Basically all species included live almost exclusively in terrestrial habitats.
Lamtostylides gen. nov. Nomenclature: Lamtostylides is a composite of Lamtostyla (see there for derivation), and the suffix -ides (similar, especially in shape; in present case because of similar cirral pattern), indicating a similarity with Lamtostyla. According to Article 30.1.4.4 of the ICZN (1999), a compound genus-group name ending with -ides is to be treated as masculine.
Lamtostylides
323
Characterisation (A = supposed apomorphy): Amphisiellidae with continuous adoral zone of membranelles. Undulating membranes rather short, roughly straight and in parallel. Three more or less distinctly enlarged frontal cirri. One buccal cirrus. One cirrus left of anterior portion of amphisiellid median cirral row because cirral anlage IV lacking (A?). Amphisiellid median cirral row usually terminates ahead of mid-body, originates from anlage V (posterior portion) and VI (anterior portion). Postperistomial cirrus lacking. Pretransverse ventral cirrus(i) usually present. Transverse cirri present. One right and one left marginal row. Two or three dorsal kineties. Caudal cirri lacking. Oral primordium originates apokinetally, that is, without contact to parental cirri. Terrestrial. Additional characters: body length usually below 110 µm; body flexible and elongate elliptical; two macronuclear nodules; contractile vacuole at about 50% of body length or ahead of it, near left body portion or slightly displaced inwards; dorsal bristles less than 5 µm long. Type species: Lamtostyla edaphoni Berger & Foissner, 1987. Remarks: Lamtostyla comprised species which form the frontal-ventraltransverse cirri from six anlagen (I–VI) or only from five (I–III, V, VI, that is, anlage IV is lacking). Although the ontogenesis of Lamtostyla lamottei, type of Lamtostyla, is not known, one can conclude from the cirral pattern that this species also has six anlagen (Fig. 30a). Species with six anlagen have more than one cirrus left of the anterior portion of the amphisiellid median cirral row. By contrast, three Lamtostyla species have only one cirrus left of the anterior portion of the row due to the lack of anlage IV. Lamtostylides is established for these species. I designate L. edaphoni as type because two populations have been studied in detail showing that the single cirrus left of the amphisiellid median cirral row is constant; in addition, its cell division is described in detail (Petz & Foissner 1996). When I reviewed Hemisincirra I recognised the similarity of Lamtostylides halophilus and Hemisincirra pori, which I therefore also include in the present genus. The cirral pattern of Lamtostyla hyalina is difficult to interpret, although it has, like the other three species, only one cirrus left of the anterior end of the row. However, it is not known whether this is cirrus III/2 as in the other species, or the somewhat posteriorly displaced right frontal cirrus. Further, it usually has six transverse cirri, including one small pretransverse ventral cirrus. However, five transverse cirri usually require six anlagen. Ontogenetic data are needed for a correct interpretation. In spite of that Lamtostyla hyalina is preliminary also transferred to Lamtostylides. Interestingly, all species included have two macronuclear nodules, which is, however, an old feature (Berger 2006). For comparison with Terricirra, see there. Afroamphisiella (p. 371) lacks transverse cirri and can thus be easily distinguished from Lamtostylides and Lamtostyla (p. 161). Species included in Lamtostylides (alphabetically arranged basionyms are given): (1) Lamtostyla edaphoni Berger & Foissner, 1987; (2) Lamtostyla halophila Foissner, Agatha & Berger, 2002; (3) Lamtostyla kirkeniensis Berger & Foissner,
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1988; (4) Perisincirra pori Wilbert & Kahan, 1986; (5) Tachysoma hyalina Berger, Foissner & Adam, 1984.
Key to Lamtostylides species The separation of the species included is not quite simple and requires very detailed live observation (interference contrast) and in some cases even protargol impregnation. 1 Body length about 50 µm (Fig. 69a–d). . . . . . . . . Lamtostylides hyalinus (p. 347) - Body length usually more than 50 µm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2 Cortical granules present, however, difficult to recognise because colourless and only 0.5–1.0 µm across (Fig. 66a–f). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lamtostylides halophilus (p. 336; see also L. pori) - Cortical granules lacking. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3 Inhabits saline soil (Fig. 68a, b). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lamtostylides pori (p. 344; see also L. halophilus) - Inhabits non-saline soil (Fig. 64a–h, 65a–e). . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4 13–27 right marginal cirri; 3 dorsal kineties (Fig. 64a–h, Table 22). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lamtostylides edaphoni (p. 324) - 38–46 right marginal cirri; 2 dorsal kineties (Fig. 65a–e, Table 22). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lamtostylides kirkeniensis (p. 333)
Lamtostylides edaphoni (Berger & Foissner, 1987) comb. nov. (Fig. 64a–u, Table 22) 1987 Lamtostyla edaphoni nov. spec.1 – Berger & Foissner, Zool. Jb. Syst., 114: 215, Fig. 56–60, Table 9 (Fig. 64a–e; original description; the holotype slide [accession number 1986/81; Aescht 2003, p. 384] and a paratype slide [1986/82] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria). 1988 Lamtostyla edaphoni Berger and Foissner, 1987 – Berger & Foissner, Zool. Anz., 220: 124, Fig. 19–23, Table 1 (Fig. 64a–e; revision of Lamtostyla). 1996 Lamtostyla edaphoni Berger and Foissner, 1987 – Petz & Foissner, Acta Protozool., 35: 258, Fig. 1–16, Table 1 (Fig. 64f–u; redescription and morphogenesis; voucher slides are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria). 1999 Amphisiella edaphoni (Berger and Foissner, 1987) nov. comb. – Eigner, Europ. J. Protistol., 35: 44 (combination with Amphisiella). 2001 Lamtostyla edaphoni Berger and Foissner, 1987 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 45 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Lamtostyla edaphoni – Lynn & Small, Phylum Ciliophora, p. 459, Fig. 57B, C (Fig. 64d, e; guide to ciliate genera). 1
Berger & Foissner (1987) provided the following diagnosis: In vivo about 70–85 × 20–30 µm (n = 4), long ellipsoid, c. 7–9 cirri in the frontal row. 1 cirrus left of the frontal row at about the level of the buccal cirrus. About 17 adoral membranelles.
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Nomenclature: No derivation of the name is given in the original description. The species-group name edaphoni (from edaphon; Greek noun; the soil flora and fauna) refers to the habitat (soil) where the species was discovered. “Lamtostyla edaphoni Berger & Foissner” in Lüftenegger et al. (1986, p. 152) is a nomen nudum because published without description (ICZN 1999, Article 13). Type species of Lamtostylides. Remarks: For a foundation of the transfer from Lamtostyla to Lamtostylides, see genus section remarks of both taxa. Lamtostylides edaphoni was discovered in Europe (Austria) and later recorded from the Antarctica and Africa. The Antarctic populations have been studied in detail (Petz & Foissner 1996), and the following differences to the type population have been found: (i) number of cirri forming amphisiellid median cirral row (9.5 vs. 8.0); (ii) size (mainly length) of macronuclear nodules (16–17 µm vs. 9–10 µm); (iii) position of contractile vacuole (left margin vs. displaced inwards). The differences are inconspicuous, that is, the conspecificity of the Austrian and Antarctic populations is beyond reasonable doubt. This is sustained by the significantly changed numbers of marginal cirri in specimens from pure cultures (Table 22). The very similar Lamtostylides kirkeniensis can be separated from L. edaphoni by having twice the number of marginal cirri (Table 22) and a lower number of dorsal kineties (2 vs. 3). Lamtostyla islandica has, inter alia, three cirri left of the anterior portion of the amphisiellid median cirral row (vs. one in present species). Morphology: The type population is described first, followed by supplementary observations by Petz & Foissner (1996). Body size of type population 70–85 × 20–30 µm (n = 4) in life; body length:width ratio in protargol preparations 3.8:1 on average (Table 22). Body outline long elliptical, that is, margins nearly parallel, both ends rounded (Fig. 64a). Body flattened about 2:1 dorsoventrally. Macronuclear nodules ellipsoidal with large chromatin bodies, lying slightly left of midline. Each nodule usually with one micronucleus, anterior nodule rarely with two. Contractile vacuole about in midbody, conspicuously displaced inwards, during diastole without collecting canals (Fig. 64b). Cortical granules lacking. Cytoplasm colourless, sometimes with some 1–3 µm large globules and many 4–10 µm large food vacuoles with bacteria and unidentifiable content. Movement rapid. Adoral zone occupies about 30% of body length, composed of an average of 17 membranelles of usual fine structure; bases of largest membranelles in life about 5 µm wide. Buccal area flat and narrow, undulating membranes almost parallel, slightly bent; paroral commences more anteriorly than endoral (Fig. 64a, d; according to Petz & Foissner 1996, who studied ontogenesis, the anterior membrane is the paroral. Berger & Foissner 1987, 1988 did not label the membranes because we were uncertain about the correct designation). Pharyngeal fibres recognisable in life and in protargol preparations; extend obliquely backwards.
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Fig. 64a–e Lamtostylides edaphoni (from Berger & Foissner 1987. a–c, from life; d, e, protargol impregnation). a: Ventral view of a representative specimen, 81 µm. b, c: Dorsal view showing adoral zone and contractile vacuole and right lateral view. d, e: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 68 µm. Arrow marks cirrus (= cirrus III/2) left of amphisiellid median cirral row. Arrowhead denotes inconspicuous break in amphisiellid median cirral row. ACR = amphisiellid median cirral row, AZM = adoral zone of membranelles, LMR = left marginal row, P = paroral (see description), RMR = right marginal row, TC = transverse cirri, 1–3 = dorsal kineties. Page 324.
Cirral pattern as shown in Fig. 64a, d, variability of number of cirri and arrangement relatively low (Table 22). Frontal cirri almost transversely arranged, slightly enlarged. Buccal cirrus right of anterior portion of paroral and ahead of anterior end of endoral. One cirrus (= cirrus III/2) left of amphisiellid median cirral row, usually slightly behind level of buccal cirrus. Amphisiellid median cirral row commences, as is usual, near right frontal cirrus, respectively, distal end of adoral zone, composed of eight cirri on average; row terminates at 31 % of body length on average, that is, slightly behind level of proximal end of adoral zone; sometimes with an indistinct discontinuity in middle portion (= site where the two portions of the row meet; see cell division). Transverse cirri not enlarged, in life about 15 µm long, distinctly pro-
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Fig. 64f–i Lamtostylides edaphoni (from Petz & Foissner 1996. f, from life; g–i, protargol impregnation). f: Ventral view of a representative specimen, 82 µm. g, h: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 56 µm. i: Ventral view of specimen with surplus (arrow) cirral row right of amphisiellid median cirral row, 78 µm. ACR = amphisiellid median cirral row, BC = buccal cirrus, E = endoral, P = paroral, 1–3 = dorsal kineties. Page 324.
truding beyond rear body margin. Right marginal row distinctly shortened anteriorly, that is, usually commences behind level of buccal cirrus; ends, like left row, at or ahead of level of transverse cirri; left row commences at level of buccal vertex. Dorsal cilia about 3 µm in life, arranged in three kineties (Fig. 64e). Kinety 1 distinctly, kinety 2 moderately shortened anteriorly; bristles within kineties rather widely spaced. Caudal cirri lacking. Supplementary data from Antarctic population I (Petz & Foissner 1996; morphometric data of all Antarctic populations see Table 22): Body size in vivo 55–80 (rarely up to 120) × 18–25 µm in life; body flexible, usually rather transparent at low magnification. Macronuclear nodules 15–18 × 7–8 µm in life. 1–4 micronuclei, 2.5–3.0 µm across in life, in variable position attached to macronuclear nodules.
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Contractile vacuole sometimes slightly ahead of mid-body, close to left cell margin. Food vacuoles with dark and bright green debris. Crawls moderately rapidly, quickly going back and forth; thigmotactic, that is, not easy to take with pipette. Cilia of adoral membranelles up to about 16 µm long. Each undulating membrane possibly composed of single basal body row, cilia in life 5–6 µm long. Pharyngeal fibres 9–12 µm long in protargol-impregnated specimens, extend obliquely backwards. Frontal cirri composed of 2–3, all other cirri of two basal body rows; however, bases of frontal cirri, buccal cirrus, and cirrus III/2 slightly elongated and thus also larger. Very rarely (in two out of 70 specimens), a short second row of 3–4 cirri left of posterior portion of amphisiellid median cirral row, possibly remnants from last division (Fig. 64i, arrow). Number of marginal cirri distinctly higher in cultured specimens, but other characters similar to field material (Table 22). Cell division (Fig. 64j–u): Morphogenesis of L. edaphoni is described in great detail by Petz & Foissner (1996). As in other small species, the process is very difficult to study because all cirri and primordia are very close together. Thus, they could not clarify the origin of some anlagen unambiguously. Petz & Foissner (1996) designated the five frontal-ventral-transverse cirri anlagen with 1 to 5. The morphogenesis shows that in the present species (and in other species with only one cirrus left of the anterior portion of the amphisiellid median cirral row) anlage IV is lacking. Thus, Lamtostylides edaphoni forms the cirri from the anlagen I, II, III, V, and VI. Stomatogenesis begins with the proliferation of basal bodies close ahead of the left transverse cirrus (Fig. 64j). The primordium then elongates anteriorly to the parental adoral zone of membranelles. At the same time, the rearmost cirrus of the amphisiellid median cirral row disintegrates and a small basal body patch is formed posteriorly (Fig. 64k). It is uncertain whether this patch originates from a cirrus of the amphisiellid row or originates de novo. Moreover, Petz & Foissner (996) could not clarify whether this patch is incorporated into the oral primordium or remains independent, forming only cirral streaks (Fig. 64k, l). The same uncertainty exists in Lamtostyla perisincirra (Berger et al. 1984) and Lamtostyla australis (Voß 1992). Afterwards, the buccal cirrus and cirrus III/2 are modified to proter’s anlagen II and III, respectively (Fig. 64l, m). At the same time, some further cirri disintegrate at the rear end of the amphisiellid median cirral row, forming the rightmost anlage (= anlage VI) of the opisthe. This anlage terminates near the oral primordium, respectively, the basal body patch mentioned above (Fig. 64l, m). A further cirrus of the amphisiellid row then dedifferentiates ahead of the rightmost anlage, forming
b
Fig. 64j–o Lamtostylides edaphoni (from Petz & Foissner 1996. Protargol impregnation). j: Infraciliature of ventral side of very early divider (93 µm) showing origin of oral primordium (arrows). k, l: Infraciliature of ventral side of early dividers (k = 87 µm, l = 90 µm) showing basal body patch (k, long arrow) right of oral primordium. Short arrows denote disintegrating rearmost cirrus of amphisiellid median cirral row (k) and buccal cirrus (l); arrowhead in (l) denotes disintegrating cirrus III/2. m, n: Infraciliature of ventral side of early dividers (m = 83 µm, n = 73 µm) showing organisation of cirral anlagen. Arrows mark first sign of proter’s anlage V (m; designated as anlage 4 by Petz & Foissner) and VI (n). o: Infraciliature of dorsal side of specimen shown in (n). TC = transverse cirri. Page 324.
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proter’s anlage V (Fig. 64m). The oral primordium produces membranelles from right to left in a posteriad direction (Fig. 64m). On the right side, basal bodies align to the anlage I (undulating membranes primordium) and also form the streaks II and III of the opisthe; sometimes some of the primordia contact the posterior end of the proter’s corresponding primordia (Fig. 64m, n, p; Petz & Foissner 1996). The origin of opisthe’s anlage V is uncertain; probably, it originates from the oral primordium as stated in Lamtostyla australis by Voß (1992). Subsequently, the old paroral disintegrates and the endoral probably becomes double-rowed, possibly incorporating basal bodies from the dedifferentiating paroral, which appears as a loose row extending to the oral primordium of the opisthe. Finally, a distinct, two-rowed anlage is recognisable. As is usual, the anterior part of the anlage becomes the leftmost frontal cirrus (= cirrus I/1), while the rear portion forms the new undulating membranes (Fig. 64n, p, q). Anlage VI of the proter originates ahead of anlage V, probably from a disintegrating cirrus of the old amphisiellid median cirral row (Fig. 64n, p). Later, the primordia elongate by basal body proliferation and five anlagen each are recognisable in both dividers (Fig. 64p). As is usual, the cirri are formed in a posteriad direction as follows: anlage I – left frontal cirrus and undulating membranes; anlage II – middle frontal cirrus, buccal cirrus, and sometimes also a transverse cirrus (when four transverse cirri are present; Fig. 64q); anlage III – right frontal cirrus, cirrus III/2 (= cirrus left of anterior portion of amphisiellid median cirral row; rarely two cirri are formed), transverse cirrus; anlage IV – lacking in this species and genus; anlage V – posterior portion of amphisiellid median cirral row and transverse cirrus; anlage VI – anterior portion of amphisiellid median cirral row and rightmost transverse cirrus. The cirri of anlage V migrate backwards and align behind streak VI, both forming the new amphisiellid median cirral row, when the new transverse cirri organise (Fig. 64q, r, t). Surplus basal body pairs in the anlagen are likely resorbed, as are all parental cirri, which do not contribute to anlagen formation. However, it cannot be excluded that some cirri at the front end of the parental amphisiellid median cirral row remain because these are always very close to anlage VI (Fig. 64q, r, t). The parental adoral zone is apparently not renewed, but the pharyngeal fibres are resorbed and rebuilt during cytokinesis and in postdividers (Petz & Foissner 1996). Marginal row and dorsal kinety formation proceeds in the usual, plesiomorphic mode, that is, most marginal cirri dissolve and two separate anlagen are formed per
b
Fig. 64p–s Lamtostylides edaphoni (from Petz & Foissner 1996. Protargol impregnation). p: Infraciliature of ventral side of middle divider, 66 µm. Both proter and opisthe show five frontal-ventral-transverse cirral anlagen. Note that L. edaphoni lacks anlage IV; thus, only one cirrus (= cirrus III/2) is present left of anterior portion of amphisiellid median cirral row. q: Infraciliature of ventral side of middle divider showing segregation of cirri and development of marginal primordia, 86 µm. r, s: Infraciliature of ventral and dorsal side and nuclear apparatus of a late divider, 68 µm. As in most other amphisiellids, the amphisiellid median cirral row is composed of cirri of anlage V (posterior portion) and VI (anterior portion; corresponds the frontoterminal cirri of the oxytrichids and urostyloids). I–III, V, VI = frontal-ventral-transverse cirri anlagen. Page 324.
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Fig. 64t, u Lamtostylides edaphoni (from Petz & Foissner 1996. Protargol impregnation). Infraciliature of ventral and dorsal side and nuclear apparatus of a very late divider, 96 µm. Frontal-ventral-transverse cirri of proter which originate from same anlage, are connected by broken lines (corresponding transverse cirri not included). ACR = amphisiellid median cirral row, I–III, V, VI = frontal-ventral-transverse cirri anlagen, 1–3 = dorsal kineties of proter. Page 324.
parental row (Fig. 64o, q–u). The dorsal kineties develop only by intrakinetal proliferation, that is, fragmenting kineties and dorsomarginal kineties are lacking. The anlagen within kinety 1 occur distinctly later than those in kineties 2 and 3. No caudal
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cirri occur (Petz & Foissner 1996). Nuclear division proceeds as is usual, that is, the macronuclear nodules first fuse and then divide twice (Fig. 64j, l, o, s, u). The micronuclei split once, very rarely twice (Petz & Foissner 1996). Occurrence and ecology: Lamtostylides edaphoni is very likely confined to terrestrial habitats (Foissner 1987, p. 125; 1998, p. 205); probably it is of cosmopolitan distribution. Type locality is the city of Salzburg, Austria, where Berger & Foissner (1987) discovered it in the lower part of a bundle of straw, which was in contact with soil. The bundle was used for the culture of the fungus Leccinum testaceoscabrum. Petz & Foissner (1996, 1997a) found population I of L. edaphoni in the upper (0–5 cm) soil layer (collected on 14.11.1993; mineral debris overgrown with Usnea spacelata, a lichen) at Casey Station, Budd Coast, Wilkes Land, continental Antarctica (66°17'S 110°32'E; about 40 m NN)1. Environmental parameters in this Antarctic soil: humidity up to 27% of wet mass, loss-on-ignition <1.0–7.8% of dry mass, pH 5.1–6.6. Population II was found in moss (Brachythecium austrosalebrosum) collected by R.I.L. Smith on 20.03.1981 at Charlotte Bay, Andrée Island, maritime Antarctic (64°31'S 61°30'W; see also Foissner 1996b, p. 98, 100). Further records: soil of a ski slope (about 2800 m above sea-level) in Tyrol, Austria (Lüftenegger et al. 1986, p. 152); soil of a subalpine grassland in Styria, Austria (Foissner et al. 1990, p. 18); forest stand (Stampfltal) in Lower Austria (Foissner et al. 2005, p. 629); three samples from terrestrial habitats of Namibia (Foissner et al. 2002, p. 60); algal ornithogenic soil from Shirley Island, Windmill Islands, Antarctica (Petz & Foissner 1997b, p. 309). Feeds on bacteria (Berger & Foissner 1987), flagellates, algae, and rarely fungi (Petz & Foissner 1996). Biomass of 106 specimens about 11 mg (Foissner 1987, p. 125) to 16 mg (Petz & Foissner 1996).
Lamtostylides kirkeniensis (Berger & Foissner, 1988) comb. nov. (Fig. 65a–e, Table 22) 1988 Lamtostyla kirkeniensis nov. spec.2 – Berger & Foissner, Zool. Anz., 220: 124, Fig. 24–28, Table 1 (Fig. 65a–e; original description; the holotype slide [accession number 1988/54; Aescht 2003, p. 388] and a paratype slide [1988/55] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria). 2001 Lamtostyla kirkeniensis Berger and Foissner, 1988 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 45 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs).
1
Petz & Foissner (1996) mentioned in the occurrence and ecology section a second site for population I, namely mineral debris covered with Prasiola crispa (alga) at the margin of a small snow water pool on Shirley Island (Windmil Islands, 66°17'S, 110°29'E; about 15 m altitude). 2 Berger & Foissner (1988) provided the following diagnosis: In vivo c. 100 × 27 µm. About 16 adoral membranelles and 7 cirri in the frontal row. 1 cirrus left of the anterior end of the frontal row. 41 left and 42 right marginal cirri on average. 3 transverse cirri, 2 dorsal kineties, 2 micronuclei.
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Nomenclature: The species-group name refers to the city of Kirkenes (Norway) where the species was discovered. Lamtostyla kirkensis in Foissner et al. (2002, p. 712) is an incorrect subsequent spelling. Remarks: For a foundation of the transfer from Lamtostyla to Lamtostylides, see genus section remarks of both taxa. The present species can be separated from the very similar L. edaphoni by having twice the number of marginal cirri (Table 22) and a lower number of dorsal kineties (two vs. three). Lamtostyla islandica has, inter alia, three cirri left of the anterior portion of the amphisiellid median cirral row (vs. one in present species). Foissner et al. (2002) found L. kirkeniensis in the Namib desert (see occurrence section). It matches the type population very well, indicating that it is a cosmopolitan. Morphology: Body size about 100 × 27 µm in life, body length:width ratio in protargol preparations about 4.0:1 (Table 22). Body outline long elliptical, anterior portion distinctly converging. Body very flexible and slightly contractile, about 2:1 dorsoventrally flattened (Fig. 65b). Macronuclear nodules slightly left of midline, anterior nodule behind proximal end of adoral zone; individual nodules with large chromatin bodies of different shape. Contractile vacuole about in mid-body, slightly displaced inwards (Fig. 65c). Cortical granules and cytoplasmic crystals lacking. Cytoplasm with numerous food vacuoles 5–8 µm across and small, greasy granules about 1 µm across, especially around macronuclear nodules. Movement inconspicuous. Adoral zone occupies 20% of body length on average (Table 22), composed of 16 membranelles of ordinary fine structure on average. Buccal field moderately wide. Undulating membranes slightly curved, paroral (endoral?) obviously slightly shorter than endoral (paroral?; Fig. 65d). Cytopharynx extends obliquely backwards. Cirral pattern and number of cirri of usual variability (Fig. 65d, Table 22). Frontal cirri only slightly enlarged, arranged in almost transverse row close to distal portion of adoral zone. Buccal cirrus right of anterior portion of undulating membranes. One slightly enlarged cirrus (= cirrus III/2) left of anterior portion of amphisiellid median cirral row which commences near right frontal cirrus, terminates about at same level as adoral zone, composed of seven cirri on average. Usually three terminal transverse cirri, which project distinctly beyond rear body end; of about same size as marginal cirri. Right marginal row commences about at level of cirrus III/2, ends subterminal. Left row begins behind proximal end of adoral zone, ends about at level of transverse cirri, that is, marginal rows distinctly separated posteriorly. Dorsal bristles about 3 µm long in life, arranged in two kineties; kinety 1 slightly shortened anteriorly; distance between basal body pairs of kinety 2 widens from anterior to posterior (Fig. 65e). Caudal cirri lacking. Occurrence and ecology: Very likely confined to terrestrial habitats (Foissner 1998, p. 205). Type locality of Lamtostylides kirkeniensis is the area near the airport of the city of Kirkenes, Norway, where we discovered it in the Tundra soil (Berger
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Fig. 65a–e Lamtostylides kirkeniensis (from Berger & Foissner 1988. a–c, from life; d, e, protargol impregnation). a: Ventral view of a representative specimen, 102 µm. b: Right lateral view showing dorsoventral flattening. c: Ventral view showing contractile vacuole. d, e: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 85 µm. Arrow marks buccal cirrus. ACR = amphisiellid median cirral row, AZM = adoral zone of membranelles, CV = contractile vacuole, FC = right frontal cirrus, LMR = left marginal row, MA = macronuclear nodule, MI = micronucleus, RMR = right marginal row, TC = transverse cirri, III/2 = cirrus behind right frontal cirrus, 1, 2 = dorsal kineties. Page 333.
& Foissner 1988). According to Foissner (1998) also known from Australia. Foissner et al. (2002, p. 60) found it in five out of 73 soil samples from Namibia, inter alia, in site 38, that is, in a sample of various lichens, including Teloschistes capensis, from ground and gravel surface up to 2 cm in the Central Namib Desert, Cape Cross, about 120 km north of the town of Swakopmund. Lamtostylides kirkeniensis feeds on bacteria, fungal spores, heterotrophic flagellates, and small ciliates, for example, Cyrtolophosis sp. and Colpoda sp. (Berger & Foissner 1988). Biomass of 106 specimens about 20 mg (Foissner 1998, p. 205).
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Lamtostylides halophilus (Foissner, Agatha & Berger, 2002) comb. nov. (Fig. 66a–v, 67a–e, Table 22) 2002 Lamtostyla halophila nov. spec.1 – Foissner, Agatha & Berger, Denisia, 5: 706, Fig. 157a–v, 390a–e, Table 140 (Fig. 66a–v, 67a–e; original description; one holotype slide [accession number 2002/214], one paratype slide [2002/215], and three voucher slides [2002/216–218] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria).
Nomenclature: The species-group name halophil·us, -a, -um (Latin adjective [m; f; n]; thriving in saline habitats) is a composite of the Greek words halós (salt) and philos (preferring), referring to the saline habitats where the species occurs (Foissner et al. 2002). Lamtostyla is feminine whereas Lamtostylides is masculine; thus, the species-group name has to be changed from halophila to halophilus. Remarks: For a foundation of the transfer from Lamtostyla to Lamtostylides, see remarks at genus section of both taxa. We assigned, according to the paper by Petz & Foissner (1996), the present species to Lamtostyla because the oral primordium originates apokinetally and the amphisiellid median cirral row commences anlage formation at the posterior end. Morphologically, Lamtostylides halophilus resembles, due to the cortical granules, Lamtostyla granulifera, which, however, is much larger (about 150 × 35 µm), has three cirri left of the amphisiellid median cirral row, and possesses conspicuous, curved undulating membranes. Lamtostylides edaphoni and L. kirkeniensis are very likely closely related to L. halophilus because of the single cirrus left of the amphisiellid median cirral row. However, they lack cortical granules or have more cirri (6–13 vs. 3–5) in the amphisiellid median cirral row (L. edaphoni, L. kirkeniensis). For comparison with the very similar L. pori see same chapter at this species.
Fig. 66a–i Lamtostylides halophilus (from Foissner et al. 2002. a, d–f, from life; b, c, g–i, protargol impregnation). a: Ventral view of a representative specimen, 65 µm. b, c: Infraciliature of ventral and dorsal side and nuclear apparatus of holotype specimen, 60 µm. Arrow marks cirrus III/2, which originates from same anlage as right frontal cirrus (in addition, it forms – as is usual in hypotrichs – anlage III of the proter; see Fig. 66l). Frontal cirri connected by dotted line; frontal-ventral cirri which originate from same anlage, connected by broken line. d: Oral apparatus. Arrow marks furrow bearing amphisiellid median cirral row. e, f: Ventral and right lateral view of shape variant showing, inter alia, cortical granulation and dorsoventral flattening. Arrow in (e) marks faecal mass. g–i (details from Fig. 66q, s, t): Development of proter’s paroral (arrowheads) and endoral. The paroral moves left of the endoral during shaping of the buccal cavity. AZM = adoral zone of membranelles, ACR = amphisiellid median cirral row, BL = buccal lip, FS = frontal scutum, P = paroral, RMR = right marginal row, TC = transverse cirri (possible pretransverse ventral cirri included), 1–3 = dorsal kineties. Page 336. 1
Foissner et al. (2002) provided the following diagnosis: Size about 70 × 25 µm in vivo; slenderly ellipsoidal. On average 2 macronuclear nodules, 19 cirri in left and 24 in right marginal row, 4 cirri in amphisiellid median cirral row (ACR), 1–2 cirri left of ACR, 1 buccal cirrus, 3–4 transverse cirri, and 2–3 dorsal kineties. Adoral zone continuous, consists of 17–18 membranelles on average. Buccal cavity narrow and flat. The oral primordium develops postorally and independently of the ACR.
d
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The spindle-shaped food vacuoles with the parallel arranged bacteria are somewhat reminiscent of Terricirra (see there for comparison). Morphology: Foissner et al. (2002) studied two populations, of which that from Namibian site (69) was much more variable than the type (Table 22). The diagnosis (see footnote) and description comprise both populations because conspecificity is beyond reasonable doubt. Body size 55–100 × 20–30 µm in life, usually about 70 × 25 µm, length:width ratio 2.2–4.0:1 in life, 1.7–3.3:1, on average about 2.5:1 in protargol preparations, where specimens tend to become inflated. Body outline elliptical to slenderly elliptical. Body dorsoventrally flattened up to 2:1 (Fig. 66a, e, f; Table 22). Macronuclear nodules in middle third of cell left of midline, ellipsoidal to elongate ellipsoidal (up to 4:1), usually with rather irregular outline both in life and protargol preparations; chromatin bodies large and irregularly branched. Micronuclei globular, usually one attached to each macronuclear nodule, difficult to recognise in life and protargol preparations because hyaline and faintly impregnated. Contractile vacuole in midbody at left cell margin. Cytopyge in posterior end left of midline, faecal clumps contain highly refractive bacterial spores embedded in fluffy material (Fig. 66e). Cortical granules loosely arranged, in life difficult to recognise because colourless and only 0.5–1.0 µm across, usually impregnate with protargol (Fig. 66e, 67a–c, e). Cytoplasm colourless, without distinct crystals and lipid droplets. Food vacuoles up to 10 µm across with parallel arranged content (see remarks; Fig. 66a). Glides slowly on microscope slide and organic debris. Oral apparatus inconspicuous because adoral zone narrow and occupying only 27% of body length; composed of an average of 18 membranelles, bases of largest membranelles 4 µm wide in life, only about 2 µm in anterior third of zone. Buccal cavity narrow and flat. Buccal lip rather conspicuous, arched, covers posterior third of adoral zone. Paroral and endoral very small, staggered in parallel, both near adoral zone and very likely composed of a single row of cilia (basal bodies), often slightly obliquely arranged. Paroral cilia form distinct, triangular velum with cilia 12 µm long at anterior end and only 5 µm at posterior (Fig. 66a, b, d, 67a, d). Pharyngeal fibres distinct only in protargol preparations. Cirral pattern and number of cirri of usual variability (Fig. 66b, Table 22). Cirri about 12 µm long in life, very fine because composed of only 2–4 cilia, except for frontal cirri usually comprising six cilia each; left frontal cirrus often enlarged and
b
Fig. 66j–n Lamtostylides halophilus (from Foissner et al. 2002. Protargol impregnation). Infraciliature of dividers in ventral view. The oral primordium (OP) originates apokinetally in mid-body and produces five cirral streaks (numbers I, II, III, V, VI), which unite with streaks generated by parental paroral (anlage I) and frontoventral cirri (anlage II by buccal cirrus; anlage III by cirrus III/2; anlage V by the last cirrus of amphisiellid median cirral row) to form long primary primordia (n). Anlage VI, however, is generated entirely by the oral primordium (k–n). Species with only one cirrus (= cirrus III/2) left of amphisiellid median cirral row lack anlage IV (see text and general section of the Amphisiellidae). Note that dividers become distinctly smaller and broader during this process (j = 66 µm, k = 64 µm, l = 68 µm, m = 58 µm, n = 60 µm). OP = oral primordium, I, II, III, V, VI = frontal-ventral-transverse cirri primordia. Page 336.
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SYSTEMATIC SECTION
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composed of eight cilia. Buccal cirrus invariably right of anterior end of paroral. Usually one cirrus (= cirrus III/2), rarely two cirri left of amphisiellid median cirral row which is accompanied by distinct furrow, composed of only four cirri on average, and terminates distinctly ahead of level of buccal vertex. Usually four transverse cirri, insert at right margin of flat crest, project beyond rear body end. Right marginal row commences ahead of level of buccal cirrus, terminates, like left row, about at level of transverse cirri; thus, marginal rows widely separated posteriorly; left row commences near proximal end of adoral zone. Dorsal bristles 3–4 µm long, arranged in three kineties; bristles widely spaced, especially in kinety 1, which is lacking in about 60% of specimens from Namibian site (69); no caudal cirri (Fig. 66a–c, 67a, d). Cell division (Fig. 66g–v): Division of Lamtostylides halophilus is very similar to that of L. edaphoni (Fig. 64j–u), Lamtostyla australis (Fig. 31j–z), and Lamtostyla perisincirra (Fig. 39g–m). Thus, Foissner et al (2002) described cell division not in detail, but referred the reader to the figures and figure explanations. Three main differences should be noted: (i) the oral primordium does not develop near the transverse cirri, but in mid-body (Fig. 66j, k); (ii) no patch of basal bodies develops at the posterior end of the amphisiellid median cirral row, which originates, as is typical for the group, by alignment of the cirri formed by the two rightmost (V and VI) anlagen (Fig. 66s–u; note that in Lamtostylides, which has only one cirrus left of the amphisiellid median cirral row, anlage IV is lacking; see general section); (iii) the parental adoral zone of membranelles performs a rather distinct internal reorganisation in middle dividers (Fig. 66g, q). Occurrence and ecology: To date found only in three highly saline soil samples from Namibia (sites 11, 18, 69), indicating that Lamtostylides halophilus is a halophilus species. Type locality is site 18, that is, a highly saline soil from the margin of a pond near of the town of Maltahöhe (24°55'S 16°55'E), Namibia (Foissner et al. 2002). The populations studied by Foissner et al. (2002) reproduced readily in the non-flooded Petri dish cultures so that ontogenesis could be studied. Feeds on long, spore-forming bacteria (Foissner et al. 2002).
b
Fig. 66o–r Lamtostylides halophilus (from Foissner et al. 2002. Protargol impregnation). Infraciliature of ventral (o, q) and dorsal side and nuclear apparatus (p, r) of middle dividers. The primary primordia (see previous stages) split in the middle to produce five cirral anlagen each in proter and opisthe (o, q). The dorsal kineties reproduce by intrakinetal proliferation of dikinetids (p, r; parental kinetids ciliated). According to the proter in (q), only the anlagen II, III, and VI (= anlagen 2, 3, and 5 in Foissner et al. 2002) produce transverse cirri, that is, anlage V does not produce such a cirrus; however, further data are needed to confirm this assumption. Note that anlage IV is lacking because the remaining anlagen can be easily homologised, via the cirri which they form, with the anlagen I, II, III, V, and VI of the 18-cirri hypotrichs (see text). o, p = 48 µm, q, r = 49 µm. I–III, V, VI = frontal-ventral-transverse cirri anlagen, 1–3 = dorsal kineties. Page 336.
Fig. 66s–v Lamtostylides halophilus (from Foissner et al. 2002. Protargol impregnation). Infraciliature of ventral (s–u) and dorsal side and nuclear apparatus (v) of late and very late dividers, s = 52 µm, t = 64 µm, u, v = 76 µm. Parental structures white, new black. The amphisiellid median cirral row is formed by anlagen V (posterior portion) and VI (anterior portion). Dorsal morphogenesis clearly shows that no caudal cirri are formed (v). Page 336.
342 SYSTEMATIC SECTION
Fig. 67a–e Lamtostylides halophilus (from Foissner et al. 2002. Protargol impregnation). a–c: Cortical granulation of ventral (a) and dorsal (c) side and in optical section (b). The granules have about the same size as the cirral bases, which are thus difficult to recognise. Arrowhead marks undulating membranes. d: Infraciliature of ventral anterior portion. Arrowheads mark cirri of amphisiellid median cirral row, arrow denotes single cirrus (= cirrus III/2) left of amphisiellid median cirral row. e: Higher magnification of cortical granulation in mid-body of dorsal side. Explanation of original labelling: AZM = adoral zone of membranelles, BU = buccal cirrus, CV = contractile vacuole, EM = endoral, MA = macronuclear nodules, PM = paroral. Page 336.
Lamtostylides 343
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SYSTEMATIC SECTION
Lamtostylides pori (Wilbert & Kahan, 1986) comb. nov. (Fig. 68a–h, Table 22) 1986 Perisincirra pori nov. spec. – Wilbert & Kahan, Arch. Protistenk., 131: 133, Abb. 3, 4a–f, 6a–f (Fig. 68a–h; original description; site where type slides are deposited not mentioned, perhaps in the Institut für landwirtschaftliche Zoologie und Bienenkunde, University of Bonn, Germany). 1987 Hemisincirra pori Wilbert, 1986 – Foissner, Progr. Protistol., 2: 124 (combination with Hemisincirra; incorrect authorship). 2001 Perisincirra pori Wilbert and Kahan, 1986 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 72 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs).
Nomenclature: Wilbert & Kahan (1986) dedicated this species to F. D. Por from the Hebrew University Jerusalem, who contributed significantly to the investigation of the Sinai coast. Remarks: This species is obviously closely related to Lamtostylides halophilus originally described in our monograph on Namibian soil ciliates (Foissner et al. 2002). Unfortunately, we overlooked the strong resemblance and therefore did not discuss the similarity in the comparison with related species. Possibly L. halophilus and L. pori are only subspecies; however, I preliminarily retain the species status of L. halophilus mainly because of the cortical granulation. The cortical granules of L. halophilus are difficult to recognise in life, but usually impregnate heavily with protargol (Fig. 68a–c, e). By contrast, the micrographs of the protargol-impregnated specimens of L. pori presented in Wilbert & Kahan (1986; their Abb. 6a–f) absolutely do not show a cortical granulation. This indicates that such organelles are lacking in L. pori, although this assumption has to be checked by live data from populations collected from near the type locality. Whether differences in other features (eight cirri in frontal area in L. pori vs. about nine in L. halophilus; dorsal kinety 1 roughly continuous vs. with more or less distinct break; marginal cirri composed of two cilia and roughly continuous posteriorly vs. usually composed of four cilia and distinctly separated; moves up and down vs. slowly gliding) are sufficient for the separation at species and/or subspecies level can be only decided by detailed studies on further populations, inter alia, from the type locality of L. pori. The foundation for the classification of the present species in Lamtostylides is the same as in other species (see genus section). Morphology: Body size in life 75–110 × 12–19 µm (specimen illustrated about 115 µm long); body length:width ratio 5.1:1 on average in life (Table 22). Body outline elongate elliptical. Body distinctly twisted about main axis. Two ellipsoidal macronuclear nodules left of midline; one micronucleus attached to each macronuclear nodule (Fig. 68a, b). Contractile vacuole near left cell margin slightly behind buccal vertex; in specimen illustrated at about 32% of body length (Fig. 68a). Presence/absence of cortical granules not known; in micrographs of protargolimpregnated specimens presented in original description (Abb. 6a–f in Wilbert & Kahlan 1986) no granules recognisable (see remarks). Details about cytoplasm (colour, inclusions) and consistency of cell (flexible/rigid) not described. Movement
Lamtostylides
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Fig. 68a, b Lamtostylides pori (from Wilbert & Kahan 1986. From life and after protargol impregnation). Ventral and dorsal view of same specimen (?) showing, inter alia, infraciliature and nuclear apparatus, 115 µm (note that this specimen is somewhat longer than the size given in the text). Short arrow marks cirrus (= cirrus III/2) behind right frontal cirrus, long arrows mark amphisiellid median cirral row. AZM = adoral zone of membranelles, CV = contractile vacuole, FC = right frontal cirrus, LMR = left marginal row, MA = macronuclear nodule, RMR = right marginal row, TC = transverse cirrus, 1–3 = dorsal kineties. Page 344.
conspicuous because cell is directed perpendicularly to substrate while moving up and down. Adoral zone occupies about 23% of body length, composed of 17 membranelles on average (Table 22). Buccal field narrow. Undulating membranes as in L. halophilus, that is, short paroral left of anterior half of endoral which is almost twice as long as paroral (Fig. 68a, c). Cirral pattern as shown in Fig. 68a, that is, more or less exactly as in L. halophilus. Three frontal cirri composed of four cilia each (left one possibly composed of six cilia), arranged in slightly oblique pseudorow. Buccal cirrus close to anterior end of undulating membranes. In total eight cirri on frontal field (frontal cirri, buccal cirrus, amphisiellid median cirral row), that is, row composed of four cirri (cirrus [= cirrus III/2] shifted leftwards included); row terminates, according to specimen illus-
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Fig. 68c–h Lamtostylides pori (from Wilbert & Kahan 1986. Protargol impregnation). c: Very early divider, 95 µm. The oral primordium is formed in central part of ventral side without contact to parental cirri. d: Early to middle divider, 97 µm. Parental cirri on frontal field have modified to primordia and the oral primordium produced some anlagen extending anteriad. e: Middle divider, 77 µm. The five anlagen of proter and opisthe are separated (note the three parental transverse cirri). f, g: Late dividers, f = 83 µm, g = 76 µm. Arrows in (f) mark the five cirri anlagen of opisthe. g: Very late divider, 91 µm. OP = oral primordium, 3 = new dorsal kineties. Page 344.
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trated, about at level of buccal vertex (Fig. 68a). Postperistomial cirri lacking. Usually two, rarely three, about 15 µm long transverse cirri close to rear cell end; composed of about 4–6 cilia each. Right marginal row commences dorsolaterally near distal end of adoral zone, extends sigmoidally to near rear cell end. Exact origin of left marginal row not described and not clearly recognisable from illustration, extends sigmoidally onto dorsolateral surface posteriorly. Marginal cirri and cirri of amphisiellid median row composed of two cilia only (Fig. 68c). Length of dorsal bristles not described, according to Fig. 68b about 5 µm long, arranged in three kineties with rather widely spaced bristles. All rows obviously more or less continuous, that is, break in kinety 1 of L. halophilus not present. Caudal cirri obviously lacking because neither described nor clearly recognisable in illustrations. Unfortunately, Wilbert & Kahan (1986) did not describe the formation of the dorsal infraciliature. Cell division (Fig. 68c–h): Wilbert & Kahan (1986) described the ontogenesis of L. pori. It proceeds as in L. halophilus; please refer to the illustrations and the legends. The presence of only five frontal-ventral-transverse cirri anlagen is the reason for the classification in Lamtostylides. Unfortunately, the formation of the dorsal ciliature is not described, but likely only intrakinetal proliferation occurs. Occurrence and ecology: Possibly confined to saline soils. Type locality of Lamtostylides pori is the area about 10 km south of the city of Eilat, Israel. Wilbert & Kahan (1986) discovered it in the highly saline soil (solonetz) of flat pools in the supralittoral which are fed by seawater and rainwater; however, most of the time these pools are dry. Salinity ranged from 18–60‰. No further records published. Feeds on bacteria and single-celled algae (Wilbert & Kahan 1986). Biomass of 106 specimens 19 mg (Foissner 1987, p. 124; 1998, p. 204).
Lamtostylides hyalinus (Berger, Foissner & Adam, 1984) comb. nov. (Fig. 69a–d, Table 22) 1984 Tachysoma hyalina nov. spec.1 – Berger, Foissner & Adam, Zool. Jb. Syst., 111: 359, Abb. 54–57, Tabelle 8 (Fig. 69a–d; see remarks; original description; the holotype slide [accession number 1982/60; Aescht 2003, p. 388] is deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria). 1987 Lamtostyla hyalina (Berger, Foissner, and Adam, 1984) nov. comb. – Berger & Foissner, Zool. Jb. Syst., 114: 216 (combination with Lamtostyla). 1988 Lamtostyla hyalina (Berger, Foissner, and Adam, 1984) Berger and Foissner, 1987 – Berger & Foissner, Zool. Anz., 220: 126, Fig. 29–32, Table 1 (Fig. 69a–d; revision of Lamtostyla).
1
Berger et al. (1984) provided the following diagnosis: In vivo etwa 50 × 16 µm große, ausgeprägt parallelseitige Tachysoma mit 4 Cirren in der Frontalreihe und je etwa 11 Cirren in den gleich langen Marginalreihen. Drei Dorsalkineten, von denen die rechte aus nur 1 Basalkörperpaar besteht. Durchschnittlich 11 adorale Membranellen, von denen die 3 vorderen durch eine deutliche Lücke von den hinteren getrennt sind.
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SYSTEMATIC SECTION
2001 Lamtostyla hyalina (Berger, Foissner and Adam, 1984) Berger and Foissner, 1987 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 88 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs).
Nomenclature: No derivation of the name is given in the original description. The species-group name hyalin·us, -a, -um (Latin adjective [m; f; n]; glassy, transparent) refers to the fact that the species is transparent at low magnification. Lamtostyla is feminine, whereas Lamtostylides is masculine; thus, the species-group name has to be changed from hyalina to hyalinus. The designation “Lamtostyla hyalina Berger, Foissner & Adam, 1984” in Foissner (1996a, p. 284) is incorrect because the species was not established in Lamtostyla but in Tachysoma. Tachysoma is neuter (Aescht 2001, p. 302); thus, the correct name in the original description would have been Tachysoma hyalinum. Remarks: For a foundation of the transfer from Lamtostyla to Lamtostylides, see genus section remarks of both taxa. The present species was originally assigned to Tachysoma, an oxytrichid group (for review, see Berger 1999, p. 431), because of the lack of caudal cirri. Later, we transferred it to Lamtostyla because it agrees with L. lamottei, type of Lamtostyla, in several features, namely, dorsal kineties without caudal cirri, small body length, reduced number of transverse cirri and pretransverse ventral cirri, lack of postoral ventral cirri, and possession of a short amphisiellid median cirral row which terminates about at level of buccal vertex. Lamtostylides hyalinus usually has five transverse cirri (Fig. 69c). However, five (true) transverse cirri can only be formed when six (I–VI) frontal-ventral-transverse cirri anlagen are present (see Fig. 6a in Berger 1999). By contrast, the single cirrus (right frontal cirrus in present case?) left of the anterior portion of the amphisiellid median cirral row strongly indicates that anlage IV is lacking in the present species, and in Lamtostylides in general. At least two possibilities exist to explain this inconsistency: (i) anlage IV produces only the transverse cirrus (a similar phenomenon is described for Amphisiella annulata; Fig. 17s); and (ii) the cirrus marked with TC in Fig. 69c is not a transverse cirrus, but a (relatively large) pretransverse ventral cirrus. Ontogenetic data are needed for a correct interpretation of the cirral pattern. The illustrations in the original description are rather faint. Thus, Berger & Foissner (1988) provided a better printable version of the original illustrations. Morphology: Body size about 50 × 16 µm in life; length:width ratio 2.3:1 on average in protargol preparations (Table 22). Body outline with parallel margins, both ends broadly rounded; left margin slightly indented in area of adoral zone. Body flattened about 2:1 dorsoventrally, ventral side flat, dorsal side slightly convex (Fig. 69b); hyaline at low magnification; slightly contractile under cover-glass pressure. Macronuclear nodules about in central portion of cell, narrowly spaced, in life about 7 × 4 µm, with some small chromatin bodies. Usually one micronucleus attached to each macronuclear nodule, rarely one between the nodules (Fig. 69a, d); micronuclei in protargol preparations always weakly stained. Contractile vacuole near left cell margin about in mid-body, without distinct collecting canals. Cortical granules lack-
Lamtostylides
349
Fig. 69a–d Lamtostylides hyalinus (from Berger & Foissner 1988. a, b, from life; c, d, protargol impregnation). a, b: Ventral and right lateral view of representative specimen, 50 µm. c, d: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 32 µm. Short arrow marks rear end of amphisiellid median cirral row, long arrow denotes (pretransverse?) ventral cirrus in U-cavity formed by transverse cirri; arrowhead marks buccal cirrus, which is likely composed of two basal bodies only. This specimen has only one micronucleus, which is in between the two macronuclear nodules. E = endoral, FC = left and right frontal cirrus (left one in gap of adoral zone), P = paroral, TC = rightmost transverse cirrus (whether this is a true transverse cirrus or an enlarged pretransverse ventral cirrus can only be decided after studying cell division; see remarks), 1–3 = dorsal kineties. Page 347.
ing. Cytoplasm colourless, with some small, colourless globules. Movement very rapid, sliding hastily to and fro. Adoral zone occupies 29% of body length on average in protargol preparations, composed of 11 membranelles on average (Table 22) separated by distinct gap into proximal portion and distal portion with about three membranelles; extends onto right body margin to about 11% of body length. Buccal field narrow. Undulating membranes relatively short, only slightly overlapping, that is, endoral distinctly displaced anteriorly, paroral commences slightly behind level of buccal cirrus and ter-
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minates at level of proximal end of adoral zone. Endoral likely composed of widely spaced basal bodies. Pharyngeal fibres inconspicuous (Fig. 69a, c). Frontal cirri arranged in very oblique pseudorow; left cirrus more or less in gap of adoral zone, right one left of anterior portion of amphisiellid median cirral row. Buccal cirrus right of mid-portion of endoral, likely composed of two basal bodies (cilia) only. Amphisiellid median cirral row composed of four cirri only, terminates slightly ahead of level of buccal vertex. Postperistomial cirrus lacking. Usually five, about 12 µm long transverse cirri arranged in U-shaped pattern near body end, cirri thus distinctly projecting beyond body margin; in centre of U-cavity usually a small (pretransverse?) ventral cirrus (see remarks for comment about transverse cirri). Marginal rows commence about at level of buccal vertex, end subterminally and are therefore distinctly separated posteriorly; distance between 6–8 µm long cirri increases in posterior direction. Dorsal bristles about 3 µm long, arranged in three kineties; kinety 1 composed of a single bristle. Caudal cirri lacking (Fig. 69c, d). Cell division: Morphogenesis commences with the formation of an oral primordium at the left transverse cirrus. A little later the rearmost cirrus of the amphisiellid median cirral row modifies to an anlage (Berger et al. 1984). See remarks for a reflection about the formation of the transverse cirri. Occurrence and ecology: Very likely confined to terrestrial habitats (Foissner 1987, p. 127; 1998, p. 205) and possibly an indicator for moder (Foissner 1985, p. 85). Type locality of Lamtostylides hyalinus is the Stubnerkogel, a mountain near the village of Bad Gastein, Austria, where we found it with moderate abundance in the upper soil layer (0–5 cm) of an alder stand (about 1780 m above sea-level) at the sub-alpine timber line (Berger et al. 1984); for some autecological data of this population, see Foissner & Peer (1985, p. 44). Records not substantiated by morphological data: soil from the Enns/Danube River floodplain, Upper Austria (Foissner 2004, p. 370); forest stands in eastern Austria (Foissner et al. 2005, p. 629); litter of spruce and beech forests near the city of Ulm, southern Germany (Funke 1986, p. 72; Lehle 1989, p. 142); in one out of 73 soil samples from Namibia (Foissner et al. 2002, p. 60); Sphagnum moss under mire vegetation (pH 5.0) from Tafelkop, Gough Island, and soil from occupied Wandering Albatross nest (pH 4.7) as well as muddy soil from beside grey lava outcrop near bases of Junior’s Kop, Marion Island, southern Indian Ocean (Foissner 1996a, p. 284). Food not known, likely bacteria. Biomass of 106 specimens only about 4 mg (Foissner 1987, p. 127; 1998, p. 205).
Paramphisiella
351
Paramphisiella Foissner, 1988 1988 Paramphisiella nov. gen.1 – Foissner, Stapfia, 17: 121 (original description). Type species (by original designation): Amphisiella acuta Foissner, 1982. 1994 Paramphisiella Foissner, 1988 2 – Eigner & Foissner, J. Euk. Microbiol., 41: 258 (redefinition). 1996 Paramphisiella Foissner, 1988 3 – Petz & Foissner, Acta Protozool., 35: 277 (redefinition). 2001 Paramphisiella Foissner 1988 – Aescht, Denisia, 1: 117 (catalogue of generic names of ciliates). 2001 Paramphisiella Foissner, 1988 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 69 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Paramphisiella Foissner, 1988 – Lynn & Small, Phylum Ciliophora, p. 452 (guide to ciliate genera).
Nomenclature: Paramphisiella is, according to Foissner (1988), a composite of the Greek prefix par- or para- (close to; related; deviating) and the genus-group name Amphisiella (see there for derivation). Likely it should indicate a relationship or difference to Amphisiella. Like Amphisiella of feminine gender (Foissner 1988). Characterisation (A = supposed apomorphy): Amphisiellidae with continuous adoral zone of membranelles. Body slender, spindle-shaped body (A?). Three frontal cirri. Buccal cirrus present. Only one cirrus (= cirrus III/2) left of anterior portion of amphisiellid median cirral row because frontal-ventral-transverse cirri anlage IV lacking (A?). Amphisiellid median cirral row originates from anlagen V and VI. Postperistomial cirrus and transverse cirri lacking (type species), but short, longitudinal row present in P. caudata (A?). One left and one right marginal row. Only bipolar kineties present, that is, dorsomarginal row and/or kinety fragmentation lacking. Caudal cirri present. Mainly terrestrial. Additional characters: body very flexible; contractile vacuole ahead of midbody, at left cell margin; dorsal bristles less than 5 µm long; three dorsal kineties. Remarks: Foissner (1988) established Paramphisiella for Amphisiella acuta, which lacks transverse cirri (see species description). Eigner & Foissner (1994) and Petz & Foissner (1996) redefined Paramphisiella. Interestingly, they included the presence of longitudinally arranged transverse cirri into the diagnosis, although the type species lacks such cirri. In addition, Eigner & Foissner (1994, p. 258) wrote that P. caudata is the single (assignable?) species. However, this is formally incorrect because the type species (P. acuta) is always assignable. The presence of a single cirrus (= cirrus III/2) left of the anterior portion of the amphisiellid median cirral row implies that Paramphisiella lacks the frontal-ventral 1
Foissner (1988) provided the following diagnosis: Amphisiellidae mit 1 Cirrus links der Ventralreihe im Frontalfeld. Caudalcirren vorhanden. 2 Eigner & Foissner (1994) provided the following improved diagnosis: The amphisiellid median cirral row originates from two rightmost anlagen. One cirrus left of amphisiellid median cirral row. Transverse cirri longitudinally arranged, originate from single anlage. Caudal cirri present. 3 Petz & Foissner (1996) provided the following improved diagnosis: The oral primordium originates in close contact with the amphisiellid median cirral row. The amphisiellid median cirral row commences anlagen formation within-row and originates from two rightmost anlagen. All dorsal kineties develop intrakinetally. One cirrus left of amphisiellid median cirral row. Transverse cirri longitudinally arranged, originate from single anlage. Caudal cirri present.
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SYSTEMATIC SECTION
cirral anlage IV because the amphisiellid median cirral row is usually produced by the anlagen V and VI. For details see general section. Shi (1999, p. 253) and Shi et al. (1999, p. 100) synonymised Paramphisiella and Hemiamphisiella with Uroleptoides. Preliminarily I do not accept this synonymy because at present we do not know whether or not Uroleptoides kihni, type of Uroleptoides, has transverse cirri, which are lacking in P. acuta, type of Paramphisiella. In addition, Uroleptoides kihni has very likely more than one cirrus left of the anterior portion of the amphisiellid median cirral row (vs. one in Paramphisiella). Synonymy of Paramphisiella and Hemiamphisiella can be excluded because Paramphisiella has only five frontal-ventral cirri anlagen (anlage IV lacking), whereas anlage IV is present in H. terricola, type of Hemiamphisiella. In addition, the anlage IV of Hemiamphisiella produces a postperistomial cirrus (= homologous to cirrus IV/2 of the 18-cirri hypotrichs), indicating that it is not closely related to the remaining, typical amphisiellids which lack this cirrus. Species included in Paramphisiella (alphabetically arranged basionyms are given): (1) Amphisiella acuta Foissner, 1982; (2) Uroleptoides caudata Hemberger, 1985.
Key to Paramphisiella species 1 8–16, on average 14, macronuclear nodules; amphisiellid median cirral row composed of 38–42 cirri on average (Fig. 73a–e). . . Paramphisiella caudata (p. 358) - 24–38, on average about 32, macronuclear nodules; amphisiellid median cirral row composed of 19 cirri on average (Fig. 70a–g). Paramphisiella acuta (p. 352)
Paramphisiella acuta (Foissner, 1982) Foissner, 1988 (Fig. 70a–j, Table 24) 1982 Amphisiella acuta nov. spec.1 – Foissner, Arch. Protistenk., 126: 35, Abb. 2a–f, 42, Tabelle 5 (Fig. 70a–g; original description; the holotype slide [accession number 1981/85; Aescht 2003, p. 379] is deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria). 1988 Paramphisiella acuta (Foissner, 1982) nov. comb. – Foissner, Stapfia, 17: 121 (combination with Paramphisiella). 1994 Amphisiella acuta Foissner, 1982 – Shin, Dissertation, p. 42, Fig. 3A–C, Table 4 (Fig. 70h–j; redescription of a Korean population). 2001 Paramphisiella acuta (Foissner, 1982) Foissner, 1988 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 7 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs).
1
Foissner (1982) provided the following diagnosis: In vivo etwa 110–170 × 18–22 µm große, nach hinten keilförmig verjüngte, leicht S-förmig gebogene Amphisiella mit ungefähr ½ körperlanger Ventralreihe. Durchschnittlich 32 ellipsoide Makronucleus-Teile. Adoral Membranellenzone etwa 1/8 körperlang. 3 Dorsalkineten.
Paramphisiella
353
Nomenclature: No derivation of the name is given in the original description. The species-group name acut·us, -a, -um (Latin adjective [m; f; n]; pointed) obviously refers to the pointed posterior body portion. Type species of Paramphisiella. Remarks: Paramphisiella acuta was discovered by Foissner (1982) in an Austrian alpine soil and later recorded from various terrestrial habitats in Austria, Slovakia, Himalayas, and Korea (see occurrence and ecology section). Interestingly, it was never recorded from the southern hemisphere, indicating that this species has a holarctic distribution (Foissner 1987, 1998). Foissner (1982) described transverse cirri between the rear end of the marginal rows. Later, in his important paper on amphisiellids (Foissner 1988), he re-examined the slides and found that he had misinterpreted the caudal cirri as transverse cirri. Thus, he established Paramphisiella, which differs from Uroleptoides and Lamtostyla by the presence of caudal cirri and the lack of transverse cirri. Shin (1994), who did not know this paper, described three transverse cirri located at the ventral surface. However, this statement must not be overinterpreted because in tailed species it is very difficult to recognise, even in protargol preparations, whether terminal cirri are transverse or caudal cirri. But there is no doubt that ontogenetic data are needed for a final decision about the correct designation of these terminal cirri. The illustrations by Shin (1994) are modified redrawings of the type population, as indicated, inter alia, by the identical nuclear apparatus and dorsal kinety pattern. The present species differs from Paramphisiella caudata mainly by the number of macronuclear nodules and the number of cirri composing the amphisiellid median cirral row (see key). Afroamphisiella species, which also lack transverse cirri, do not have caudal cirri (vs. present – although not very conspicuous – in Paramphisiella species). Lamtostyla and Uroleptoides species have transverse cirri and lack caudal cirri. Morphology: This chapter is based on the original description by Foissner (1982) unless otherwise indicated. For a morphometric characterisation of the Korean population studied by Shin (1994), see Table 24. Body size in life about 110–170 × 18–22 µm, body length:width ratio of specimen shown in Fig. 70a about 8.3:1, of prepared specimens 7.3:1 on average (Table 24); body size of Korean specimens about 110–170 × 20–40 µm in life (Shin 1994). Body outline very slender, slightly to distinctly sigmoidal and cuneate, posterior body portion often twisted by half a turn about main body axis; anterior and posterior end narrowly rounded. Body flexible; dorsoventrally flattened about 2:1 only in anterior and posterior body regions. Macronuclear nodules in main body portion, that is, lacking in anterior and posterior regions and near right cell margin (Fig. 70e); individual nodules 6.0 × 3.5 µm in life, with a moderate number of small chromatin bodies. Micronuclei did not impregnate with protargol method used. Contractile vacuole slightly ahead of mid-body near left cell margin, during diastole with inconspicuous collecting canals. Pellicle colourless and flexible. Cortical granules lacking. Cytoplasm colourless, with many about 9-µm-sized food vacuoles with unknown content and some vacuoles 4–7 µm across with small, yellow crystals. Movement slow, twitching to and fro.
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SYSTEMATIC SECTION
Fig. 70a–g Paramphisiella acuta (from Foissner 1982. a–c, from life; d–g, protargol impregnation). a: Ventral view of a representative specimen, 150 µm. b: Left lateral view. c: Dorsal view. d, e: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 112 µm. f, g: Infraciliature of ventral side of anterior and posterior body portion. Arrow marks buccal cirrus. Frontal cirri connected by dotted line, broken line connects right frontal cirrus and cirrus III/2. In the original description the caudal cirri were misinterpreted as transverse cirri. ACR = amphisiellid median cirral row, AZM = adoral zone of membranelles, CC = caudal cirri (see remarks), CV = contractile vacuole, FC = right frontal cirrus, LMR = left marginal row, MA = macronuclear nodule, RMR = right marginal row, 1–3 = dorsal kineties. Page 352.
Adoral zone very short, that is, occupies on average only 14% of body length and composed of 15 membranelles of usual fine structure (Fig. 70a, d, f, Table 24). Bases of largest membranelles about 5 µm in life. Buccal field deep. Undulating
Paramphisiella
355
membranes distinctly curved and roughly in parallel. Pharyngeal fibres conspicuous, form sabre-shape structure in life. Cirral pattern and number of cirri of usual variability (Fig. 70a, d, f, g, Table 24). Three slightly enlarged frontal cirri. Buccal cirrus fine, likely composed of a single, short row of basal bodies/cilia only. Cirrus behind right frontal cirrus (= cirrus III/2) enlarged (thus, Foissner designated it as [fourth] frontal cirrus); no further cirri left of anterior portion of amphisiellid median cirral row, which commences about at level of right frontal cirrus, terminates at 40% of body length on average, in specimen illustrated at about 54% (Fig. 70d); distance between cirri gradually increases in posteriad direction. Postperistomial cirrus and transverse cirri lacking (see remarks). Right marginal row extends onto dorsal side anteriorly, commences about at level of right frontal cirrus, terminates – like left row – near cell end, that is, marginal rows separated posteriorly, but indistinctly set off from caudal Fig. 70h–j Paramphisiella acuta (from Shin 1994. cirri (see remarks). Left row commences h, from life; i, j, protargol impregnation). Ventral left of proximal end of adoral zone. view and infraciliature of ventral and dorsal side Marginal cirri fine, in life about 9 µm and nuclear apparatus. h = 140 µm, i, j = 155 µm. I long; distance among cirri in posterior suppose that these are modified redrawings of Foissner’s illustrations (compare nuclear pattern portion about twice as long as in ante- shown in [e] and [j]). Arrow marks rear end of amrior region. phisiellid median cirral row, which is composed of Dorsal bristles about 5 µm long in only 14 cirri in the Korean population (vs. 19 in life, arranged in three kineties extending type population). Page 352. to rear cell end; kinety 1 distinctly, kinety 2 slightly, and kinety 3 not shortened anteriorly (Fig. 70e). Bristles within each kinety about equally spaced. Three caudal cirri present, that is, each kinety forms such a cirrus; in life 14 µm long (see remarks). Occurrence and ecology: Very likely confined to terrestrial habitats and possibly of holarctic (and paleotropic? Himalayan region) distribution (Foissner 1987, 1998). Type locality of P. acuta is the Schlossalm area near the village of Bad Hofgastein, Austria, where Foissner (1982) discovered it in soil of an alpine pasture
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Table 24 Morphometric data on Paramphisiella acuta (ac1, type population, from Foissner 1982; ac2, Korean population, from Shin 1994) and Paramphisiella caudata (ca1, type population, from Hemberger 1982, 1985; ca2, from Eigner & Foissner 1994; ca3, from Shin 1994) Characteristics a Body, length
Species mean
ac1 ac2 ca2 ca3 Body, width ac1 ac2 ca2 ca3 Body length:width, ratio ac2 ca1 ca3 Adoral zone of membranelles, length ac1 ac2 ca2 ca3 Body length:length of adoral zone, ratio ac2 ca1 ca3 Undulating membranes, length ac2 ca3 Undulating membrane length:length of ac2 adoral zone, ratio ca3 Pharyngeal fibres, length ac2 Anterior body end to rear end of ac1 amphisiellid median cirral row, distance ca2 Macronuclear nodule, length ac1 ac2 ca2 ca3 Macronuclear nodule, width ac1 ac2 ca2 ca3 Macronuclear nodules, number ac1 ac2 ca1 ca2 ca3 Micronucleus, diameter ac2 Micronucleus, length ca2 ca3 Micronucleus, width ca2 Micronuclei, number ac2 ca1 ca2 ca3 Adoral membranelles, number ac1
111.4 137.1 148.1 172.1 15.2 24.7 41.0 53.6 5.7 – 3.3 14.7 19.2 37.8 46.9 7.1 5.0 3.7 14.3 32.5 0.7 0.7 16.7 44.6 121.3 5.0 5.8 10.4 7.6 1.9 2.6 6.8 5.2 31.7 23.9 – 14.8 18.5 1.7 4.5 3.1 3.7 2.5 – 4.0 3.8 14.7
M
SD
SE
108.0 143.0 146.5 168.0 15.0 23.0 41.0 57.0 5.5 – 3.2 15.0 20.0 38.0 46.0 7.1 – 3.6 14.0 32.4 0.7 0.7 19.0 43.0 123.5 5.3 6.0 10.5 7.6 1.9 2.5 7.0 5.3 31.5 21.0 14.0 14.0 20.0 1.8 4.4 3.0 3.8 2.0 – 4.0 4.0 15.0
17.6 2.0 9.6 20.3 0.8 5.5 5.1 10.5 1.1 – 0.4 0.9 1.9 1.7 5.0 0.7 – 0.3 2.2 2.8 0.1 0.0 3.6 5.6 8.6 0.8 1.2 1.9 1.1 0.3 0.4 1.0 0.7 4.2 5.4 – 1.1 4.4 0.3 0.5 0.4 0.5 1.0 – 0.6 0.9 0.7
5.1 6.7 2.8 5.6 0.2 1.8 1.5 2.9 0.4 – 0.1 0.3 0.6 0.5 1.4 0.2 – 0.1 0.7 0.8 0.0 0.0 1.4 1.6 2.5 0.2 0.4 0.5 0.3 0.1 0.1 0.3 0.2 1.2 1.8 – 0.3 1.2 0.2 0.1 0.1 0.1 0.5 – 0.2 0.3 0.2
CV Min 15.8 14.6 6.5 11.8 5.4 22.3 12.5 19.6 18.4 – 12.4 6.3 10.0 4.4 10.6 9.4 – 8.3 15.2 8.7 8.3 5.2 21.5 12.7 7.1 15.3 20.8 18.1 14.0 18.0 15.3 14.3 12.8 13.1 22.6 – 7.2 23.7 18.0 11.4 13.3 12.3 40.0 – 15.1 24.6 4.9
86.0 105.0 134.0 135.0 14.0 20.0 32.0 37.0 4.1 4.0 2.8 13.0 16.0 35.0 39.0 6.2 – 3.3 11.0 28.2 0.7 0.6 10.0 39.0 105.0 4.0 4.0 8.0 5.6 1.4 2.0 5.0 3.6 24.0 18.0 8.0 12.0 6.0 1.3 4.2 2.5 3.0 2.0 4.0 3.0 2.0 13.0
Max
n
147.0 160.0 168.0 214.0 17.0 37.0 51.0 66.0 7.4 5.0 3.9 16.0 22.0 41.0 54.0 8.0 – 4.3 17.0 38.0 0.9 0.8 20.0 57.0 136.0 6.4 8.0 14.0 9.4 2.6 3.0 8.0 5.9 38.0 35.0 – 16.0 23.0 2.0 5.6 3.7 4.0 4.0 6.0 5.0 5.0 16.0
12 9 12 13 12 9 12 13 9 >100? 13 12 9 12 13 9 >100? 13 9 13 9 13 7 12 12 12 9 12 12 12 9 12 12 12 9 >100? 12 13 4 12 12 12 4 >100? 12 13 12
Paramphisiella
357
Table 24 Continued Characteristics a Adoral membranelles, number
Species mean
ac2 ca1 ca2 ca3 Frontal cirri, number ac1 b ac2 b ca1 ca2 d ca3 f Buccal cirri, number ac1 ac2 ca1 ca2 ca3 Cirri left of anterior portion of amphisiellid ac1 b median cirral row, number ac2 b ca1 ca2 Amphisiellid median cirral row, number of ac1 cirri ac2 ca1 ca2 ca3 Cirri between posterior portion of marginal ca2 e rows, number Left marginal cirri, number ac1 ac2 ca1 ca2 ca3 Right marginal cirri, number ac1 ac2 ca1 ca2 ca3 Dorsal kineties, number ac1 ac2 ca1 ca2 Caudal cirri, number c ac1 ca1 ca2 ca3
15.1 35.0 31.8 32.5 3.0 3.0 3.0 4.2 4.1 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0
M
SD
15.0 – 32.0 32.0 3.0 3.0 – 4.0 4.0 1.0 1.0 – 1.0 1.0 1.0 1.0 –
SE
CV Min
Max
n 9 >100? 12 13 12 7 >100? 12 13 12 8 >100? 12 13 12 7 >100?
0.2 – 0.4 1.1 0.0 0.0 – 0.2 0.1 0.0 0.0 – 0.0 0.0 0.0 0.0 –
4.0 – 4.2 12.6 0.0 0.0 – 13.6 6.8 0.0 0.0 – 0.0 0.0 0.0 0.0 –
14.0 – 30.0 27.0 3.0 3.0 – 4.0 4.0 1.0 1.0 – 1.0 1.0 1.0 1.0 –
16.0 – 34.0 39.0 3.0 3.0 – 6.0 5.0 1.0 1.0 – 1.0 1.0 1.0 1.0 –
18.8 14.2 – 42.3 37.2 3.5
0.6 – 1.3 4.1 0.0 0.0 – 0.6 0.3 0.0 0.0 – 0.0 0.0 0.0 0.0 – usually 1 d 19.0 2.5 15.0 1.6 38.0 – 42.5 1.4 37.0 6.2 3.0 0.7
0.7 0.5 – 0.4 1.7 0.2
13.3 11.0 – 3.4 16.7 19.3
15.0 12.0 – 41.0 27.0 3.0
24.0 12 16.0 9 – >100? 46.0 12 48.0 13 5.0 12
44.6 32.6 – 41.0 33.2 45.9 34.3 – 44.6 41.4 3.0 3.3 3.0 3.0 3.0 3.0 3.3 3.0
45.0 33.0 30.0 40.5 34.0 45.5 34.0 – 44.0 43.0 3.0 3.0 – 3.0 3.0 – 3.0 3.0
1.6 0.5 – 1.0 1.6 1.6 0.5 – 0.8 1.8 0.0 0.2 – 0.0 0.0 – 0.2 0.0
12.8 4.6 – 8.1 17.6 12.4 4.1 – 6.2 16.0 0.0 14.9 – 0.0 0.0 – 19.5 0.0
36.0 30.0 24.0 37.0 24.0 35.0 33.0 37.0 41.0 29.0 3.0 3.0 – 3.0 3.0 – 3.0 3.0
55.0 35.0 35.0 48.0 45.0 54.0 36.0 39.0 50.0 50.0 3.0 4.0 – 3.0 3.0 – 5.0 3.0
5.7 1.5 – 3.3 5.9 5.7 1.4 – 2.7 6.6 0.0 0.5 – 0.0 0.0 – 0.7 0.0
12 9 >100? 12 13 12 9 >100? 12 13 12 7 >100? 12 12 >100? 12 3
a
All measurements in µm. CV = coefficient of variation in %, M = median, Max = maximum value, mean = arithmetic mean, Min = minimum value, n = number of individuals investigated (Hemberger 1985 wrote: “Basis: n = über 100 Individuen”), SD = standard deviation, SE = standard error of arithmetic mean. Data based on protargol-impregnated specimens. b
Foissner (1982) counted invariably four frontal cirri, that is, three frontal cirrus plus one cirrus (= cirrus III/2) left of anterior portion of amphisiellid median cirral row.
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SYSTEMATIC SECTION
Table 24 Continued c
See remarks for Shin’s (1994) data.
d
Usually this population has three frontal cirri (cirrus I/1, II/3, III/3) and one cirrus (= cirrus III/2) behind the right frontal cirrus; rarely, one or two further cirri occur left of the anterior portion of the amphisiellid median cirral row (see cell division). e
Designated as transverse cirri by Eigner & Foissner (1994).
f
Frontal cirri plus cirrus III/2 (rarely a further cirrus is obviously present).
(about 1950 m above sea level). He found it also in soil samples from another mountain (Stubnerkogel) nearby (Foissner & Peer 1985, p. 37; this paper also contains detailed autecological data). Further records: subalpine grassland field trial in Styria, Austria (Foissner et al. 1990, p. 18); alluvial soils from the Tullner Feld region, Lower Austria (Foissner et al. 1985, p. 107); soil from the Enns/Danube River floodplain, Upper Austria (Foissner 2004, p. 370); agricultural soils from near the village of Ostrov, southwest Slovakia (Tirjaková 1988, p. 499); mull from a mountain slope (about 3300 m above sea level) grown with shrubs from near the Pisang-Peak, Himalayan-region (Foissner 1986, p. 46; sample collected by Eduard Vierthaler, Filzmoos); terrestrial moss from a forest at Kimpo-up, Kimpo-gun, and from forest and grassland soil from Tokchok Is, Tokchok-myon, Ongijin-gun, Kyonggi-do, Korea (Shin 1994, samples 12 and 14; p. 258, 259). Not recorded during detailed surveys of Australian and Namibian soils (Blatterer & Foissner 1988, Foissner et al. 2002). Food not known. Biomass of 106 specimens about 18 mg (Foissner 1987, p. 121; 1998, p. 207).
Paramphisiella caudata (Hemberger, 1985) Foissner, 1988 (Fig. 71a–s, 72a–c, 73a–o, Table 24) 1982 Uroleptoides caudata n. sp.1 – Hemberger, Dissertation2, p. 54, Abb. 9a–k (Fig. 71a–s; see nomenclature; morphology and ontogenesis). 1985 Uroleptoides caudata n. spec. – Hemberger, Arch. Protistenk., 130: 402, Abb. 6 (Fig. 71a, b; original description; no formal diagnosis provided; type slides are deposited in the Institut für landwirtschaftliche Zoologie, University of Bonn, Germany). 1988 Paramphisiella caudata (Hemberger, 1985) nov. comb. – Foissner, Stapfia, 17: 121 (combination with Paramphisiella). 1994 Uroleptoides caudata Hemberger, 1985 – Shin, Dissertation, p. 56, Fig. 6A–C, Table 6 (Fig. 72a–c; description of Korean population). 1994 Paramphisiella caudata (Hemberger, 1985) Foissner, 1988 – Eigner & Foissner, J. Euk. Microbiol., 41: 247, Fig. 25–39, Table 2 (Fig. 73a–o; redescription and cell division; voucher slides [accession numbers 1994/12, 13] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Austria).
1 2
This name is disclaimed for nomenclatural purposes (ICZN 1999, Article 8.3). See next footnote. See corresponding footnote at Uroleptoides binucleatus.
Paramphisiella
359
2001 Paramphisiella caudata (Hemberger, 1985) Foissner, 1988 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 96 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Paramphisiella caudata – Lynn & Small, Phylum Ciliophora, p. 452, Fig. 36A–C (Fig. 73d, e; guide to ciliate genera).
Nomenclature: The species-group name caudat·us, -a, -um (Latin adjective [m; f; n]; having a tail, tailed) refers to the tailed posterior body portion (Hemberger 1985). Hemberger (1982, 1985) classified the present species in Uroleptoides, which is masculine (ICZN 1999, Article 30.1.4.4). Thus, Foissner (1988, p. 121) introduced the nomen corrigendum Uroleptoides caudatus. Lynn & Small (2002) used the term “midventral cirral file” for the designation of the amphisiellid median cirral row. This is somewhat misleading because the term midventral (cirri/rows/complex) should be confined to the urostyloids (for review, see Berger 2006) or non-urostyloid groups with zigzagging ventral cirri (e.g., Neokeronopsis; Berger 2006, p. 1190). Remarks: Hemberger (1985) did not study live specimens in detail. Thus, it is not known whether or not the specimens of the type population – which is very likely from a Peruvian soil – had cortical granules. The population studied by Eigner & Foissner (1994) is from Kenya and has distinct cortical granules (Fig. 73c). Consequently, it cannot be excluded that these populations, which are very similar in terms of the infraciliature, belong to different species. Thus, the descriptions are kept separate. A noteworthy difference in the cirral pattern of these two populations consists in the absence/presence of transverse cirri. According to Hemberger’s (1982, 1985) illustration (Fig. 71a) and description, interphasic specimens lack transverse cirri. However, during cell division the rearmost 2–3 cirri of the rightmost anlage (= anlage VI) migrate posteriorly to form a short row (Fig. 71n). Very likely, these cirri are resorbed in very late dividers and postdividers in the type population. By contrast, in the population described by Eigner & Foissner (1994) these cirri are retained in postdividers. However, only the rearmost cirrus of this row can be designated as transverse cirrus (Fig. 71d), because a transverse cirrus is the rearmost (often more or less distinctly enlarged and set off) cirrus of a frontal-ventral-transverse cirri anlage; usually several rearmost cirri form a distinct oblique [“transverse”] pseudorow; see general section and Berger 1999, 2006). The population studied by Shin (1994) agrees very well with the type material; unfortunately, Shin made no comment about the presence/absence of cortical granules. Interestingly, Shin (1994, p. 216, 220) found in a phenetic analysis that Uroleptoides caudata clusters with a uroleptid and a urostylid, indicating a serious mistake in the analysis because the present species lacks a midventral pattern, making such a relationship very unlikely. In addition, Shin neither described nor illustrated caudal cirri, which are, however, very small in the type population. Thus, one cannot exclude that Shin overlooked them. If the Korean population indeed lacks caudal cirri, then it is perhaps an Afroamphisiella.
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SYSTEMATIC SECTION
Fig. 71a–i Paramphisiella caudata (from Hemberger 1982. Protargol impregnation). a, b: Infraciliature of ventral side and nuclear apparatus, 200 µm. Body outline from a live specimen. Arrow marks cirrus III/2, which is formed from the same anlage as the right frontal cirrus. c–i: Infraciliature of ventral side and nuclear apparatus of early and middle dividers, c = 200 µm. Note that in this species and in the type species anlage IV is lacking. Details see text. The black transverse stripes in the macronuclear nodules are the replication bands. ACR = amphisiellid median cirral row, CC = caudal cirri, OP = oral primordium, I–III, V, VI = frontalventral cirri anlagen. Page 358.
Paramphisiella
361
Morphology: As mentioned above, it cannot be excluded that the Peruvian type population (Hemberger 1985) and the Kenyan population studied by Eigner & Foissner (1994) belong to different subspecies or species. Thus, the descriptions are kept separate. Unless otherwise indicated the data are from Hemberger (1985). Body size of type population in life(?) about 200 × 50 µm, body length:width ratio 4–5:1. Body outline variable, usually lanceolate with anterior end broadly rounded and posterior portion with narrowly rounded tail. Body very flexible and slightly contractile. Korean specimens 150–220 × 35–70 µm, very soft and flexible, elongate and slender, flattened dorsoventrally; anterior end slightly narrowed and rounded, posterior end tapered tail-like; ventral surface slightly concave, dorsal side convex. Nuclear apparatus in central body portion left of midline; macronuclear nodules ellipsoidal, micronuclei globular (Fig. 71b). Contractile vacuole behind proximal end of adoral zone (in specimen illustrated at 31% of body length; Fig. 71a), near left cell margin. Presence/absence of cortical granules not known. Cytoplasm and movement not described. Movement of Korean population rapid, changing direction frequently. For cirral pattern and variability, see Fig. 71a and Table 24; basically as in population described by Eigner & Foissner (1994; see below). Amphisiellid median cirral row extends to near last body fifth, in specimen illustrated it terminates at 70% of body length (Fig. 71a). Postperistomial cirrus and transverse cirri lacking (see remarks). Marginal cirri in life about 17 µm long. Dorsal cilia about 5 µm long, arranged in three, likely bipolar kineties (Fig. 71r). One caudal cirrus at end of each kinety; cirri fine because composed of four basal bodies/cilia only (Fig. 71r, s). Shin (1994) described and illustrated no caudal cirri (Fig. 72c). Body size of Kenyan specimens in life about 150 × 40 µm (Eigner & Foissner 1994). Body outline lanceolate, posterior end usually more distinctly tapered than anterior (Fig. 73a, b). Body highly flexible and able to contract to about 70% of usual body length. Macronuclear nodules in left body portion, in life about 10 × 6 µm. Contractile vacuole ahead of mid-body near left cell margin, in specimen illustrated at about 33% of body length (Fig. 73a); during diastole with indistinct collecting canals. Cortical granules arranged in conspicuous rows, 1.0–1.2 µm across, colourless (Fig. 73c). Cytoplasm colourless with lipid inclusions 1–5 µm across, yellowish crystals mainly in posterior body portion, and food vacuoles. Movement rather slow and clumsy. Adoral zone occupies about 25% of body length, composed of 32 membranelles of usual fine structure on average (Fig. 73d, Table 24). Bases of largest membranelles in life 7 µm wide. Buccal cavity flat and narrow. Buccal lip parallels slightly curved undulating membranes, left edge slightly thickened and wavy. Paroral and endoral about of same length and optically slightly intersecting. Pharyngeal fibres extend obliquely backwards (Fig. 73a, d). Cirral pattern and number of cirri of usual variability (Fig. 73d, Table 24). Frontal cirri enlarged, 15 µm long, arranged in oblique row with right cirrus, as is usual, near
362
SYSTEMATIC SECTION
Paramphisiella
363
distal end of adoral zone. Buccal cirrus slightly enlarged, right of anterior end of paroral. Usually one enlarged cirrus (= cirrus III/2) behind right frontal cirrus (likely rarely a second and/or third cirrus is/are present behind the right frontal cirrus because the maximum value of “frontal cirri” is six; no further details about variability given). Cirri of amphisiellid median cirral row and marginal rows about 10 µm long; row commences right of right frontal cirrus, extends obliquely backwards and terminates on average at 82% of body length near left marginal row (Table 24). A short longitudinal row composed of 3–5 cirri (designated as transverse cirri by Eigner & Foissner 1994) between rear portion of marginal rows (see remarks). Right marginal row extends onto dorso- Fig. 71r, s Paramphisiella caudata (from Hemberger 1982. lateral surface anteriorly, com- Protargol impregnation). Infraciliature of dorsal side of a mences about at level of distal middle and a late divider. Formation of dorsal kineties proend of adoral zone, terminates, ceeds in usual way, that is, two primordia are formed like left row, at rear cell end. Left within each parental kinety. At the posterior end of each kinety a small caudal cirrus – composed of four basal bodies marginal row more or less dis- each – is formed. CC = parental (r) and new (s) caudal cirri, tinctly curved rightwards anteri- 1–3 = dorsal kineties. Page 358. orly, commences left of proximal end of adoral zone; distance between cirri gradually increases posteriorly. Dorsal cilia 3 µm long, arranged in three kineties; kinety 3 posteriorly distinctly shortened in about 80% of specimens (Fig. 73e). Caudal cirri very small, in life 15–20 µm long; usually three cirri, rarely up to five cirri present; details about distribution not described; specimen shown in Fig. 73e with each two cirri on kineties 1 and 2. No caudal cirri described for Korean population (see remarks).
b Fig. 71j–q Paramphisiella caudata (from Hemberger 1982. Protargol impregnation). Infraciliature of ventral side and nuclear apparatus of middle, late, and very late dividers, sizes not indicated. Broken lines in (n) connects cirri which are formed by the same anlage (parental structures white, new black). Note that anlage IV is lacking in this species. I–III, V, VI = frontal-ventral cirral anlagen. Page 358.
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Fig. 72a–c Paramphisiella caudata (from Shin 1994. a, from life?; c, d, protargol impregnation). a: Ventral view, 200 µm. b, c: Infraciliature of ventral and dorsal side and nuclear apparatus, 200 µm. ACR = amphisiellid median cirral row, III/2 = cirrus behind right frontal cirrus (= paramalar cirrus), 1–3 = dorsal kineties. Page 358.
Cell division: This part of the life cycle is described in great detail both by Hemberger (1982; Fig. 71c–s) and Eigner & Foissner (1994; Fig. 73f–o). The two descriptions basically agree rather well, although they are difficult to compare because not exactly the same stages are described and illustrated. As discussed in the general section, Paramphisiella is a group with only five frontal-ventral-(transverse) cirri anlagen because it (usually) has only one cirrus (= cirrus III/2) left of the anterior portion of the amphisiellid median cirral row. This means that anlage IV, which forms the additional cirri left of the amphisiellid cirral row, is lacking. Eigner & Foissner (1994) did not use Wallengren’s designation of the cirral anlagen. Their anlagen 1, 2, and 3 are identical with Wallengren’s anlagen I, II, and III (see also general section); anlage 4 corresponds anlage V and anlage 5 is anlage VI. The follow-
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Fig. 73a–e Paramphisiella caudata (from Eigner & Foissner 1994. a–c, from life; d, e, protargol impregnation). a: Ventral view of a representative specimen, 150 µm. Note that this specimen has ingested, inter alia, one specimen of the colpodid ciliate, Colpoda inflata. b: Rare shape-variant. c: The colourless cortical granules are 1.0–1.2 µm across and arranged in longitudinal rows. d, e: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 151 µm. Dotted line connects frontal cirri. Broken line connects right frontal cirrus and cirrus III/2. Transverse arrow in (d) marks region where the anterior and posterior portion of the amphisiellid median cirral row overlap. Longitudinal arrow in (d) denotes the rear portion formed by the rightmost cirral anlage; Eigner & Foissner designated this row as transverse cirri; according to my opinion, only the rearmost cirrus of this short row can be designated, if at all, as true transverse cirri (details see text). Arrow in (e) marks rear end of dorsal kinety 3. ACR = rear end of amphisiellid median cirral row, AZM = adoral zone of membranelles, CC = caudal cirri, CV = contractile vacuole during diastole, FC = right frontal cirrus, MA = macronuclear nodule, MI = micronucleus, RMR = anterior end of right marginal row (extends dorsolaterally), 1–3 = dorsal kineties. Page 358.
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Fig. 73f–h Paramphisiella caudata (from Eigner & Foissner 1994. Protargol impregnation). f: Infraciliature of ventral side of an early divider (203 µm) showing proliferation of basal bodies left of the middle portion of the amphisiellid median cirral row. g: The oral primordium splits into a small front and a large rear portion. h: Dorsal view of an early divider, 125 µm. Dorsal morphogenesis commences with the formation of anlagen (arrow) within the parental rows. AZM = proximal end of (parental) adoral zone. Page 358.
Fig. 73i–l Paramphisiella caudata (from Eigner & Foissner 1994. Protargol impregnation). Infraciliature of ventral side of dividing specimens. i: Early divider, 140 µm. The anlagen I–III are formed by the oral primordium, anlage V develops from the amphisiellid median cirral row, and anlage VI possibly forms de novo. Arrow marks disorganising parental buccal cirrus. j: Middle divider, 175 µm. In this stage five primary primordia are recognisable. Arrows mark marginal row primordia of opisthe. k: Middle to late divider, 172 µm. The primary primordia divide and form five anlagen each in proter and opisthe. l: Late divider (160 µm) showing formation of cirri (dorsal side, see Fig. 73m). I–III, V, VI = frontal-ventral-(transverse) cirri anlagen. Page 358.
d
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SYSTEMATIC SECTION
Fig. 73m–o Paramphisiella caudata (from Eigner & Foissner 1994. Protargol impregnation). m: Infraciliature of dorsal side and nuclear apparatus of a late divider, 170 µm (same specimen as shown in Fig. 73l). Formation of new dorsal kineties and division of nuclear apparatus proceeds in ordinary way (see text for details). n, o: Infraciliature of dorsal and ventral side and nuclear apparatus of a very late divider, 158 µm. Arrows in (n) mark new caudal cirri (no cirri are formed at end of kinety 3). New structures black, parental white. Broken lines connect cirri which originate from same anlage (only shown for proter). Posterior cirral portion of anlage VI circled (designated as transverse cirral row by Eigner & Foissner). I–III, V, VI = frontal-ventral-(transverse) cirri anlagen. Page 358.
ing description is only an excerpt from the original papers, which have to be consulted when required. The oral primordium originates left of the middle portion of the amphisiellid median cirral row (Fig. 71c, 73f). A large field is formed and divides into a small anterior and a large posterior portion (Fig. 71e, g, 73g). The anterior portion extends to the proximal end of the parental undulating membranes. According to Hemberger (1982), a small anlage occurs right of the proximal end of the parental paroral. The amphisiellid median cirral row is obviously unchanged (Fig. 71e, g, 73g).
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The small anterior portion produces one short (anlage I) and two long (anlagen II, III) streaks in the African population (Fig. 73i). Anlage II extends to the parental buccal cirrus, which has modified to anlage II of the proter (Fig. 71h; 73i, arrow). The cirri of the central portion of the amphisiellid median cirral row are modified and form anlage V. Anlage VI is generated de novo (Hemberger 1982, Eigner & Foissner 1994) or is also formed by disorganised cirri of the amphisiellid row (Eigner & Foissner 1994). The organising centre of the amphisiellid row is obviously the site where the anterior and posterior portion of the anlagen V and VI aligned in the previous generation (Fig. 71h, 73i, j). In later stages, five long cirral streaks – so-called “primary primordia” (term introduced by Foissner 1983) – are formed. Streak I comprises basal bodies from the disorganising parental paroral and, possibly, from the opisthe’s oral primordium (Fig. 71h, 73j). Streak II develops from the oral primordium and connects with the anlage originating from the parental buccal cirrus. Streak III originates from the oral primordium and joins more or less distinctly with the anlage formed by the parental cirrus III/2 (Fig. 73j). Streak V develops from disintegrating cirri of the anterior segment of the amphisiellid median cirral row. Anlage VI possibly develops de novo or from the amphisiellid median cirral row too; anlagen V and VI form a V-shaped pattern (Fig. 73j). Somewhat later the primary primordia split about in the middle, producing five anlagen per filial product (Fig. 71j, 73k). In late and very late dividers, the cirri are formed and migrate to their final position (Fig. 71l, n, p, 73l, o). Anlage I forms the undulating membranes and the left frontal cirrus (= cirrus I/1); anlage II forms the middle frontal cirrus and the buccal cirrus; anlage III forms the right frontal cirrus and the cirrus behind (= cirrus III/2); anlage IV is lacking in Paramphisiella; anlage V forms the posterior portion of the amphisiellid median cirral row; anlage VI forms the anterior portion of the amphisiellid median cirral row (homologous to the frontoterminal cirri of the urostyloids, respectively, the cirri VI/3 and VI/4 of the 18-cirri hypotrichs) and some terminal cirri, which are arranged between the posterior portion of the marginal rows (only the posteriormost cirrus can be designated as transverse cirrus). The anterior portion of anlage VI aligns more or less perfectly with the cirri of anlage V, that is, in some interphasic specimens the site where the two portions meet is clearly recognisable (Fig. 73d, long arrow). According to Eigner & Foissner (1994), an additional anlage (likely anlage IV) occurs between the anlagen III and V in one or both filial products in few specimens to form one or two additional cirri left of the anterior portion of the amphisiellid median cirral row. The formation of the marginal rows and the dorsal kineties proceeds in the common (plesiomorphic) way, that is, two anlagen each occur within a parental row/kinety to form the new rows/kineties of the proter and the opisthe. At the rear end of each dorsal kinety anlage, a new caudal cirrus, composed of four basal bodies only,
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is formed. In the African population usually two cirri each are formed at the end of kineties 1 and 2 (Fig. 71h, j, l, n, p, r, s, 73h–o). The nuclear apparatus divides plesiomorphically way, that is, the macronuclear nodules fuse to a single mass and later divide (Fig. 71d, f, i, k, m, o, q, 73m, n). Occurrence and ecology: Paramphisiella caudata is basically terrestrial (Foissner 1987, p. 128), however, according to Foissner (1998, p. 207) also reliably recorded from freshwater habitats (according to the present review no published limnetic record is available; see below). So far not recorded from the Holarctic and Antarctic region (Foissner 1998). Type locality is very likely a soil sample from the Puerto Maldonado region (Madre de Dios), Peru. Unfortunately, Hemberger (1982, 1985) did not mention the exact site in the description; according to a personal communication, he found P. caudata in forest soil. Eigner & Foissner (1994) and Foissner (1999) found P. caudata at two sites in the Shimba Hills Nature Reserve (E39°25’ S5°). The population studied by Eigner & Foissner (1994) is from the upper 5 cm soil litter and sandy soil layer (pH 6.1) under a leguminose tree from near a forest surrounding the Sheldrick waterfalls; in addition, the present species occurred in a forest around a picnic site, where Foissner collected the upper litter and soil layer (pH 6.1). Foissner (1995, p. 39) found it in the upper 3 cm litter and soil layer from near a small path from the ranch house La Casona to the Pacific Ocean, Santa Rosa National Park in Costa Rica. The Korean population is from moss-covered soils and grassland at Namhansansong in Kwangju-gun, South Korea (Shin 1994, station 32, p. 259). Not recorded during detailed surveys of Australian and Namibian soils (Blatterer & Foissner 1988, Foissner et al. 2002). Paramphisiella caudata feeds on bacteria, heterotrophic flagellates (Polytoma sp.), and ciliates, like Colpoda inflata (Eigner & Foissner 1994). Biomass of 106 specimens about 300 mg (Foissner 1987, p. 128; 1998, p. 207).
Incertae sedis in the Amphisiellidae In “true” members of the Amphisiellidae, the amphisiellid median cirral row is formed by the anlagen V (forms posterior portion), IV (middle portion; usually not present), and VI (anterior portion) (see general section for further information). For the genera reviewed below, the formation of the frontoventral row is not known in detail, so they cannot be unequivocally assigned to the amphisiellids. If further studies show that the frontoventral row forms only from one anlage, like, for example, in Circinella (Foissner 1994), the corresponding genus has to be removed from the Amphisiellidae. The genera included in the present chapter are arranged alphabetically.
Afroamphisiella
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Afroamphisiella Foissner, Agatha & Berger, 2002 2002 Afroamphisiella nov. gen.1 – Foissner, Agatha & Berger, Denisia, 5: 698 (original description). Type species (by original designation): Afroamphisiella multinucleata Foissner, Agatha & Berger, 2002.
Nomenclature: Afroamphisiella is a composite of the Latin noun Africa and the genus-group name Amphisiella (see there for derivation), indicating that the type species was discovered in Africa (Foissner et al. 2002, p. 698). Feminine gender (Foissner et al. 2002). Characterisation (A = supposed apomorphy): Amphisiellidae(?) with slightly bipartite (type species) or continuous adoral zone of membranelles. Three frontal cirri. One buccal cirrus (type species) or more. One or more cirri left of anterior portion of amphisiellid median cirral row. Postperistomial cirrus and transverse cirri lacking (A). One left and one right marginal row. Two or three more or less bipolar dorsal kineties, that is, dorsomarginal kinety and kinety fragmentation lacking. Caudal cirri lacking (A?). Terrestrial. Additional characters: body flexible; contractile vacuole at left cell margin, about in mid-body, during diastole with collecting canals; dorsal bristles short, that is, less than 5 µm long. Remarks: The phylogenetic position of Afroamphisiella is uncertain (Foissner et al. 2002). We assumed that it belongs to the amphisiellids because the long cirral row (= amphisiellid median cirral row?) is involved in the formation of the oral primordium (Fig. 74k, 75d). Unfortunately, we could not find middle and late dividers showing the main feature of the amphisiellids, namely, that the frontoventral row is formed by the two rightmost anlagen, whose cirri align one behind the other (see chapter cell division below). In Orthoamphisiella, the long frontoventral row does not participate in the origin of the oral primordium and develops from a single primordium within the central portion of the parental row (Eigner & Foissner 1993). The two species included differ in the number of cirri left of the anterior portion of the amphisiellid median cirral row. The type species has only one cirrus on average, indicating that anlage IV is lacking. Afroamphisiella abdita has distinctly more cirri arranged in two rows and therefore likely does not lack anlage IV; perhaps Afroamphisiella is not monophyletic. Ontogenetic data are needed to get a deeper insight into the phylogeny of Afroamphisiella, which comprises two species discovered in the southern hemisphere. Afroamphisiella is mainly characterised by some cirri left of the anterior end of the amphisiellid median cirral row and the lack of transverse cirri, caudal cirri, and a postperistomial cirrus. This combination of features separates it from all other amphisiellids, which have transverse cirri and/or caudal cirri. 1
Foissner et al. (2002) provided the following diagnosis: Amphisiellidae with one or several cirri left of amphisiellid median cirral row, which is involved in oral primordium formation. Transverse cirri, caudal cirri, and postperistomial cirri lacking.
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Species included in Afroamphisiella (alphabetically arranged basionyms are given): (1) Afroamphisiella multinucleata Foissner, Agatha & Berger, 2002; (2) Lamtostyla abdita Foissner, 1997.
Key to Afroamphisiella species 1 2–4 macronuclear nodules; cortical granules lacking (Fig. 75a, c). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Afroamphisiella abdita (p. 377) - 14–29, on average about 18 macronuclear nodules; cortical granules present (Fig. 74a, j). . . . . . . . . . . . . . . . . . . . . . . . . . . . . Afroamphisiella multinucleata (p. 372)
Afroamphisiella multinucleata Foissner, Agatha & Berger, 2002 (Fig. 74a–k, m, n, Table 25) 1979 Uroleptoides kihni Wenzel, 1953 – Borror & Evans, J. Protozool., 26: 54, Fig. 13 (Fig. 74l; misidentification, see remarks). 2002 Afroamphisiella multinucleata nov. spec.1 – Foissner, Agatha & Berger, Denisia, 5: 699, Fig. 155a–k, 381j, k, Table 139 (Fig. 74a–k, m, n; original description; the holotype slide [accession number 2002/111] and two paratype slides [2002/112, 113] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; Foissner et al. 2002, p. 37 and Aescht 2003, p. 391).
Nomenclature: The species-group name multinucleata is a composite of the Latin quantifier mult·us (many), the thematic vowel ·i-, and the Latin adjective nucleát·us, -a, -um ([m; f; n]; kernel-like), referring to the many macronuclear nodules (Foissner et al. 2002). Type species of Afroamphisiella. Remarks: Uroleptoides kihni Wenzel sensu Borror & Evans (1979) is probably Afroamphisiella multinucleata because the nuclear and cirral pattern are very similar, even in details such as the beginning of the right marginal row at the level of the buccal cirrus (Fig. 74l). The only difference worth mentioning is the number of dorsal kineties: three in Borror & Evans’ population, usually two in the type population of A. multinucleata. However, we also found specimens with a short third row, indicating that this feature may be somewhat variable in this species. Indeed, the nuclear apparatus and the body size of A. multinucleata and Uroleptoides kihni (type of Uroleptoides) are very similar (Fig. 74a). However, they differ in body shape (elongate rectangular vs. slender, almost fusiform) and cirral pattern, such as length of amphisiellid median cirral row (55% of body length vs. almost of body length), number of cirri in frontal field (usually 5, rarely 6 or 7 vs. 10; cirri of 1
Foissner et al. (2002) provided the following diagnosis: Size about 85 × 13 µm in vivo; outline elongate rectangular. Cortical granules in distinct rows, yellowish, about 1.0 × 0.5 µm. On average 18 macronuclear nodules, 17 cirri in left and 21 in right marginal row, 14 cirri in amphisiellid median cirral, 1 cirrus behind right frontal cirrus, 1 buccal cirrus, and 2 dorsal kineties. Adoral zone composed of 17 membranelles on average, bipartited by an inconspicuous gap at left anterior margin of cell into a distal portion with 3–4 membranelles and a proximal portion with about 14 membranelles.
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Fig. 74a–h Afroamphisiella multinucleata (from Foissner et al. 2002. From life). a: Ventral view of a representative specimen, 85 µm. Note that the numerous macronuclear nodules are mainly arranged left of midline. b, c: The ellipsoidal cortical granules are about 1.0 × 0.5 µm, highly refractive, yellowish, and arranged in longitudinal rows both on ventral and dorsal side. d, e: Right lateral views showing dorsoventral flattening. f–h: Rare shape variants showing oral apparatus and contractile vacuole (f). BL = buccal lip, CV = contractile vacuole with collecting canals. Page 372.
amphisiellid row not included!), and cirri (caudal? transverse?) at posterior body end (both groups certainly lacking vs. at least one of these groups very likely present). Furthermore, the present species has distinct cortical granules, whereas such organelles are not described for U. kihni (lacking? overlooked?). Consequently, synonymy of A. multinucleata and U. kihni is rather unlikely. The congener Afroamphisiella abdita has four macronuclear nodules, usually two buccal cirri, and two short rows left of the amphisiellid median cirral row, which has on average about the same relative length as in A. multinucleata (55%). Orthoamphisiella stramenticola and O. grelli also lack transverse and caudal cirri; however, they have four, respectively, two macronuclear nodules against 14–29 in A. multinucleata (Eigner & Foissner 1991, 1993). Orthoamphisiella breviseries has a dumb-bell-shaped macronucleus and very short frontoventral rows not extending beyond buccal vertex (Foissner et al. 2002).
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Fig. 74i–k Afroamphisiella multinucleata (from Foissner et al. 2002. Protargol impregnation). i, j: Infraciliature of ventral and dorsal side and nuclear apparatus of holotype specimen, 70 µm. Arrow denotes the cirrus behind the right frontal cirrus (= cirrus III/2). Note that most cirri consist of four cilia only. k: Very early divider (90 µm), showing that the oral primordium originates from the posteriormost cirri (arrows) of the amphisiellid median cirral row. The arrowhead marks a gap in the adoral zone, which is also recognisable in the type specimen (i). Page 372. Fig. 74l Uroleptoides kihni sensu Borror & Evans (1979; protargol impregnation). This specimen is very likely an Afroamphisiella multinucleata (cp. with Fig. 74i), 53 µm. Note that even details (for example, position of buccal cirrus ahead of endoral; anterior end of right marginal row at level of buccal cirrus) are identical in these two illustrations. Morphometric data of this specimen, see Table 25. Page 372. ACR = posterior end of amphisiellid median cirral row, AZM = adoral zone of membranelles (bipartite by more or less distinct gap), BC = buccal cirrus, LMR = anterior end of left marginal row, MA = macronuclear nodule, MI = micronucleus, P = paroral, RMR = posterior end of right marginal row, 1, 2 = dorsal kineties.
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In life, Afroamphisiella multinucleata is easily identified by the following combination of features (Foissner et al. 2002): rows of yellowish, cortical granules; many macronuclear nodules; single, long frontoventral cirral row, one buccal cirrus; transverse cirri lacking. Morphology: For a brief morphometric characterisation of Borror & Evans’ specimen, see Table 25. Body size 70–100 × 18–30 µm in life, on average about 85 × 23 µm; body length:width ratio 3.1–4.5:1, on average 3.7:1 in life, while 3:1 in protargol preparations (Table 25). Body outline usually elongate rectangular, that is, with parallel margins and broadly rounded ends, rarely other shapes occur (Fig. 74f–h). Body very flexible, but acontractile, dorsoventrally flattened up to 1.5:1, ventral side flat, dorsal distinctly vaulted in mid-body or posterior half (Fig. 74a, d–h). Macronuclear nodules mainly left of midline (Table 25), ellipsoidal or dumb-bell-shaped, rarely globular, with few chromatin bodies 1–2 µm across. Micronuclei globular, attached or near macronuclear nodules. Contractile vacuole with two collecting canals, slightly ahead of mid-body at left cell margin. Cortical granules in longitudinal, rather widely spaced rows, do not impregnate with protargol, but when methyl green-pyronin is added, they become red, released, and swell to a voluminous, membranous coat; individual granules ellipsoidal, yellowish, highly refractive and thus conspicuous although only about 1.0 × 0.5 µm (Fig. 74b, c, m, n). Cytoplasm colourless, packed with granules about l µm across and, in posterior third, 2–5 µmsized lipid droplets. Food vacuoles about 6 µm across. Glides slowly on microscope slide and soil particles. Adoral zone occupies 26–42%, on average 31% of body length, composed of an average of 17 membranelles, bases of largest membranelles about 5 µm wide in life. Distalmost 3–4 membranelles separated from proximal portion of zone by more or less distinct gap (Fig. 74a, f, i, k; Table 25). Buccal cavity flat and narrow, anterior margin, however, rather distinct, forming bow with dorsal buccal wall (Fig. 74f). Buccal lip distinctly curved, covers right half of buccal cavity and proximal portion of adoral zone. Exact structure and arrangement of undulating membranes not unequivocally recognisable; paroral and endoral short and almost straight, parallel to each other with posterior, respectively, anterior portion slightly overlapping; paroral cilia about 8 µm long in life. Pharyngeal fibres clearly recognisable in vivo and after protargol impregnation, of ordinary length and structure, extend obliquely backwards. Cirral pattern very constant, number of cirri of usual variability (Fig. 74a, i, k; Table 25). Most cirri about 8 µm long in life, fine because usually composed of four cilia only. Three frontal cirri with left cirrus often slightly enlarged. One buccal cirrus ahead of anterior end of endoral. Usually one cirrus (= cirrus III/2; rarely two or three cirri) behind right frontal cirrus (the lack of further cirri right of cirrus III/2 indicate that anlage IV is lacking). Amphisiellid median cirral row begins right of right frontal cirrus and extends to body midline at 55% of body length on average; straight, that is, does not have, as in some other species, a slight bow or interruption
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Fig. 74m, n Afroamphisiella multinucleata (from Foissner et al. 2002. m, interference contrast; n, methyl greenpyronin staining). The yellowish cortical granules (1.0 × 0.5 µm) are arranged in longitudinal, widely spaced rows; when methyl green-pyronin is added, they swell to a voluminous, membranous coat. ACR = amphisiellid median cirral row, CG = cortical granules, RMR = right marginal row. Page 372.
where the row-fragments unite during late ontogenesis. Postperistomial cirrus and pretransverse ventral and transverse cirri absent. Marginal rows widely open posteriorly, right row begins near level of buccal cirrus and ends subterminally; left row usually commences at level of buccal vertex, slightly shorter than right posteriorly. Dorsal bristles about 3 µm long in life, arranged in two rows with distances between dikinetids increasing from anterior to posterior, rarely a short third row between anterior portion of row 2 and right body margin. Row l anteriorly slightly more shortened than row 2; posteriormost dikinetid in cell’s midline and thus not unequivocally assignable to one of the two rows. Caudal cirri absent (Fig. 74j). Cell division: We found only one divider (Fig. 74k). It showed that the oral primordium very likely originates from the posteriormost cirri of the amphisiellid median cirral row. According to Borror & Wicklow (1982), the cirral row of Uroleptoi-
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des originates from only one streak. However, it is unclear whether or not their statement on Uroleptoides is based on their U. kihni, which is likely identical with Afroamphisiella multinucleata (see remarks). More detailed cell division data are needed for a proper discussion. Occurrence and ecology: The type locality of A. multinucleata is a highly saline soil from the margin of the Etosha Pan, Namibia (Foissner et al. 2002). Borror & Evans (1979) found this or a very similar species in protargol slides made by E. Small during ecological investigations in the Chesapeake Bay estuary, Maryland, USA (Fig. 74l, Table 25; see also remarks). Feeds on bacteria (Foissner et al. 2002).
Afroamphisiella abdita (Foissner, 1997) Foissner, Agatha & Berger, 2002 (Fig. 75a–e, Table 25) 1997 Lamtostyla abdita n. sp.1 – Foissner, Biol. Fert. Soils, 25: 330, Fig. 20–24, Table 6 (Fig. 75a–e; original description; the holotype slide [accession number 1998/114; Aescht 2003, p. 379] and a paratype slide [1998/115] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Austria). 2001 Lamtostyla abdita Foissner, 1997 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 45 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Afroamphisiella abdita (Foissner, 1997) nov. comb. – Foissner, Agatha & Berger, Denisia, 5: 698 (combination with Afroamphisiella).
Nomenclature: The species-group name abdit·us, -a, -um (Latin adjective [m; f; n]; concealed, distant from) refers to the interphase cirral pattern, which would classify the present species in Orthoamphisiella (Foissner 1997, p. 331). Remarks: The present species resembles, according to the ventral interphase cirral pattern, Orthoamphisiella spp. because it has two short cirral rows left of the amphisiellid median cirral row and lacks transverse and caudal cirri (Eigner & Foissner 1991, 1993). However, two dividers (Fig. 75d, e) show that the long frontoventral row does not originate by within-row proliferation, as in Orthoamphisiella, but forms an anlage at its rear end, as is typical for Lamtostyla. Foissner (1997) already discussed that L. abdita could be a representative of a new genus because it lacks transverse cirri. The present species is very similar to Afroamphisiella multinucleata both in interphase (lack of transverse and caudal cirri) and early cell division (participation of amphisiellid median cirral row in oral primordium formation). Thus, we transferred it from Lamtostyla to Afroamphisiella (Foissner et al. 2002). Further ontogenetic
1
Foissner (1997) provided the following diagnosis: Size in vivo about 100 × 25 µm. Slenderly elliptical and slightly sigmoidal. Usually four macronuclear nodules. Two short rows and one long row of frontoventral cirri extending far beyond adoral zone of membranelles. On average 19 adoral membranelles, 32 right and 28 left marginal cirri, 3 frontal cirri, 2 buccal cirri, 17 cirri in right frontoventral row, 3 cirri in middle frontoventral row, and 2 cirri in left frontoventral row. Three dorsal kineties.
Fig. 75a–e Afroamphisiella abdita (from Foissner 1997. a, from life; b–e, protargol impregnation). a: Ventral view of a representative specimen, 100 µm. b, c: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 116 µm. Note lack of transverse and caudal cirri and discontinuity (arrow) in amphisiellid median cirral row, indicating that is composed of two segments (from two rightmost anlagen). d, e: Ventral infraciliature of early dividers, d = 106 µm, e = 103 µm. The pattern is similar to that found in Lamtostyla species. ACR = amphisiellid median cirral row, OP = oral primordium, 1–3 = dorsal kineties. Page 377.
378 SYSTEMATIC SECTION
Afroamphisiella
379
data are needed to show whether the amphisiellid median cirral row is formed by cirri originating from only one anlage or from two anlagen. Morphology: Body size in life 85–120 × 20–30 µm. Body outline inconspicuous, slenderly elliptical and indistinctly sigmoidal, usually widest close behind adoral zone and slightly narrowing posteriorly, both ends evenly rounded. Body dorsoventrally only slightly flattened, acontractile. Macronuclear nodules slightly ellipsoidal, with large chromatin bodies, arranged in two groups with two nodules each left of midline, second anterior nodule about of same size (11 × 7 µm, n = 17) as anterior nodule (Table 25); number of nodules highly variable, very likely because many postdividers were present, as indicated by the rather high proportion (about 20%) of specimens having one or two dividing nuclei; thus, four nodules is very probably the usual number. Micronuclei globular, usually one in each macronuclear group (Fig. 75a, c). Contractile vacuole near left cell margin about in mid-body, during diastole with two collecting canals. Cortex flexible, colourless, without special granules. Cytoplasm without crystals, usually containing many small and large globules and food vacuoles. Glides quickly on soil particles and slide. Adoral zone continuous, conspicuously short, that is, occupies only about 23% of body length on average, composed of an average of 20 membranelles; distal end extends only very slightly onto right body margin (Fig. 75a–c, Table 25). Bases of largest membranelles in life about 7 µm wide. Buccal cavity rather conspicuous because deep and distinctly curved. Paroral and endoral likewise curved and almost of same length, form wedge-shaped pattern because very close together anteriorly and widely separated posteriorly. Pharyngeal fibres conspicuous because long and numerous, extend obliquely backwards. Number of cirri of usual variability (Fig. 75b, Table 25). Cirri about 12 µm long, very thin. Three slightly enlarged frontal cirri almost transversely arranged; right one must not be interpreted as distalmost adoral membranelle. Usually two buccal cirri right of anterior half of paroral. Amphisiellid median cirral row commences near distal end of adoral zone, terminates in midline on average at 52% of body length (Table 25); frequently with small discontinuity in anterior third, indicating that it is composed of at least two fragments. Left of anterior portion of amphisiellid median cirral row usually two short rows not extending beyond adoral zone; left row originates from cirral anlage III, usually composed of two cirri; right row composed of 2–5, on average 3.4 cirri, obviously originating from anlage IV (note that this anlage is likely lacking in the type species!). Rarely specimens with a third row left of anterior portion of amphisiellid median cirral row (Fig. 75d). Pretransverse ventral cirri and transverse cirri lacking. Right marginal row commences slightly behind level of distal end of adoral zone, terminates somewhat ahead of posterior body end. Left row begins left of proximal end of adoral zone, terminates near rear body end; marginal rows distinctly separated posteriorly, cirri rather evenly spaced (Fig. 75c). Dorsal cilia about 3 µm long, arranged in three more or less bipolar kineties, which are rather evenly spaced. Row 2 more densely ciliated than rows 1 and 3 (Fig. 75c). Caudal cirri lacking.
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SYSTEMATIC SECTION
Table 25 Morphometric data on Afroamphisiella abdita (abd, from Foissner 1997) and Afroamphisiella multinucleata (mu1, type population from Foissner et al. 2002; mu2, specimen from Borror & Evans 1979, data from Fig. 74l) Characteristics a Body, length
Species
abd mu1 mu2 Body, width abd mu1 mu2 Body length:width, ratio mu1 mu2 Adoral zone of membranelles, length abd mu1 mu2 Body length:length of adoral zone, ratio mu1 mu2 Anterior body end to rear end of abd amphisiellid median cirral row, distance mu1 mu2 Anterior body to buccal cirrus, distance mu1 Anterior body end to right marginal row, mu1 distance Anterior body end to first macronuclear mu1 nodule, distance Nuclear figure, length mu1 Macronuclear nodule, length abd b mu1 b Macronuclear nodule, width abd b mu1 b Macronuclear nodules, number abd mu1 mu2 Macronuclear nodules, number right of mu1 midline mu2 Micronuclei, largest diameter abd Micronuclei, length mu1 Micronuclei, width mu2 Micronuclei, number abd mu1 mu2 Adoral membranelles, number abd mu1 mu2 Proximal adoral membranelles, number mu1 mu2 Distal adoral membranelles, number mu1 mu2 Frontal cirri, number abd mu1 mu2
mean
M
SD
SE
CV
Min
Max
n
97.9 71.5 53.0 28.9 24.0 15.0 3.0 3.5 22.2 22.1 17.0 3.2 3.1 51.2 39.3 33.0 10.3 9.2
97.0 72.0 – 29.0 24.0 – 3.0 – 22.0 22.0 – 3.3 – 49.0 39.0 – 10.0 9.0
10.1 6.5 – 2.9 2.4 – 0.3 – 1.3 2.0 – 0.3 – 6.7 5.2 – 1.3 2.0
2.5 1.2 – 0.7 0.4 – 0.1 – 0.3 0.4 – 0.1 – 1.6 1.0 – 0.2 0.4
10.3 9.1 – 9.9 9.9 – 9.3 – 6.0 9.2 – 10.2 – 13.1 13.2 – 12.4 21.4
80.0 117.0 59.0 84.0 – – 24.0 34.0 18.0 29.0 – – 2.5 3.7 – – 20.0 25.0 19.0 30.0 – – 2.4 3.8 – – 40.0 65.0 31.0 52.0 – – 7.0 15.0 6.0 15.0
17 29 1 17 29 1 29 1 17 29 1 29 1 17 29 1 28 29
8.8
9.0
2.4
0.4
26.9
4.0
13.0
29
57.6 10.6 5.5 7.3 3.0 3.3 18.4 14.0 5.2 6.0 2.2 2.1 2.1 2.2 2.8 3.0 19.6 17.5 18.0 14.2 15.0 3.3 3.0 3.0 3.0 3.0
58.0 10.0 6.0 7.0 3.0 3.0 17.0 – 5.0 – 2.0 2.0 2.0 2.0 3.0 – 19.0 17.0 – 14.0 – 3.0 – 3.0 3.0 –
5.3 1.8 1.1 0.9 0.4 0.7 3.7 – 1.5 – – – – – 1.1 – 1.1 0.9 – 1.0 – – – 0.0 0.0 –
1.0 0.4 0.2 0.2 0.1 0.1 0.7 – 0.3 – – – – – 0.2 – 0.3 0.2 – 0.2 – – – 0.0 0.0 –
9.2 16.6 19.1 12.5 14.1 19.8 20.0 – 28.8 – – – – – 38.3 – 5.4 5.0 – 6.9 – – – 0.0 0.0 –
47.0 8.0 3.0 6.0 2.0 2.0 14.0 – 2.0 – 2.0 1.5 1.5 2.0 1.0 – 18.0 16.0 – 13.0 – 3.0 – 3.0 3.0 –
66.0 14.0 7.0 9.0 4.0 4.0 29.0 – 8.0 – 3.0 3.0 2.5 3.0 6.0 – 22.0 19.0 – 16.0 – 4.0 – 3.0 3.0 –
29 17 29 17 29 30 29 1 29 1 17 28 28 17 28 1 17 29 1 29 1 29 1 17 28 1
Afroamphisiella
381
Table 25 Continued Characteristics a
Species
mean
M
SD
SE
CV
Min
Max
n
abd mu1 mu2 Cirri behind right frontal cirrus, number abd mu1 mu2 Cirri left of anterior portion of amphisiellid abd c median cirral row, number Amphisiellid median cirral row, number of abd cirri mu1 mu2 Left marginal cirri, number abd mu1 mu2 Right marginal cirri, number abd mu1 mu2 Dorsal kineties, number abd mu1 mu2 Dorsal kinety 1, number of kinetids mu1 Dorsal kinety 2, number of kinetids mu1 Dorsal kinety 3, number of kinetids mu3
1.5 1.0 1.0 2.1 1.3 2.0 3.4
2.0 1.0 – 2.0 1.0 – 3.0
– 0.0 – – – – 0.8
– 0.0 – – – – 0.2
– 0.0 – – – – 23.3
1.0 1.0 – 2.0 1.0 – 2.0
2.0 1.0 – 3.0 3.0 – 5.0
17 28 1 17 29 1 17
17.5 14.4 17.0 27.8 16.6 17.0 32.4 20.9 25 3.0 2.1 3.0 9.2 8.1 3.3
17.0 14.0 – 28.0 17.0 – 32.0 21.0 – 3.0 2.0 – 9.0 8.0 2.0
1.7 2.2 – 3.7 2.4 – 2.1 3.7 – 0.0 – – 1.5 1.2 –
0.4 0.4 – 0.9 0.4 – 0.5 0.7 – 0.0 – – 0.3 0.2 –
9.9 15.3 – 13.3 14.5 – 6.5 17.5 – 0.0 – – 16.8 15.1 –
14.0 11.0 – 22.0 11.0 – 30.0 13.0 – 3.0 2.0 – 6.0 6.0 2.0
21.0 22.0 – 34.0 22.0 – 37.0 34.0 – 3.0 3.0 – 13.0 10.0 6.0
17 29 1 17 29 1 17 29 1 17 29 1 27 27 3
Buccal cirri, number
a All measurements in µm. CV = coefficient of variation in %, M = median, Max = maximum value, mean = arithmetic mean, Min = minimum value, n = number of individuals investigated, SD = standard deviation, SE = standard error of arithmetic mean. Data based on protargol-impregnated specimens. b
Anterior macronuclear nodule.
c
Cirri behind right frontal cirrus not included.
Cell division: Foissner (1997) found two dividers (Fig. 75d, e). The oral primordium originates at two sites, namely at the posterior end of the amphisiellid median cirral row and likely in the area where the transverse cirri are located in other species (Fig. 75d). The somewhat later stage shows that some parental cirri (e.g., buccal cirri, cirri behind right frontal cirrus) are involved in the formation of the cirral anlagen (Fig. 75e). The discontinuity in the amphisiellid median cirral row (Fig. 75b, arrow) indicates that the row is composed of two portions, which very likely originate – as is usual – from the two rightmost anlagen. Occurrence and ecology: Very likely confined to terrestrial habitats (Foissner 1998, p. 205). Type locality of Afroamphisiella abdita is a rain forest near Cairns, Australia, where Foissner (1997) discovered it in the upper (2–5 cm) root carpet containing heavily decomposed litter mixed with brownish humic soil. Foissner (1997, p. 331) wrote that he has found it at two sites; however, in his Table 2 only sample 7 is marked.
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SYSTEMATIC SECTION
Afroamphisiella abdita feeds on rod-shaped bacteria, small hyphae, heterotrophic flagellates, and naked amoebas (Foissner 1997). Biomass of 106 specimens about 27 mg (Foissner 1998, p. 205).
Cossothigma Jankowski, 1978 1978 Cossothigma gen. n. – Jankowski, Tezisy Dokl. zool. Inst. Akad. Nauk SSSR, 1978: 40 (original description). Type species (by original designation and monotypy): Trachelostyla dubia Dragesco, 1954. 1979 Cossothigma Jk., 1978 – Jankowski, Trudy zool. Inst., Leningr., 86: 62 (generic catalogue). 1999 Cossothigma Jankowski, 1978 – Shi, Song & Shi, Progress in Protozoology, p. 150 (generic revision). 2001 Cossothigma Jankowski 1978 – Aescht, Denisia, 1: 49 (catalogue of generic names of ciliates). 2001 Cossothigma Jankowski, 1978 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 17 (nomenclator containing all basionyms, combinations, and higher taxa of euplotids and hypotrichs).
Nomenclature: Likely, no derivation of the name is given in the original description. Cossothigma is possibly a composite of cossus (Latin noun; woodworm, Cossus cossus = goat moth) and to thigma (Greek noun; touch, contact). I do not know to which feature this name should refer; possibly to the psammotic occurrence. Neuter gender because ending with the Greek suffix -ma (Aescht 2001, p. 50). Characterisation (A = supposed apomorphy): Amphisielidae(?) with trachelostylid body, that is, anterior portion (= head) narrowed, posterior portion narrowed or broadly rounded (A). Oral apparatus trachelostylid, that is, adoral zone along anterior and left margin of head. Three(?) frontal cirri. Several frontoventral cirri right of adoral zone (buccal cirri?). Ventral row extends obliquely from buccal vertex onto left dorsolateral side to near body end. Transverse cirri present. One longitudinal right and one spiralled left marginal row. Caudal cirri likely present. Marine. Additional characters: Body (very likely) flexible. Two ellipsoidal macronuclear nodules narrowly spaced in cell centre. Contractile vacuole and cortical granules likely lacking. Remarks: See type species. Species included in Cossothigma (alphabetically arranged basionyms are given): (1) Trachelostyla dubia Dragesco, 1954; (2) Trachelostyla sp. sensu Kahl (1932).
Key to Cossothigma dubium and a related species If you know that your specimen/population belongs to Cossothigma, then species determination is simple. Main features for the identification are the body size and the body shape. 1 Body length 200–650 µm; posterior body portion narrowed (Fig. 76a–d). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cossothigma dubium (p. 383)
Cossothigma -
383
Body length around 120 µm; posterior body end broadly rounded (Fig. 76e). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trachelostyla sp. sensu Kahl (1932; p. 386)
Cossothigma dubium (Dragesco, 1954) Jankowski, 1978 (Fig. 76a–d) 1954 Trachelostyla dubia n. sp. – Dragesco, Bull. Soc. zool. Fr., 79: 69, Fig. 3d (Fig. 76a; original description; no formal diagnosis provided and very likely no type material available). 1960 Trachelostyla dubia Dragesco – Dragesco, Trav. Stn biol. Roscoff, 12: 316, Fig. 167 (Fig. 76b; more detailed description). 1972 Gastrostyla dubia (Dragesco, 1954) n. comb. – Borror, J. Protozool., 19: 14 (combination with Gastrostyla; see remarks). 1978 Trachelostyla dubia Dragesco– Jankowski, Tezisy Dokl. zool. Inst. Akad. Nauk SSSR, 1978: 40 (combination with Cossothigma, see nomenclature. 1979 Trachelostyla dubia Dragesco, 1954 – Jankowski, Trudy zool. Inst., Leningr., 86: 62 (generic catalogue). 1985 Trachelostyla dubia Dragesco – Madrazo-Garibay & López-Ochoterena, Anales Instituto de Ciencias del Mar y Limnologia Universidad Nacional Autónomia de México, 12: 207, Fig. 31 (Fig. 76c; brief description of Mexican population). 1990 Trachelostyla dubia Dragesco, 1954 – Aladro Lubel, Martínez Murillo & Mayén Estrada, Manual de Ciliados, p. 134 and figure on same page (Fig. 76d; description of Mexican population, likely based on previous entry). 2001 Cossothigma dubium nom. corr. – Aescht, Denisia, 1: 50 (nomen corrigendum). 2001 Cossothigma dubium (Dragesco, 1954) Jankowski, 1978 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 90 (nomenclator containing all basionyms, combinations, and higher taxa of euplotids and hypotrichs).
Nomenclature: No derivation of the name is given in the original description. The species-group name dubi·us, -a, -um (Latin adjective [m, f, n]; doubtful, questionable, uncertain) possibly refers to the uncertain systematic position. Trachelostyla is feminine, Cossothigma is neuter. Thus, the species-group name had to be changed from dubia to dubium (Aescht 2001, p. 50; ICZN 1999, Articles 31.2, 34.2). Jankowski (1978) did not mention the name Cossothigma dubium; however, since he fixed it as type species of the present genus, he is automatically the combining author. Remarks: The present species is described after live observations only. Thus, the cirral pattern is not known in detail. Dragesco (1954, 1960) probably classified this huge ciliate (length up to 650 µm!) in Trachelostyla because the spindle-shaped body closely resembles that of T. caudata (cp. Fig. 76a, 101a). However, Trachelostyla lacks postoral ventral cirri, whereas the present species obviously has a distinct postoral ventral row; Trachelostyla pediculiformis, the type species, is an 18cirri hypotrich with anteriorly displaced postoral ventral cirri. Likely for that reason Borror (1972) transferred it to Gastrostyla Engelmann, 1862 (for review, see Berger 1999, p. 789). Indeed, the marine Gastrostyla stenocephala (Borror, 1963) Borror, 1972 (at present Hemigastrostyla stenocephala; for review, see Berger 1999, p. 933) has a similar appearance and cirral pattern, but the head is shorter and the transverse
384
SYSTEMATIC SECTION
cirri are more prominent, so synonymy of these two species can be excluded. Further, the oral apparatus – although not described in detail in both species – looks rather different so that the classification of the present species in Gastrostyla or Hemigastrostyla seems to be incorrect. Consequently, I (preliminarily) follow Jankowski (1978), who established Cossothigma for the present species, likely because of the deviating cirral pattern. There is no proposal for a higher level classification of the present genus/species, except by Dragesco (1960), who arranged it, like all other non-euplotid hypotrichs, in the Oxytrichidae. Shi et al. (1999) classified T. dubia as incertae sedis in the Hypotrichida. Hu & Song (2002, p. 178) correctly stated that further studies are needed for a final classification of C. dubium, whereas Gong et al. (2006) made no comment about the systematic position. I preliminarily assign it to the amphisiellids because of the single ventral row, which can also be interpreted as amphisiellid median cirral row, although it commences rather far posteriorly, which does not agree with the general appearance of an amphisiellid. The narrowed, trachelostylid anterior body portion is reminiscent of Trachelostyla (p. 474). However, the cirral pattern of the two type species (C. dubium, T. pediculiformis) is rather different so that a close relationship is very unlikely. Consequently, one has to assume that the cephalisation evolved convergently. Anyhow, detailed data about the ventral and dorsal ciliature including morphogenesis are needed for a more proper classification. The redescriptions by Madrazo-Garibay & López-Ochoterena (1985) and Aladro Lubel et al. (1990) match the data by Dragesco (1954, 1960) very well so that conspecificity of the French and Mexican populations is beyond reasonable doubt; however, note that Fig. 76d is likely a (superficial) redrawing of a previously published illustration (Fig. 76a, b). Both Dragesco (1954, 1960) and Madrazo-Garibay & López-Ochoterena (1985) described the species as very large (650 µm; 200–600 µm; 350 µm). By contrast, the number of cirri within the marginal rows and the ventral row seems rather normal, that is, similar to that of a normal-sized (100–200 µm) species. Whether this discrepancy is a special feature of this species or a misobservation (underestimation of number of cirri or overestimation of body size) is not known. The habitus of the present species is reminiscent of Stichotricha Perty, 1849. However, Stichotricha is characterised by two ventral rows and the lack of transverse cirri (e.g., Foissner et al. 1991, p. 203), whereas transverse cirri and only one ventral row are present in C. dubium. By contrast, Spirotrachelostyla lacks, like Trachelostyla, postoral ventral cirri (p. 502). Thus, the bimacronucleate S. simplex and S. tani must not be confused with C. dubium. Spiroamphisiella hembergeri is smaller (100–190 µm), has very prominent frontal cirri, and a second right marginal row, and the amphisiellid median cirral row commences near distal end of adoral zone (Fig. 27a–f). Cossothigma dubium differs from the specimen described by Kahl (1932) in body shape and size (further details, see Trachelostyla sp. below).
Cossothigma
385
Fig. 76a–d Cossothigma dubium (a, from Dragesco 1954; b, from Dragesco 1960; c, from MadrazoGaribay & López-Ochoterena 1985; d, from Aladro Lubel et al. 1990. a–d, from life). Ventral views, a = 650 µm, b = 417 µm, c = 350 µm, d = 424 µm (for a comment on body size see text). This species must not be confused with Spiroamphisiella hembergeri, Trachelostyla rostrata, and some bimacronucleate Spirotrachelostyla species, which lack postoral ventral cirri. Page 383. Fig. 76e Trachelostyla sp. (from Kahl 1932. From life). Ventral view, 120 µm. This specimen is very likely congeneric with Cossothigma dubium because of the identical cirral pattern (cp. with Fig. 76b) and nuclear apparatus (two closely spaced macronuclear nodules). Page 386. AZM = adoral zone of membranelles, CC = caudal cirri(?), DB = dorsal bristles, LMR = left marginal row, MA = macronuclear nodule, RMR = right marginal row, TC = transverse cirri, VR = ventral row.
Morphology: The description is mainly based on the data (illustration, description) by Dragesco (1954, 1960) unless otherwise indicated, and is supplemented by additional and/or deviating data from the Mexican populations. Body length according to Dragesco (1954) 650 µm, that is, very large; according to Dragesco (1960) 200–600 µm; body length:width ratio of specimens illustrated 5.9–6.3:1 (Fig. 76a, b). Mexican specimens about 350 × 50 µm (Madrazo-Garibay & López-Ochoterena 1985; Aladro Lubel et al. 1990); specimen shown in Fig. 76d, however, more than 400 µm long according to scale bar. Body spindle-shaped, that is, anterior portion (head) and posterior portion (tail) distinctly narrowed. Two narrowly spaced ellipsoidal macronuclear nodules in cell centre. Micronucleus neither described nor illustrated by Dragesco (1954, 1960); according to Madrazo-Garibay & López-Ochoterena
386
SYSTEMATIC SECTION
(1985) a single micronucleus between the two macronuclear nodules (original observation or misinterpretation of Dragesco’s illustrations?). Contractile vacuole and cortical granules neither mentioned nor illustrated by Dragesco; contractile vacuole according to Madrazo-Garibay & López-Ochoterena (1985) lacking. Movement not known. Oral apparatus trachelostylid, that is, adoral zone along anterior and left margin of narrowed head, extends to about 26% of body length (Fig. 76a, b). No details (e.g., number of membranelles, undulating membranes) known. Cirral pattern not known in detail. However, Kahl (1932, Fig. 76e) described the same pattern for a second species, proving that it is roughly correctly shown. Likely three frontal cirri; several (four in specimen shown in Fig. 76b) frontoventral cirri right of anterior portion of adoral zone. Spiral (oblique) row of ventral cirri (about 20 in number) extends from buccal vertex onto left dorsolateral side to near body end. Five transverse cirri, extend by about half of their length beyond rear cell end. Right marginal row commences at about 12% of body length, extends longitudinally (not spirally) to near right transverse cirri. Left marginal row commences near proximal end of adoral zone, extends spirally onto dorsolateral surface. Dorsal bristles obviously rather long (about 7 µm when for the specimen shown in Fig. 76b a body length of 300 µm is assumed); number of kineties and arrangement not known; at least rightmost kinety likely with caudal cirri (Fig. 76b). Occurrence and ecology: Marine; benthic, respectively, psammotic (Patterson et al. 1989, p. 211). Type locality of Cossothigma dubium is the small, intertidal Roscoff Aber Bay (France), Atlantic Ocean, where Dragesco (1954) found it in fine sediment (see also Dragesco 1960 and Bocquet 1971, p. 388). Madrazo-Garibay & López-Ochoterena (1985) found it in the Laguna de Términos, Campeche, Mexico (see also Aladro-Lubel et al. 1988, p. 446). Records not substantiated by morphological data: Romanian littoral of Black Sea (Petran 1968, p. 442; 1971, p. 154; see also Kovaleva & Golemansky 1979, p. 275). Food not known.
Trachelostyla sp. sensu Kahl (1932) (Fig. 76e) 1932 Trachelostyla-ähnliche Form mit ganz abweichender Bewimperung – Kahl, Tierwelt Dtl., 25: 597, Fig. 1142 (Fig. 76e; no material available).
Remarks: Kahl (1932) found only one specimen. It had a body shape very similar to that of T. pediculiformis, type of this genus. Because of the conspicuous differences (mainly in the cirral pattern) he was almost certain that it is not an anomalous specimen of T. pediculiformis. Thus, he classified it as uncertain form within Trachelostyla. As major differences he described the adoral zone, which extends distinctly onto the right side, and the spiralling ventral row and left marginal row. Jankowski (1979, p. 62) recognised the similarity of Cossothigma dubium and the present specimen. Indeed, their cirral pattern is almost identical although in both
Hemisincirra
387
cases details of the pattern are not known, for example, straight right marginal row, spiralling ventral row and left marginal row, caudal cirri on rightmost dorsal kinety. In addition, both have two narrowly spaced macronuclear nodules. Differences are in the shape of the rear body end (narrowed in C. dubium vs. broadly rounded in present specimen) and the body size (up to 650 µm vs. 120 µm). Thus, conspecificity is very unlikely. Trachelostyla rostrata has – like the present specimen and Cossothigma dubium – two macronuclear nodules, which are, however, distinctly separated (vs. narrowly spaced). Further, it lacks postoral ventral cirri (Fig. 76a, b). Detailed description (as new species) necessary. Morphology: Body size 120 × 26 µm (width calculated from Fig. 76e). Body outline more or less as in T. pediculiformis, that is, anterior body fourth (head) distinctly narrowed, body proper with parallel margins, and rear end broadly rounded. Two narrowly spaced macronuclear nodules in cell centre. Contractile vacuole neither illustrated nor mentioned, thus, possibly lacking. Cytoplasm honeycombed (Fig. 76e). Oral apparatus obviously trachelostylid, including short undulating membranes (details must not be overinterpreted); distal end of adoral zone extends onto right body margin. Three frontal cirri; five frontoventral cirri right of adoral zone; a ventral row composed of relatively strong cirri (about 17 in number) extends from buccal vertex to near body end; five transverse cirri, rightmost protrude distinctly beyond rear cell end; right marginal row commences about at 15% of body length, extends longitudinally (not spirally) to near right transverse cirri; left marginal row commences near proximal end of adoral zone, extends spirally onto left dorsolateral surface. Dorsal bristles about 3 µm long; number of kineties and their arrangement not known; at least rightmost kinety likely with caudal cirri (Fig. 76e). Occurrence and ecology: Marine. Kahl (1932) found this specimen in a culture from the Bay of Kiel (Germany), Baltic Sea.
Hemisincirra Hemberger, 1985 1982 Perisincirra Jankowski – Hemberger, Dissertation, p. 206 (redefinition of Perisincirra; see nomenclature and remarks). 1982 Perisincirra Jankowski, 1978 – Foissner, Arch. Protistenk., 126: 87 (discussion of genus and description of several species; see nomenclature and remarks). 1984 Hemisincirra Hemberger, 1984 – Foissner, Stapfia, 12: 119 (brief note and new combination of several species; see nomenclature). 1985 Hemisincirra n. gen.1 – Hemberger, Arch. Protistenk., 130: 408 (original description). Type species (by original designation): Uroleptus kahli Buitkamp, 1977. 1994 Hemisincirra, Hemberger 1985 – Tuffrau & Fleury, Traite de Zoologie, 2: 143 (classification). 1
Hemberger (1985) provided the following diagnosis which is the same as that by Hemberger (1982) for Perisincirra: Je 1 rechte und linke Marginalreihe; keine Cirren auf der postoralen Ventralfläche außer (meist 2) unscheinbaren Transversalcirren; Körper langgestreckt bis wurmförmig; Cilienzahl der Cirren reduziert; Morphogenesebeginn apokinetal unterhalb des Peristoms; 5 longitudinale FVT-Anlagen; zahlreiche Makronuklei.
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1997 Hemisincirra – Berger & Foissner, Arch. Protistenk., 148: 127 (brief note about systematic position). 1999 Hemisincirra Hemberger in Foissner, 1984 – Berger, Monographiae biol., 78: 893 (brief note about systematic position; see nomenclature). 1999 Hemisincirra Hemberger, 1985 – Shi, Song & Shi, Progress in Protozoology, p. 124 (generic revision of hypotrichs). 2000 Hemisincirra Hemberger, 1985 – Shi, Acta Zootax. sinica, 25: 12 (generic revision of hypotrichs; see remarks). 2001 Hemisincirra Hemberger in Foissner, 1984 – Aescht, Denisia, 1: 159 (catalogue of generic names of ciliates; see nomenclature). 2001 Hemisincirra Hemberger, 1985 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 30 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Hemisincirra Foissner, 1984 – Lynn & Small, Phylum Ciliophora, p. 458 (guide to ciliate genera; incorrect author and year).
Nomenclature: No etymology is given in the original description. The derivation is difficult because I do not understand the meaning of the middle part sin- in the present context; according to Werner (1972, p. 376), the stem sin- means damage or to damage. Most species now included in Hemisincirra were previously classified in Perisincirra (Hemberger 1982, Foissner 1982). However, since the type species of these two taxa are rather different, Hemberger (1985) established a new genus. Perhaps he simply replaced peri- by hemi- to get a different genus-group name. Feminine gender (Aescht 2001, p. 284). Incorrect subsequent spellings: Hemicinsirra sp. (Acosta-Mercado & Lynn 2003, p. 370); Hemisincira gellerti (Foissner) (Tirjaková 1991, p. 41). Foissner (1984) used the name “Hemisincirra Hemberger, 1984”, assuming that Hemberger’s paper would appear in 1984. He transferred several species previously classified in Perisincirra to Hemisincirra. However, the original description of Hemisincirra appeared in 1985, so that the combinations were made before Hemisincirra was available. To overcome this nomenclatural problem I proposed the authorship “Hemisincirra Hemberger in Foissner, 1984” (Berger 1999). Aescht (2001) accepted this proposal. In the catalogue of ciliate names (Berger 2001) I overlooked my 1999 comment and incorrectly assumed that Aescht (2001) had made this proposal. Simultaneously, I proposed accepting Hemberger (1985) as author, inter alia, because Foissner (1984) did not provide a diagnosis (Berger 2001). Further, “Hemisincirra Hemberger, 1985” would be a junior homonym and synonym of “Hemisincirra Hemberger in Foissner, 1984”. As a consequence of the rather confusing situation, Lynn & Small (2002) incorrectly considered Foissner (1984) as author of Hemisincirra. To bring this rather tricky situation to an end, I follow the proposal made by Berger (2001) and consider Hemberger (1985) as the author of Hemisincirra. Nomenclatural problems arising at the species Perisincirra gellerti, P. gracilis, and P. interrupta have already been solved by Berger (2001). For some further comments on the nomenclature of Perisincirra Jankowski, 1978 and Hemisincirra, see type species H. buitkampi Jankowski, 1979. Shi (1999, 2000) and Shi et al. (1999) put Perisincirra Jankowski, 1978 into the synonymy of Hemisincirra Hemberger, 1985, which is both nomenclaturally (disre-
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spect of the principle of priority) and taxonomically (Perisincirra and Hemisincirra are rather different; Hemberger 1985, Foissner 1984, Foissner et al. 2002) incorrect. Characterisation (A = supposed apomorphy): Amphisiellidae(?) with very elongate elliptical to vermiform body. Adoral zone of membranelles continuous or interrupted. Undulating membranes usually inconspicuous and thus difficult to recognise. Three frontal cirri. Buccal cirrus small (likely lacking in H. rariseta, H. interrupta, and H. vermicularis). Amphisiellid median cirral row about as long as or slightly longer than adoral zone of membranelles. Postperistomial cirrus lacking. Transverse cirri present, usually few in number. One right and one left marginal row. Most cirri composed of four or two cilia only. Three (inter alia, type species), one, two, or four dorsal kineties. Dorsomarginal kineties, dorsal kinety fragmentation lacking. Caudal cirri lacking (A). Terrestrial. Additional characters: body flexible; more than two macronuclear nodules; contractile vacuole near mid-body or ahead of it; dorsal bristles short (<4 µm). Remarks: Hemberger (1982) and Foissner (1982) described several species belonging to the present genus. Originally, they classified them in Perisincirra Jankowski, 1978. Hemberger (1982, p. 134, 206) incorrectly assumed that Uroleptus kahli Buitkamp, 1977 is the type species of Perisincirra. However, Jankowski (1978, 1979) unequivocally fixed U. kahli Grolière, 1975, the senior homonym, as type. Later, Hemberger (1985) recognised that Perisincirra – as defined via the type species U. kahli Grolière by Jankowski (1978, 1979) – is rather different from Perisincirra as defined by himself, and therefore he established Hemisincirra for U. kahli Buitkamp and his species. Since then, Hemisincirra has been a melting pot for slender, terrestrial hypotrichs with a short to moderately long, continuous or slightly irregular frontoventral row (Berger & Foissner 1997, p. 127). Foissner (1982, p. 87, 90) already recognised the heterogeneity of Perisincirra sensu Hemberger (1982) and correctly stated that ontogenetic data are needed for a more proper classification of some species. Berger & Foissner (1989), Foissner (1994), Berger (1999), Foissner et al. (2002), and Berger (2006) excluded some species from Hemisincirra and transferred them to Terricirra (p. 447), Circinella (Foissner 1994), Urosoma (Berger 1999, p. 396), Vermioxytricha (p. 596), Hemiurosoma (p. 614), and Caudiholosticha (Berger 2006, p. 232). In the present review, two species are transferred to Anteholosticha because they have a more or less distinctly zigzagging midventral complex (p. 640). In spite of these “purification-processes”, Hemisincirra is a difficult group because we cannot interpret the frontal-ventral cirral pattern of the type species due to the lack of ontogenetic data, that is, we do not know how the frontoventral row originates. The somewhat irregular arrangement indicates that at least two anlagen are involved (Fig. 77a, c). Several Hemisincirra species have distinct frontoterminal cirri (e.g., Fig. 80g), while in other species these cirri are possibly part of the amphisiellid median cirral row (Fig. 83a). The cirral and dorsal kinety pattern (1–4 kineties) of the species now included in Hemisincirra is heterogeneous, indicating that it is still a non-monophyletic assemblage. Consequently, the characterisation above
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is not very specific, that is, it is likely only a combination of more or less old plesiomorphies. As is usual, ontogenetic and meaningful molecular data are needed for a more proper classification. Previously, Hemisincirra (= Perisincirra sensu Hemberger 1982) was classified in the Oxytrichidae by Hemberger (1982; 1985, p. 405, 408), Foissner (1982), and Tuffrau & Fleury (1994). Shi (1999, 2000) and Shi et al. (1999) put it in the Gastrostylidae1. The oxytrichids are characterised by the dorsal kinety fragmentation (Berger 1999, p. 102; Berger 2006, Fig. 14a), and since the oxytrichids are a subgroup of the Dorsomarginalia, they also have a dorsomarginal kinety (Berger 2006, p. 33). Both features are obviously lacking in H. buitkampi (type species) and the other species classified in Hemisincirra, strongly indicating that Hemisincirra does not belong to the Dorsomarginalia (and therefore not to the oxytrichids). Of course, one cannot exclude that both features have been lost, but such an assumption is not in agreement with the principle of parsimony. Hemisincirra inquieta, a species very similar to the type species, has a frontoventral cirral pattern which is likely identical to that of 18-cirri oxytrichids, except that the postoral ventral cirri are displaced anteriorly (Fig. 78e, 79l). However, this is also not an argument for a classification in the oxytrichids – which comprise many 18-cirri hypotrichs (Berger 1999) – because the 18-frontal-ventral-transverse cirri pattern is an apomorphy of the hypotrichs (Berger 2006, p. 33ff), and not of the oxytrichids as incorrectly supposed in the first monograph (Berger 1999). As a consequence of the discussion above I exclude Hemisincirra from the oxytrichids (dorsal kinety fragmentation lacking) and even from the Dorsomarginalia (dorsomarginal kineties lacking). Since the type species and the other species have a more or less distinct frontoventral row likely originating from two anlagen, I put it preliminarily (that is, as incerta sedis) in the amphisiellids.2 Uroleptoides magnigranulosus, as species also classified in the amphisiellids (Fig. 52a–l), clusters outside the Dorsomarginalia in a molecular tree, but “unfortunately” in close relationship to some urostyloids (Schmidt et al. 2007, their Fig. 2). However, we know that molecular trees (like morphological) change relatively often and therefore this detail should not be over-interpreted. Lynn & Small (2002) assigned Hemisincirra, together with Trachelostyla (p. 474), Lamtostyla (p. 161), and Gonostomum (Berger 1999, p. 367), to the Trachelostylidae (p. 471). Trachelostyla pediculiformis, type of the group, is – according to a phylogenetic analysis inferred from the nuclear small subunit rRNA gene sequences – the sister group to all other hypotrichs (Schmidt et al. 2007, their Fig. 2;
1
The gastrostylids are, via the type species Gastrostyla steinii (see Berger 2001), a subgroup of the oxytrichids as clearly indicated by morphological features (dorsal kinety fragmentation; Berger 1999, p. 789). Later this position was confirmed by molecular data (e.g., Foissner et al. 2004, Schmidt et al. 2007). The rather rigid body and molecular data of G. steinii assigns it to the stylonychines. Further details, see Maregastrostyla. 2 As a consequence of the (preliminary) transfer from the oxytrichids to the amphisiellids I use the specific term “amphisiellid median cirral row” for the frontoventral row, although this row is not as continuous as the row of most true amphisiellids.
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note that the hypotrichs of the present review are the Stichotrichia in Schmidt et al. 2007). Vermioxytricha arenicola, type of Vermioxytricha Foissner, Agatha & Berger, 2002, closely resembles several Hemisincirra species (Fig. 126a–z). However, it forms a dorsomarginal kinety, a feature not described for Hemisincirra species, indicating that these two genera are not closely related. The habitus of Hemisincirra species is also very similar to that of Periholosticha species (for review, see Berger 2006, p. 498). However, these urostyloids lack a buccal cirrus (vs. present, although sometimes rather inconspicuous in Hemisincirra) and have caudal cirri (vs. lacking) and a zigzagging midventral complex (vs. absent). Species of Lamtostylides, a new genus established in the present book, have a wider body and only two macronuclear nodules. Hemisincirra species are very likely confined to terrestrial habitats (Foissner 1987, 1998, Cowling 1994, p. 30). Only occasionally, for instance after flooding, some specimens can be found in running waters (e.g., Packroff & Zwick 1996, p. 258; Tirjaková 2004, p. 7). Species included in Hemisincirra (alphabetically arranged basionyms are given): (1) Hemisincirra inquieta Hemberger, 1985; (2) Hemisincirra namibiensis Foissner, Agatha & Berger, 2002; (3) Hemisincirra octonucleata Hemberger, 1985; (4) Hemisincirra quadrinucleata Hemberger, 1985; (5) Hemisincirra rariseta Foissner, Agatha & Berger, 2002; (6) Hemisincirra vermicularis Hemberger, 1985; (7) Perisincirra gellerti Foissner, 1982; (8) Perisincirra interrupta Foissner, 1982; (9) Uroleptus kahli Buitkamp, 1977a. Incertae sedis: (10) Hemisincirra wenzeli Foissner, 1987. Species misplaced in Hemisincirra: The following species were originally assigned or transferred (mainly from Perisincirra) to Hemisincirra. However, various data indicate that some of them have to be classified in other genera. Hemisincirra gellerti verrucosa Foissner & Schade in Foissner, 2000. Remarks: Transferred to Anteholosticha in present book (p. 647). Hemisincirra heterocirrata Hemberger, 1985. Remarks: Transferred to Anteholosticha in present book (p. 540). Hemisincirra livida Berger & Foissner, 1987. Remarks: Now classified in Terricirra (p. 458). Hemisincirra muelleri Foissner, 1986. Remarks: Now classified in Vermioxytricha (see p. 607). Hemisincirra polynucleata Foissner, 1984, Remarks: Now classified in Hemiurosoma (p. 634). Hemisincirra vettersi Berger & Foissner, 1989, Bull. Br. Mus. nat. Hist. (Zool.), 55: 35, Fig. 48–55, Table 9. Remarks: Now classified in Circinella Foissner, 1994. The frontoventral row of Circinella arenicola Foissner, 1994, type of the genus, is formed from a single anlage. Thus, Circinella is not classified in the amphisiellids which form the row from two or three anlagen.
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Perisincirra filiformis Foissner, 1982, Arch. Protistenk., 126: 99, Abb. 27a–i, 69, 70, Tabelle 23. Remarks: Originally established as P. filiformis and later transferred to Hemisincirra by Foissner (1984, p. 119). However, this combination is invalid because the transfer was made before Hemisincirra Hemberger, 1985 was available. A correction is not necessary because at present this species is classified in Circinella Foissner, 1994. The frontoventral row of Circinella arenicola Foissner, 1994, type of the genus, is formed from a single anlage. Thus, Circinella is not classified in the amphisiellids which form the row from two or three anlagen. Perisincirra gracilis Foissner, 1982, Arch. Protistenk., 126: 95, Abb. 25a–g, Tabelle 22. Remarks: Transferred to Hemisincirra by Foissner (1984, p. 119; invalid combination) and by Foissner in Berger (2001, p. 71). Recently classified in Caudiholosticha Berger, 2003 (as C. gracilis) by Berger (2006) because it very likely has a midventral complex, that is, this species probably has more than the ordinary six (I–VI) anlagen and the cirri form a more or less distinct zigzag-pattern. For review see Berger (2006, p. 266). Perisincirra similis Foissner, 1982, Arch. Protistenk., 126: 94, Abb. 24a–f, 66, 72, Tabelle 21. Remarks: Now classified in Hemiurosoma (p. 633). Perisincirra viridis Foissner, 1982. Remarks: Now the type species of Terricirra (p. 450).
Key to Hemisincirra species and similar species Hemisincirra species are, like other slender species, difficult to identify. Detailed live observations (cortical granulation) and protargol preparations (cirral and dorsal kinety pattern) are usually needed for a reliable identification. If you cannot determine your specimen/population with the key below, see also Vermioxytricha (p. 596), Lamtostylides (p. 322), Periholosticha (Berger 2006, p. 498), Paragastrostyla (Berger 2006, p. 613), Caudiholosticha (Berger 2006, p. 232), or Circinella (Foissner 1994). The key also contains the two Hemisincirra species transferred to Anteholosticha in the present book (for review of Anteholosticha, see Berger 2006, p. 292). 1 Usually 4 macronuclear nodules (Fig. 82a, 83a). . . . . . . . . . . . . . . . . . . . . . . . . 2 - Usually 8 or more macronuclear nodules (e.g., Fig. 85a). . . . . . . . . . . . . . . . . . . 3 2 Two dorsal kineties; undulating membranes almost in parallel (Fig. 82b–d). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemisincirra namibiensis (p. 418) - Three dorsal kineties; undulating membranes at acute angle (Fig. 83a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemisincirra quadrinucleata (p. 419) 3 (1) About 8–10 macronuclear nodules (e.g., Fig. 85a, 134a, c, i). . . . . . . . . . . . . 4 - Invariably more than 10 macronuclear nodules. . . . . . . . . . . . . . . . . . . . . . . . . . . 7 4 Body length:width ratio about 16:1; body length about 200 µm; four contractile vacuoles (Fig. 88a–c). . . . . . . . . . . . . . . . . . . . Hemisincirra vermicularis (p. 435)
Hemisincirra 5 6 7 8 9 10 11
-
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Body length:width ratio about 5:1; body length below 120 µm; one contractile vacuole. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Three dorsal kineties; presence/absence of cortical granules not known (Fig. 84a, b). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemisincirra octonucleata (p. 421) Four dorsal kineties; cortical granules present (Fig. 85c, 134b, e, f). . . . . . . . . . 6 Cortical granules arranged in narrowly spaced, longitudinal rows, colourless (Fig. 85c). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemisincirra gellerti (p. 424) Cortical granules around cirri and dorsal bristles, colourless or yellowish (Fig. 134b, f). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha verrucosa (p. 647) (3) One dorsal kinety (Fig. 87a–f). . . . . . . . . . . . Hemisincirra interrupta (p. 432) Two or more dorsal kineties. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Two dorsal kineties (Fig. 86a–g, 133a, r). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Three dorsal kineties. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Body length:width ratio about 15:1 (Fig. 77a–d; note that Fig. 77a, b are incorrect in this respect). . . . . . . . . . . . . . . . . . . . . . . . Hemisincirra buitkampi (p. 393) Body length:width ratio about 7:1 (Fig. 78a, 79a, f, 80a, 89a). . . . . . . . . . . . . . 10 Caudal cirri present (Fig. 89a–h). . . . . . . . . . . . . . . Hemisincirra wenzeli (p. 437) Caudal cirri usually lacking (Fig. 78a, b, 79a–l, 80a–h, 81a–h). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemisincirra inquieta (p. 403) (8) Body length:width ratio on average 10:1, that is, body very slender; body length on average 170 µm; frontoventral row (amphisiellid median cirral row) slightly shorter than adoral zone (Fig. 86a–g). . . . Hemisincirra rariseta (p. 428) Body length:width ratio 5–6:1, that is, body elliptical; body length 100–140 µm; frontoventral row (midventral complex) slightly longer than adoral zone (Fig. 133a, b). . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha heterocirrata (p. 640)
Hemisincirra buitkampi (Jankowski, 1979) comb. nov. (Fig. 77a–c, Table 26) 1977 Uroleptus kahli n. spec. – Buitkamp, Decheniana, 130: 121, Abb. 4 (Fig. 77a, b; original description; see nomenclature; type slides are deposited in the Institut für landwirtschaftliche Zoologie und Bienenkunde, University of Bonn, Germany). 1979 Perisincirra buitkampi nom. n. – Jankowski, Trudy zool. Inst., 86: 84 (replacement name because of primary homonymy; see nomenclature). 1982 Perisincirra kahli (Buitkamp, 1977) n. comb. – Hemberger, Dissertation, p. 206 (revision of hypotrichs). 1985 Hemisincirra kahli (Buitkamp) – Hemberger, Arch. Protistenk., 130: 408 (combination with Hemisincirra). 1986 Perisincirra kahli (Buitkamp, 1977) Hemberger, 1981 – Dragesco & Dragesco-Kernéis, Faune tropicale, 26: 475, Planche 140, I, J (the illustrations are somewhat inexact redrawings of Fig. 77a, b and thus not shown in present review; guide to ciliates of tropical Africa). 2001 Perisincirra buitkampi Jankowski, 1979 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 71 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs).
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Nomenclature: The present species is the type species of Hemisincirra. Buitkamp (1977a) did not provide a derivation of the name, but obviously he dedicated the present species to Alfred Kahl, the great ciliatologist (e.g., Kahl 1930, 1931, 1932, 1935). Unfortunately, Buitkamp (1977a) overlooked that Grolière (1975) already established a species with the same name, that is, U. kahli Buitkamp, 1977a is a junior primary homonym (but not a synonym!) of U. kahli Grolière, 1975. Jankowski (1979) recognised this nomenclatural problem and therefore correctly introduced, according to Article 59a1 of the relevant code (ICZN 1964, p. 57), a replacement name (Perisincirra buitkampi) which is used in the present review as species-group name. However, since this species-group name was never used in Hemisincirra, it has to be transferred to this genus via a new combination (see heading). Hemberger (1982) knew – via a personal communication by Jankowski to Buitkamp who was, like Hemberger, at the Bonn University – that Jankowski has changed the name of Buitkamp’s species to Perisincirra buitkampi, but he did not know the second P. kahli species and therefore did not accept the replacement name proposed by Jankowski. Interestingly, Hemberger (1982) incorrectly mentioned “Perisincirra kahli (Buitkamp)” as type species of Perisincirra Jankowski, 1978, although Jankowski (1978, 1979) unequivocally fixed Uroleptus kahli Grolière, 1975 as type of Perisincirra. Hemisincirra buitkampi should be redescribed and the ontogenesis has to be studied so that Hemisincirra can be defined more precisely. Remarks: According to the text of the original description, the present species has a body length:width ratio of 15:1. By contrast, the illustration indicates a ratio of 9:1 (Fig. 77a). Hemberger (1982) checked the type slides of H. buitkampi and confirmed that the ratio of 15:1 is correct. The short, irregular amphisiellid median cirral row (four cirri plus one cirrus left of it) indicates that this species has the ordinary number of six frontal-ventral-transverse cirri anlagen (possibly even only five anlagen are present). For comparison with some other very similar Hemisincirra species (H. inquieta, H. wenzeli) see there and key above. According to Buitkamp (1977a), “Uroleptus (?) spec.” sensu Kahl (1932, p. 548, his Fig. 11021) is probably identical with the present species. This population is thus briefly described at the end of the morphology section (Fig. 77d). Morphology: Buitkamp (1977a) likely studied only few, possibly only one specimen because for meristic variables (e.g., number of macronuclear nodules or adoral membranelles) no variability is given. Body length (in life?) 80–180 µm, usually about 150 µm; body length:width ratio of life and prepared specimens 15:1 (see remarks for problems), that is, body vermiform. Consistency of cell (flexible or rigid) not mentioned, but very likely H. buitkampi is a very flexible species. 24 macronuclear nodules (variability not given) mainly in left body portion behind oral apparatus (Fig. 77b). Micronuclei not recognisable. Contractile vacuole, as is usual, 1
According to this article, a species-group name that is a junior primary homonym must be permanently rejected. The current code (ICZN 1999, Article 23.9.5) is less rigorous, but in the present case not relevant. Since the homonymy exists in Uroleptus, it would have been better to introduce the new speciesgroup name buitkampi in this genus and not in Perisincirra.
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395
Fig. 77a–c Hemisincirra buitkampi (from Buitkamp 1977a. Protargol impregnation). a, b: Infraciliature of ventral and dorsal side and nuclear apparatus, 90 µm. Note that the body length:width ratio is incorrectly illustrated (see text). Transverse cirri circled by dotted line. c: Detail of Fig. 77a. Frontal cirri connected by dotted line, cirri of amphisiellid median cirral row circled. Cirri (very likely) originating from same anlage connected by broken lines (only shown for anlagen I–III). Arrow marks buccal cirrus. AZM = distal end of adoral zone of membranelles, CV = contractile vacuole, LMR = left marginal row, MA = macronuclear nodule, RMR = right marginal row, 1–3 = dorsal kineties. Page 393. Fig. 77d Uroleptus (?) spec. sensu Kahl (1932). Ventral view, 200 µm. According to Buitkamp (1977a) it is likely identical with H. buitkampi. Page 402.
near left cell margin at 42% of body length in specimen illustrated (Fig. 77b). Presence/absence of cortical granules not known and cytoplasm (colour, inclusions) and movement not described. Adoral zone occupies about 17% of body length in specimen illustrated (Fig. 77a); note, however, that the length:width ratio of this specimen is incorrect, that is, this value must not be over-interpreted. Specimen illustrated with 13 adoral membranelles; distalmost three membranelles separated from each other and from proximal portion of zone. Undulating membranes almost in parallel and longitudinally arranged. Paroral two-rowed, endoral single-rowed (Fig. 77c). Cirral pattern as shown in Fig. 77a, variability, unfortunately not described. All cirri about 10 µm long and composed of four cilia, except for buccal cirrus which consist of two cilia only. Frontal cirri arranged in slightly oblique pseudorow. Buccal cirrus right of anterior end of paroral. Amphisiellid median cirral row composed of five cirri (cirrus left of it [= cirrus III/2] included), obviously arranged in an indistinct zigzag-pattern, indicating that it is formed not from a single, but likely from
396
SYSTEMATIC SECTION
Table 26 Morphometric data on Anteholosticha heterocirrata (het, from Hemberger 1982, 1985), Anteholosticha verrucosa (ve1, type population from Tenerife; ve2, Berlin population; both from Foissner 2000), Hemisincirra buitkampi (bui, from Buitkamp 1977a), Hemisincirra gellerti (gel, type population from Foissner 1982), Hemisincirra inquieta (in1, from Hemberger 1985; in2, from Berger & Foissner 1987; in3, from Berger & Foissner 1989; in4, from Foissner et al. 2002), Hemisincirra interrupta (int, from Foissner 1982), Hemisincirra namibiensis (nam, from Foissner et al. 2002), Hemisincirra octonucleata (oct, from Hemberger 1985), Hemisincirra quadrinucleata (qua, from Hemberger 1985), Hemisincirra rariseta (rar, from Foissner et al. 2002), Hemisincirra vermicularis (ver, from Hemberger 1982, 1985), Hemisincirra wenzeli (wen, from Foissner 1987a) Characteristics a Body, length
Body, width
Body length:width, ratio
Adoral zone of membranelles, length
Species mean bui gel het b in1 b in2 in3 in4 int nam oct b qua b rar ver b ve1 ve2 wen gel het b in1 in2 in3 in4 int nam oct b qua b rar ver b ve1 ve2 wen bui het nam oct qua ver gel in2 in3 in4
150.0 63.7 – 100.0 78.0 84.6 111.8 102.3 57.7 – – 143.2 200.0 64.4 60.5 75.9 12.6 – 15.0 10.1 12.6 14.9 9.3 11.7 – – 13.9 12.0 14.4 12.4 11.8 15.0 – 5.0 5.0 – 16.0 15.4 13.2 15.1 18.6
M
SD
SE
– 67.0 – – 75.0 90.0 108.0 106.0 57.0 – – 140.0 – 65.0 60.0 75.0 12.0 – – 10.5 13.0 15.0 10.0 11.5 – – 13.0 – 14.0 12.0 11.0 – – 4.8 – – – 15.2 13.0 15.0 18.0
– 8.0 – – 12.1 11.4 16.2 6.7 5.2 – – 17.6 – 11.5 5.7 8.1 1.3 – – 1.3 1.3 1.3 2.1 1.4 – – 1.1 – 1.9 2.1 2.0 – – 0.7 – – – 1.1 0.8 1.4 0.8
– 2.5 – – 3.8 4.3 4.5 2.5 1.3 – – 4.0 – 3.5 1.5 – 0.4 – – 0.4 0.5 0.4 0.8 0.4 – – 0.3 – 0.6 0.6 – – – 0.2 – – – 0.4 0.2 0.5 0.2
CV
Min
Max
n
– 80.0 12.6 50.0 – 100.0 90.0 15.5 63.0 13.4 70.0 14.5 77.0 6.5 93.0 9.1 48.0 – 95.0 – 110.0 12.3 115.0 – – 17.9 46.0 9.3 53.0 10.7 63.0 10.2 11.0 – 15.0 – – 12.7 7.0 10.1 11.0 8.6 13.0 22.8 7.0 11.8 10.0 – 18.0 – 15.0 7.9 13.0 – – 12.9 12.0 17.9 10.0 17.1 10.0 – – – 5.0 13.3 4.1 – – – 5.0 – – 7.5 14.0 5.9 12.0 8.9 13.0 4.1 18.0
180.0 79.0 140.0 125.0 100.0 98.0 140.0 112.0 68.0 120.0 130.0 170.0 – 90.0 70.0 89.0 15.0 25.0 – 11.0 14.0 17.0 12.0 14.0 25.0 20.0 16.0 – 18.0 18.0 16.0 – 6.0 6.5 – 6.0 – 18.0 15.0 17.0 20.0
? 10 >100 >100 10 7 13 7 17 15 >50 19 15 11 14 10 10 >100 >100 10 7 13 7 14 15 >50 19 15 11 14 10 ? >100 14 15 >50 15 10 10 7 13
Hemisincirra
397
Table 26 Continued Characteristics a Adoral zone of membranelles, length
Species mean
int nam rar ve1 ve2 wen Body length: length of adoral zone, ratio nam Anterior body end to paroral, distance in4 rar Paroral, length rar Anterior body end to amphisiellid median rar cirral row, distance Anterior body end to end of amphisiellid gel median cirral row, distance in2 in4 int nam rar ve1 o ve2 o wen Body length:length of amphisiellid nam median cirral row, ratio Anterior body end to buccal cirrus, nam distance Anterior body end to right marginal row, in4 distance nam rar Anterior body end to nuclear apparatus, in4 distance nam rar Posterior body end to last right marginal nam cirrus, distance Posterior body end to last left marginal nam cirrus, distance Anterior macronuclear nodule, length gel j int j nam rar j ve1 j ve2 j wen j Anterior macronuclear nodule, width gel j int j nam rar j ve1 j ve2 j wen j Posterior macronuclear nodule, length in2
M
SD
SE
CV
Min
Max
n
15.1 13.6 22.6 17.1 18.6 13.9 4.3 7.8 9.4 5.2 5.7
15.0 14.0 22.0 17.0 19.0 14.0 4.2 8.0 9.0 5.0 6.0
1.5 1.3 1.5 1.7 1.4 1.1 0.6 1.0 1.1 0.6 1.2
0.5 0.3 0.3 0.5 0.4 – 0.1 0.3 0.3 0.1 0.3
9.6 9.3 6.7 9.9 7.5 7.9 13.9 13.0 11.9 11.7 21.2
13.0 11.0 20.0 15.0 16.0 12.0 3.5 6.0 7.0 4.0 3.0
17.0 17.0 25.0 20.0 21.0 15.0 6.2 9.0 11.0 6.0 7.0
7 17 19 11 14 10 17 13 19 19 19
27.2 16.2 25.7 12.3 12.9 20.5 27.0 28.3 15.3 4.5
26.8 16.5 26.0 12.0 13.0 20.0 25.0 28.0 15.0 4.6
2.2 2.5 2.3 1.7 1.4 3.3 6.1 2.0 2.1 0.4
0.7 0.8 0.6 0.7 0.4 0.8 1.8 0.5 – 0.1
8.3 15.6 8.8 14.2 11.1 16.0 22.6 7.0 13.8 9.1
23.0 13.0 22.0 10.0 10.0 15.0 20.0 25.0 12.0 3.9
31.0 21.0 29.0 16.0 15.0 26.0 41.0 31.0 18.0 5.3
10 10 13 7 16 19 11 13 10 16
6.4
6.0
0.9
0.2
13.9
5.0
8.0
16
14.7 13.5 15.8 21.2 13.8 24.7 6.1
15.0 14.0 16.0 20.0 14.0 25.0 6.0
3.0 1.9 2.0 2.1 1.9 3.0 1.7
0.8 0.5 0.5 0.6 0.5 0.7 0.4
20.1 13.9 12.8 9.6 14.0 12.0 28.5
11.0 11.0 13.0 18.0 10.0 17.0 3.0
22.0 17.0 20.0 25.0 18.0 29.0 9.0
13 17 19 13 17 19 16
5.7
6.0
2.1
0.5
37.5
2.0
11.0
15
6.0 2.9 7.4 6.8 7.2 6.6 3.6 3.4 2.2 3.7 4.0 3.9 4.0 1.9 3.1
6.6 2.6 8.0 7.0 6.0 7.0 3.5 4.0 2.1 4.0 4.0 4.0 4.0 2.0 2.7
1.6 0.7 1.9 1.1 3.0 1.3 1.0 1.1 0.4 0.8 0.6 0.7 0.8 0.5 1.0
0.5 0.3 0.5 0.3 0.9 0.3 – 0.3 0.2 0.2 0.1 0.2 0.2 – 0.3
27.0 26.0 25.3 16.0 41.5 19.4 26.8 32.5 19.2 30.8 14.6 17.9 20.0 25.6 32.2
2.6 2.1 3.0 5.0 4.0 5.0 2.0 1.5 1.4 3.0 3.0 3.0 3.0 1.5 1.7
8.0 4.0 10.0 8.0 14.0 10.0 5.0 5.3 2.7 5.0 5.0 5.0 5.0 3.0 4.2
10 7 17 19 11 14 10 10 7 17 19 11 14 10 10
398
SYSTEMATIC SECTION
Table 26 Continued Characteristics a Posterior macronuclear nodule, length Posterior macronuclear nodule, width
Anterior micronucleus, length
Anterior micronucleus, width
Posterior micronucleus, length
Posterior micronucleus, width
Nuclear figure, length
Macronuclear nodules, number
Micronuclei, number
Species mean in3 in4 j in2 in3 in4 nam rar j ve1 j ve2 j wen m nam rar j ve1 j ve2 j in2 in3 in4 j in2 in3 in4 in4 nam rar bui gel het in1 in2 in3 in4 int nam c oct qua rar ver ve1 ve2 wen het in1 in2 in3 in4 nam d oct rar ver ve1 ve2
4.4 4.4 1.8 1.9 2.2 2.2 3.6 2.5 2.3 2.0 2.1 2.5 1.6 1.9 1.7 2.3 1.9 1.3 1.6 1.5 65.5 25.6 83.8 24.0 8.8 – – 27.7 31.7 28.3 28.9 4.1 8.0 4.0 14.9 10.0 8.7 8.1 28.8 – 2.0 2.0 2.1 2.8 1.9 2.0 1.9 2.0 1.9 1.9
M
SD
SE
CV
4.0 4.0 1.7 2.0 2.0 2.4 4.0 2.5 2.5 2.0 2.4 3.0 1.6 2.0 1.6 2.5 2.0 1.2 1.5 1.5 67.0 25.0 85.0 – 8.0 – – 29.0 32.0 30.0 30.0 4.0 – – 16.0 0 9.0 8.0 29.0 – – 2.0 2.0 3.0 2.0 – 2.0 – 2.0 2.0
1.3 1.1 0.3 0.4 – – – 0.5 0.5 0.1 – – 0.2 0.5 0.3 0.5 – 0.3 0.1 – 12.3 4.1 14.5 – 2.1 – – 3.6 2.1 4.0 3.9 0.6 – – 2.4 – 2.8 1.9 3.3 – – 0.0 0.7 0.9 – – 0.8 – 0.3 0.3
0.5 0.3 0.1 0.1 – – – 0.1 0.1 – – – 0.1 0.1 0.2 0.2 – 0.2 0.1 – 3.4 1.0 3.3 – 0.7 – – 1.2 0.8 1.1 1.5 0.1 – – 0.5 – 0.8 0.5 – – – 0.0 0.3 0.3 – – 0.2 – 0.1 0.1
28.7 25.6 18.3 19.2 – – – 19.6 20.0 4.3 – – 15.0 25.8 18.3 21.8 – 20.4 8.8 – 18.8 15.8 17.3 – 23.7 – – 13.2 6.7 14.1 13.5 14.6 – – 15.8 – 32.2 23.7 11.6 – – 0.0 32.2 33.5 – – 42.7 – 15.8 14.2
Min
Max
n
3.0 7.0 7 3.0 6.0 13 1.4 2.5 10 1.4 2.5 7 2.0 3.0 13 1.6 2.4 17 3.0 4.0 19 2.0 3.5 11 1.5 3.0 14 1.8 2.0 10 1.6 2.4 17 2.0 3.0 19 1.2 2.0 11 1.5 3.0 14 1.4 2.0 3 1.6 2.8 7 1.5 2.0 13 1.1 1.6 3 1.4 1.8 7 1.5 2.0 13 38.0 90.0 13 19.0 32.0 17 59.0 108.0 19 – – ? 6.0 13.0 10 12.0 14.0 >100 14.0 17.0 >100 22.0 32.0 10 30.0 36.0 7 19.0 32.0 13 21.0 33.0 7 3.0 6.0 17 8.0 8.0 15 4.0 8.0 >50 7.0 18.0 19 – – 15 5.0 16.0 11 5.0 13.0 14 24.0 34.0 10 2.0 3.0 >100 – – >100 2.0 2.0 3 1.0 3.0 7 1.0 4.0 13 1.0 2.0 17 – – 15 1.0 4.0 19 – – 15 1.0 2.0 11 1.0 2.0 14
Hemisincirra
399
Table 26 Continued Characteristics a Micronuclei, number Adoral membranelles, number
Frontal adoral membranelles, number
Ventral adoral membranelles, number Frontal ciliature, number of cirri g
Frontal cirri, number
Buccal cirri, number
Species mean wen bui gel het in1 in2 in3 in4 int nam e oct qua e rar e ver e ve1 ve2 wen bui nam oct qua rar ver nam rar bui s het in1 in4 oct qua ver gel k in2 in3 in4 int nam rar ve1 ve2 wen bui gel in2 in3 in4 nam h ve1 ve2
2.4 13.0 15.0 15.0 12.0 13.0 13.1 13.9 14.3 12.0 – – 15.9 12.0 15.2 15.6 12.4 3.0 4.0 4.0 – 3.1 4.0 8.0 12.9 8.0 13.0 11.0 14.1 13.0 9.0 10.0 4.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 2.9 3.0 1.0 1.0 1.0 1.3 1.0 0.9 1.0 1.0
M
SD
SE
CV
Min
Max
n
2.0 – 15.0 – – 13.0 13.0 14.0 14.0 12.0 – – 16.0 – 15.0 16.0 13.0 – 4.0 – – 3.0 – 8.0 13.0 – – – 14.0 – – 0 4.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 – 1.0 1.0 1.0 1.0 1.0 1.0 1.0
0.7 – 1.3 – – 0.5 0.7 – 1.3 0.0 – – 0.8 – 0.8 0.7 1.1 – 0.0 – – – – 0.0 0.8 – – – 1.1 – – – 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 – – – 0.0 0.0 0.8 0.0 – 0.0 0.0
– – 0.4
29.1 – 8.4 – – 3.6 5.3 – 8.9 0.0 – – 5.1 – 4.9 4.7 8.9 – 0.0 – – – – 0.0 6.3 – – – 7.9 – – – 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 – – – 0.0 0.0 61.2 0.0 – 0.0 0.0
2.0 – 13.0 – – 12.0 12.0 13.0 12.0 12.0 13.0 15.0 15.0 – 14.0 14.0 10.0 – 4.0 – 4.0 3.0 – 8.0 12.0 – – – 13.0 – 9.0 – 4.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 2.0 3.0 – 1.0 1.0 1.0 1.0 0.0 1.0 1.0
4.0 – 17.0 – – 14.0 14.0 14.0 16.0 12.0 15.0 16.0 17.0 – 16.0 17.0 13.0 – 4.0 – 5.0 4.0 – 8.0 14.0 – – – 16.0 – 11.0 – 4.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 – 1.0 1.0 3.0 1.0 1.0 1.0 1.0
10 ? 10 >100 >100 10 7 13 7 16 15 >50 19 15 11 14 10 ? 16 15 >50 19 15 16 19 ? >100 >100 13 15 >50 15 10 10 10 10 7 14 19 11 14 10 ? 10 10 7 10 17 11 14
– 0.1 0.3 – 0.5 0.0 – – 0.2 – 0.2 0.2 – – 0.0 – – – – 0.0 0.2 – – – 0.3 – – – 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 – – – 0.0 0.0 0.3 0.0 – 0.0 0.0
400
SYSTEMATIC SECTION
Table 26 Continued Characteristics a Buccal cirri, number Frontoterminal cirri, number t
Amphisiellid median cirral row, number of cirri
Transverse cirri, number
Left marginal cirri, number
Right marginal cirri, number
Species mean wen in4 nam rar ve1 ve2 gel in2 p in4 int nam f rar k ve1 o ve2 o wen bui gel het in1 in2 in3 in4 oct l qua nam i ve1 ve2 wen bui gel het in1 in2 in3 in4 int nam oct qua rar ver ve1 ve2 wen bui gel het in1 in2 in3
1.0 1.9 2.0 2.2 2.0 2.0 10.1 5.0 8.2 6.6 5.1 7.0 10.5 11.9 4.4 2.0 2.9 – 2.0 2.0 2.0 1.9 2.0 2.0 4.3 3.7 3.7 2.0 15.0 16.6 – 17.0 17.7 19.9 24.3 25.4 24.1 – – 25.1 54.0 15.3 15.7 18.3 15.0 15.9 – 18.0 16.8 21.4
M
SD
SE
CV
1.0 2.0 2.0 2.0 2.0 2.0 10.0 5.0 9.0 6.0 5.0 7.0 11.0 12.0 4.0 – 2.5 2.0 – 2.0 2.0 2.0 – – 4.5 4.0 4.0 2.0 – 16.0 – – 18.0 20.0 25.0 24.0 25.0 – – 25.0 – 15.0 16.0 19.0 – 15.5 – 0 17.0 21.0
0.0 – 0.0 – 0.0 0.0 2.1 0.0 1.1 0.7 0.8 1.7 2.0 1.8 0.5 – 1.1 – – 0.0 0.0 – – – 0.8 0.5 0.7 0.0 – 1.4 – – 2.0 2.0 4.2 3.8 2.3 – – 2.3 – 1.0 1.6 2.3 – 1.8 – – 1.9 2.3
– – 0.0 – 0.0 0.0 0.6 0.0 0.3 0.3 0.2 0.4 0.6 0.5 – – 0.4 – – 0.0 0.0 – – – 0.2 0.1 0.2 0.0 – 0.4 – – 0.6 0.8 1.2 1.4 0.6 – – 0.5 – 0.3 0.5 – – 0.6 – – 0.6 0.9
0.0 – 0.0 – 0.0 0.0 20.5 0.0 14.0 11.1 16.3 23.7 19.2 15.4 11.7 – 39.2 – – 0.0 0.0 – – – 18.0 12.7 17.6 – – 8.6 – – 11.0 10.3 17.4 15.1 9.4 – – 9.3 – 6.6 9.9 12.6 – 11.1 – – 11.5 10.7
Min
Max
n
1.0 1.0 1.0 2.0 2.0 2.0 2.0 3.0 2.0 2.0 2.0 2.0 6.0 14.0 5.0 5.0 7.0 10.0 6.0 8.0 3.0 7.0 5.0 11.0 7.0 13.0 9.0 15.0 4.0 5.0 – – 2.0 5.0 2.0 3.0 – – 2.0 2.0 2.0 2.0 0.0 2.0 – – 2.0 3.0 3.0 5.0 3.0 4.0 2.0 4.0 2.0 2.0 – – 15.0 20.0 18.0 21.0 16.0 19.0 15.0 22.0 17.0 23.0 16.0 30.0 20.0 30.0 20.0 27.0 14.0 16.0 20.0 24.0 21.0 29.0 – – 14.0 17.0 12.0 18.0 15.0 22.0 – – 14.0 20.0 17.0 20.0 16.0 19.0 13.0 20.0 18.0 24.0q
10 13 14 19 11 14 10 10 13 7 14 19 11 12 10 ? 10 >100 >100 10 6 13 15 >50 12 11 11 10 ? 10 >100 >100 10 7 13 7 16 15 >50 19 15 11 11 10 ? 10 >100 >100 10 7
Hemisincirra
401
Table 26 Continued Characteristics a
Species mean
Right marginal cirri, number
Dorsal kineties, number
Caudal cirri, number
in4 int nam oct qua rar ver ve1 ve2 wen bui gel het in1 in2 in3 in4 int nam oct qua rar ver ve1 ve2 wen in4 r wen n
22.7 27.0 20.7 – – 26.6 64.0 15.3 16.1 19.0 3.0 4.0 2.0 3.0 3.0 3.0 3.0 1.0 2.0 3.0 3.0 2.0 1.0 4.0 4.0 3.0 1.5 3.0
M
SD
SE
CV
Min
Max
n
23.0 28.0 22.0 – – 26.0 – 15.0 15.0 19.0 – 4.0 – – 3.0 3.0 3.0 1.0 2.0 – – 2.0 – 4.0 4.0 3.0 2.0 3.0
3.0 3.6 3.2 – – 3.5 – 1.4 2.8 1.3 – 0.0 – – 0.0 0.0 0.0 0.0 0.0 – – 0.0 – 0.0 0.0 0.0 0.8 –
0.8 1.4 0.8 – – 0.8 – 0.4 0.7 – – 0.0 – – 0.0 0.0 0.0 0.0 0.0 – – 0.0 – 0.0 0.0 0.0 0.2 –
13.1 13.3 15.3 – – 13.0 – 9.3 17.4 7.0 – 0.0 – – 0.0 0.0 0.0 0.0. 0.0 – – 0.0 – 0.0 0.0 – 53.1 –
16.0 20.0 15.0 12.0 22.0 21.0 – 13.0 13.0 16.0 – 4.0 – – 3.0 3.0 3.0 1.0 2.0 – – 2.0 – 4.0 4.0 3.0 0.0 2.0
26.0 13 32.0 7 25.0 17 15.0 15 24.0 >50 36.0 19 – 15 18.0 11 21.0 14 21.0 10 – ? 4.0 10 – >100 – >100 3.0 10 3.0 7 3.0 13 1.0 7 2.0 17 – 15 – >50 2.0 19 – 15 4.0 11 4.0 8 3.0 10 3.0 13 3.0 10
a All measurements in µm. CV = coefficient of variation in %, M = median, Max = maximum value, mean = arithmetic mean, Min = minimum value, n = number of individuals investigated, SD = standard deviation, SE = standard error of arithmetic mean, ? = sample size not indicated. Data based on protargolimpregnated specimens. b
Method (from life? from protargol preparations?) not indicated.
c
Of 17 specimens investigated, one specimen each had three (Fig. 82f), five (Fig. 82n), or six (Fig. 82d) macronuclear nodules.
d
Of 17 specimens investigated, two have one micronucleus.
e
Total number of adoral membranelles.
f
Of 14 specimens investigated, one specimen each had three (Fig. 82d), six, or seven cirri in this row.
g
All cirri included, except marginal cirri and transverse cirri.
h
Lacking in one out of 17 specimens investigated.
i
Including presumed pretransverse ventral cirri ahead of transverse cirri.
j
Macronuclear nodule/micronucleus measured not specified.
k
Cirrus (= cirrus III/2) behind right frontal cirrus included.
402
SYSTEMATIC SECTION
Table 26 Continued l
Of 15 specimens one had three cirri, and one had four.
m
Largest diameter of micronucleus.
n
The presence of caudal cirri makes the classification in Hemisincirra uncertain.
o
Anteholosticha verrucosa has (very likely) a midventral complex composed of several cirral pairs.
p
The two basal body pairs left of the anterior portion of the amphisiellid median cirral row are not included.
q
In Berger & Foissner (1989) 2.0 is incorrectly given as maximum value.
r
Identity of cirri uncertain (see description).
s
All cirri included, except buccal cirrus, transverse cirri, and marginal cirri.
t
In some species the frontoterminal cirri are likely included in the next feature.
two or three anlagen; however, ontogenetic data are needed for a correct interpretation; row terminates slightly ahead of level of proximal end of adoral zone (Fig. 77c). Postperistomial cirrus/cirri lacking. Two slightly subterminal transverse cirri. Right marginal row commences about at level of buccal cirrus, terminates – like left row – about at level of transverse cirri; left row begins near proximal end of adoral zone. Marginal cirri in both rows rather widely spaced because composed of only about 15 cirri per row. Dorsal bristles about 3 µm long, arranged in three rows; kineties 1 and 3 slightly shortened anteriorly (Fig. 77b). Dorsomarginal row and dorsal kinety fragmentation very likely lacking. Caudal cirri lacking. Kahl (1932) described a very slender moss-inhabiting species which is, according to Buitkamp (1977a), identical with H. buitkampi (Fig. 77d): body length 200 µm, ratio of body length:width about 21:1, that is, worm-like. Adoral zone occupies only about 7% of body length. Details must not be over-interpreted because such slender species are very difficult to observe. Occurrence and ecology: Hemisincirra buitkampi is very likely confined to terrestrial habitats (Foissner 1987, 1998) and reliable recorded from Holarctis and Palaeotropis (Buitkamp 1979). Records from aquatic localities are probably based on misidentifications. Type locality is the so-called Venusberg in the Melbtal area near the city of Bonn, Germany. Buitkamp (1977a) discovered it in the upper (0–5 cm) soil layer (brown soil) of two localities located side by side, namely a pasture dominated by Poa annua, P. pratense, Lolium perenne, Taraxacum officinale, and Trifolium repens, and a mixed forest dominated by Quercus robur, Fagus sylvatica, and Carpinus betulus. Buitkamp (1977, p. 252) found it also in soil from a “burnt savanna” and an “unburnt savanna” and a gallery woodland from near the city of Lamto, Ivory Coast. The highest abundance (186 specimens g-1 dry soil) occurred in samples from the mixed forest in Germany at a temperature of 15°C. The German populations occurred at 15–30°C whereas the populations from Ivory Coast occurred
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in samples kept at 20–35°C (Buitkamp 1979). Records not substantiated by morphological data: sediment-water interface from the crater lake of the “Tower Hill”, about 15 km west of Warnambool, Victoria, Australia (Finlay et al. 1999, p. 140; Esteban et al. 2000, p.163). Feeds on bacteria (Buitkamp 1977a). Biomass of 106 specimens about 8 mg (Foissner 1987, p. 124) to about 11 mg (Buitkamp 1979, p. 225).
Hemisincirra inquieta Hemberger, 1985 (Fig. 78a–o, 79a–l, 80a–h, 81a–h, Tables 26, 27) 1982 Perisincirra inquieta n. spec.1 – Hemberger, Dissertation2, p. 212, Abb. 37a–i (Fig. 78a–o; description of morphology and morphogenesis). 1985 Hemisincirra inquieta n. spec. – Hemberger, Arch. Protistenk., 130: 409, Abb. 15 (Fig. 78a, b; original description, no formal diagnosis provided; the type slide is deposited in the Institut für landwirtschaftliche Zoologie und Bienenkunde, University of Bonn, Germany). 1987 Hemisincirra inquieta Hemberger, 1985 – Berger & Foissner, Zool. Jb. Syst., 114: 207, Fig. 36–39, Table 7 (Fig. 79a–e; description of German population; a voucher slide is deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria). 1989 Hemisincirra inquieta Hemberger, 1985 – Berger & Foissner, Bull. Br. Mus. nat. Hist. (Zool.), 55: 34, Fig. 42–47, Table 9 (Fig. 79f–l; description of North-Iceland population; a voucher slide [accession number 1988:2:1:10] is deposited in the British Museum for Natural History in London, see nomenclature). 2001 Hemisincirra inquieta Hemberger, 1985 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 30 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Hemisincirra inquieta Hemberger, 19853 – Foissner, Agatha & Berger, Denisia, 5: 860, Fig. 186a–k, 401a–h, Tables 163, 164 (Fig. 80a–h, 81a–h; description of Namibian population and review; six voucher slides [accession numbers 2002/419–421, 423–425] have been deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria).
Nomenclature: No derivation of the name is given in the original description. The species-group name inquiet·us, -a, -um (Latin adjective [m; f; n]; disturbed, restless) is a composite of the prefix in+ (negation, inversion) and quietus (silent; undisturbed); likely the name refers to the movement. Berger & Foissner (1989, p. 19) wrote “One slide of holotype specimens and 1 slide of paratype specimens of the new species and 1 neotype-slide of each other species described have been deposited in the British Museum (Natural History) in London”. Unfortunately, this sentence is incorrect as concerns the neotypes because 1
This name is disclaimed for nomenclatural purposes (ICZN 1999, Article 8.3). See same footnote at Uroleptoides binucleatus. 3 Foissner et al. (2002, p. 863) provided the following definition (based on all descriptions): Size usually 90–120 × 10–15 µm in vivo, very elongate (7:1) with posterior body portion narrowed or bluntly pointed. Usually about 30 scattered macronuclear nodules. Cortical granules yellowish to orange, about 1 µm in size, found only around cirri and dorsal bristles. Adoral zone indistinctly bipartite, <20% of body length, composed of 12–14 membranelles. About 11 cirri in frontal field, 20 cirri in right marginal row, and 2 transverse cirri; frontoventral row about as long as adoral zone or slightly longer. Three dorsal kineties with row 1 consisting of only one to three kinetids. 2
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Table 27 Cortical granulation of Hemisincirra inquieta (from Foissner et al. 2002, based on original notes) Population (source)
Cortical granulation
Type population (Hemberger 1985) Austrian population (studied 1981; Foissner et al. 2002, p. 863) German population (Berger & Foissner 1987) Iceland population (Berger & Foissner 1989) Kenyan population 1 (studied 1985; Foissner et al. 2002, p. 863) Kenyan population 2 (studied 1985; Foissner et al. 2002, p. 863) Namibian population (Foissner et al. 2002)
feature not checked yellowish granules around cirral bases and dorsal bristles brilliant yellow, about 1 µm-sized granules around cirral bases and dorsal bristles orange-yellowish, about 1 µm-sized, ellipsoidal granules around cirral bases and dorsal bristles yellowish granule clusters few orange granules around cirral bases and dorsal bristles brilliant yellowish, compact, 1.0–2.0 × 0.8–1.0 µm-sized, ellipsoidal to ovoid granules in clusters between cirral bases and around dorsal bristles
it was not our intention to fix a neotype for each species redescribed, like, for example, Hemisincirra inquieta, whose original type slides are deposited in the Bonn University (Hemberger 1985, p. 409). In this and some other cases voucher slide would have been the correct term. For problems with the type locality, see occurrence and ecology section. Remarks: The original description of Hemisincirra inquieta is incomplete because Hemberger (1985) provided only limited morphometrics, did not describe details of the oral apparatus, and did not study live specimens so that we do not know whether or not the type population has cortical granules as described for all other populations (Berger & Foissner 1987, 1989, Foissner et al. 2002). The arrangement and colour of the cortical granules of H. inquieta is, as in many other hypotrichs, characteristic and important. Thus, we summarised the granules-data of six populations when we described the Namibian population (Foissner et al. 2002, p. 863; see Table 27). Obviously the arrangement around the dorsal bristles and cirri is largely constant, while the colour varies from yellowish over yellow to orange. The populations described so far agree well, although there are some differences in details and morphometrics (Table 26). Most differences concern the illustration and interpretation of the amphisiellid median cirral row and undulating membranes
Fig. 78a–h Hemisincirra inquieta (from Hemberger 1982. Protargol impregnation [body outline of specimen shown in Fig. 78a from life]). a, b: Infraciliature of ventral side and nuclear apparatus of interphasic specimen, 103 µm. Large arrow marks anterior end of right marginal row; small arrow denotes buccal cirrus composed of two basal bodies (cilia) only. c–h: Infraciliature of ventral side and nuclear apparatus of very early and early dividers, c = 103 µm. Arrow in (g) marks anteriorly extending anlage. Details see text. ACR = amphisiellid median cirral row, CV = contractile vacuole, MA = macronuclear nodule, MI = micronucleus, OP = oral primordium, RE = replication band, TC = transverse cirri. Page 403.
d
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(Foissner et al. 2002). As these minute structures are difficult to analyse, minor differences should not be over-interpreted. The Namibian specimens differ from those of the other populations mainly in the amphisiellid median cirral row, which is longer and composed of more cirri; however, the typical structure of the row, that is, alternately cirri composed four and two cilia, is the same, indicating conspecificity; possibly it is an African subspecies which is also characterised by indistinct caudal cirri (see morphology). The type population of H. inquieta forms the frontal-ventraltransverse cirri from the ordinary six (I–VI) anlagen (Fig. 78k). The Namibian population possibly has more than six anlagen as indicated by the rather long, somewhat irregular amphisiellid median cirral row; if this is true then it is possibly a holostichid related to Periholosticha species, which have, however, caudal cirri (possibly also present in Namibian specimens; Fig. 80f, g), but lack a buccal cirrus. Periholosticha paucicirrata, which is rather similar to H. inquieta, has only two dorsal kineties (for review on Periholosticha see Berger 2006, p. 498). As mentioned above, the frontal-ventral-transverse ciliature of H. inquieta is formed from six (I–VI) anlagen (Fig. 78k). It is interesting that in the populations described by Hemberger (1982, 1985) and Berger & Foissner (1987, 1989) in total 11 frontal-ventral cirri (frontal cirri, buccal cirrus, cirri and basal body pairs forming the amphisiellid median cirral row) are present (Fig. 78a, 79e, l); possibly H. inquieta has the plesiomorphic 18-cirri pattern of the hypotrichs (Berger 2006, p. 33), except that some (5) pretransverse ventral cirri and transverse cirri are lacking and that the three postoral ventral cirri are displaced anteriorly like, for example, in Gonostomum (for review see Berger 1999). The type population of H. inquieta has, like our populations, three dorsal kineties. However, kinety 1 of the type population is obviously of body length (Fig. 78o), whereas it is distinctly shortened posteriorly in all other populations studied so far (Fig. 79d, k, 80h). Perhaps, Hemisincirra inquieta is a species complex or consists of several subspecies differing in various features (amphisiellid median cirral row, colour of cortical granules, length of dorsal kinety 1). Further data are needed to answer this question properly. According to Hemberger (1985), Hemisincirra inquieta very closely resembles H. buitkampi, type of Hemisincirra, inter alia, because of the similar cirral pattern and the body length:width ratio of around 9:1 of the illustrated specimen (Fig. 77a, Fig. 78a, 79a). However, Hemberger (1982, 1985) re-studied the type material of H. buitkampi and found it to be much more vermicular (about 15:1; this value is also given in the description of H. buitkampi) than shown in Fig. 77a. Thus, Hemberger
b
Fig. 78i–o Hemisincirra inquieta (from Hemberger 1982. Protargol impregnation). i–n: Infraciliature of ventral side and nuclear apparatus of middle and late dividers (i–l) and an opisthe (m, n; note the parental transverse cirri at the rear cell end). Arrows in (i, j) mark two anlagen (very likely) forming the anlagen V and VI of opisthe and proter (details see text). o: Infraciliature of dorsal side and nuclear apparatus of a middle divider. Note that kinety 1 is of body length which is a distinct difference to the other populations, where it terminates ahead of mid-body (details see remarks and text). MI = micronucleus, TC = new transverse cirri, I, VI = frontal-ventral-transverse cirri anlagen, 1–3 = dorsal kineties. Page 403.
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established a new species for his population. Interestingly, the dorsal kinety pattern of the type population of H. inquieta is more similar to that of H. buitkampi than to that of the other H. inquieta populations (see above). Berger & Foissner (1987) described a German population which agrees very well with the type material, except for the number of macronuclear nodules (22–32 in German population vs. 14–17 in Hemberger’s population). We also discussed the similarity of H. buitkampi and the present species (see below) and came to the conclusion that further populations have to be investigated to decide whether H. buitkampi and H. inquieta are real species or extremes of a polytypical species. Since then, two further population of H. inquieta have been studied, but no population of the type species was found, so the situation basically did not change. The Iceland population studied by Berger & Foissner (1989) has a more tapered body, but the remaining features agree very well with the populations studied previously. The illustration of Hemisincirra buitkampi (Fig. 77a) agrees rather well with those of H. inquieta (Fig. 78a, 79c, j, 80g). However, there are some differences which indicate that these two species are not synonyms, although it is astonishing that H. buitkampi was never recorded since its description in 1977. The differences exist in (i) the anterior end of the right marginal row (almost at level of frontal cirri in H. buitkampi vs. about at level of buccal vertex in H. inquieta); (ii) the length of dorsal kinety 1 (extends far posteriorly and composed of 6 bristles in specimen illustrated vs. terminates at mid-body or ahead of it and composed of 1–3 bristles [note, however, that the type population of H. inquieta very likely also has a long kinety 1; Fig. 78o]); (iii) basal body pairs left of anterior portion of amphisiellid median cirral row (absent vs. present); (iv) body length:width ratio (15:1 [note that Fig. 77a is incorrect in this respect] vs. about 6–9:1). Whether or not H. buitkampi has cortical granules is not known; unfortunately, this also applies to the type population of H. inquieta. Hemisincirra wenzeli (Fig. 89a–h) is also very similar to H. inquieta. However, it has colourless cortical granules (vs. yellowish to orange) and distinct caudal cirri [vs. lacking [Fig. 78o, 79d, k], respectively, inconspicuous in Namibian population [Fig. 80f, g]). For separation from other Hemisincirra species, see key. Morphology: This section begins with the detailed description of a German population investigated by Berger & Foissner (1987). Additional and/or deviating data from the populations described by Hemberger (1985) and Berger & Foissner (1989) are added below. The Namibian population described by Foissner et al. (2002) has an amphisiellid median cirral row which is longer than that of the other populations; possibly it is an African subspecies (see remarks); thus, its description is kept separate. Data on cortical granules are summarised in Table 27. Population described by Berger & Foissner (1987; Fig. 79a–e; Table 26): Body size in life about 80–100 × 14–15 µm (n = 2); body length:width ratio 7.7:1 in protargol preparations on average (Table 26). Body outline elongate, cylindrical, margins parallel or slightly converging anteriad and posteriad, both ends rounded; body shape very stable within the population. Body very fragile. On average 28 macronu-
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Fig. 79a–e Hemisincirra inquieta (from Berger & Foissner 1987. a, b, from life; c–e, protargol impregnation). a: Ventral view of a representative specimen, 102 µm. Arrow marks contractile vacuole. b: Ventral view showing arrangement of cortical granules which are less than 1 µm across and bright yellow. c–e: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 94 µm. Arrow in (d) marks rear end of dorsal kinety 1. Short arrows in (e) mark cirri which are very likely homologous to the postoral ventral cirri of the 18-cirri hypotrichs; cirri which are very likely homologous to the frontoventral cirri are circled by dotted line (for terminology see Fig. 6a in Berger 1999). Note that this pattern is basically identical to that shown in Fig. 78a and 79l. Long arrow marks gap in adoral zone. Frontal cirri connected by dotted line. Broken lines connect cirri which very likely originate from same anlage (has to be confirmed or disproved by ontogenetic data). AZM = distal end of adoral zone of membranelles, LMR = left marginal row, MA = macronuclear nodule, MI = micronucleus, RMR = right marginal row, TC = transverse cirri, I, III, V, VI = frontal-ventral-transverse cirri anlagen, 1–3 = dorsal kineties. Page 403.
clear nodules mainly arranged in the second and third quarter of cell (Fig. 79d). Contractile vacuole slightly ahead of mid-body, in specimen illustrated at 38% of body length (Fig. 79a). Pellicle and cytoplasm colourless. Cortical granules arranged in groups around bases of cirri and dorsal bristles; groups composed of 3–6 granules which are bright yellow and small, that is, less than 1 µm across. Thus, specimens are brownish at low magnification. Middle portion of cell filled with 1–4 µm-sized,
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Fig. 79f–l Hemisincirra inquieta (from Berger & Foissner 1989. f–i, from life; j–l, protargol impregnation). f: Ventral view of representative specimen, 115 µm. g: Cortical granules (about 1 µm across, orange-yellow) around dorsal bristles. h, i: Ventral views of shape variants, h = 88 µm, i = 93 µm. j–l: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 78µm. Arrow in (j) marks anterior end of right marginal row. Dotted line in (l) connects frontal cirri; broken lines connect cirri very likely originating from same anlage (has to be confirmed or disproved by ontogenetic data). Rectangle encloses cirri which are very likely homologous to the frontoventral cirri of the 18-cirri hypotrichs, ellipse encloses cirri very likely homologous to the postoral ventral cirri of the 18-cirri hypotrichs (see Fig. 6a in Berger 1999). Note that in H. inquieta only the anlage V and VI form each one transverse cirrus. ACR = posterior end of amphisiellid median cirral row (basically formed by cirri of anlagen V [posterior portion] and VI [frontoterminal cirri; anterior portion]), CV = contractile vacuole, TC = transverse cirri, I–VI = frontal-ventral-transverse cirri anlagen, 1–3 = dorsal kineties. Page 403.
colourless globules. In the posterior portion some yellowish crystals. No food vacuoles recognisable. Adoral zone of membranelles occupies 17% of body length on average in protargol preparations (Fig. 79a–c, e, Table 26). Distalmost three membranelles separated from proximal portion of adoral zone by distinct gap. Buccal area slightly deepened, undulating membranes moderately bent. Pharyngeal fibres distinct both in life and in protargol preparations.
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Cirral pattern and number of cirri of usual variability (Fig. 79a, c, e, Table 26). Frontal cirri in almost transverse pseudorow immediately behind distalmost adoral membranelles. Buccal cirrus near anterior end of undulating membranes, inconspicuous because composed of two cilia only. Amphisiellid median cirral row terminates at 21% of body length on average and thus slightly longer than adoral zone; row invariably (n =10) composed of five cirri (Table 26) with second anteriormost cirrus shifted slightly leftwards; behind and ahead of this cirrus two basal body pairs (for an in-depth interpretation of cirral pattern, see Fig. 79e). Two inconspicuous transverse cirri at rear cell end, thus projecting distinctly beyond body proper. Right marginal row begins right of posterior end of amphisiellid median cirral row, terminates – like left row – slightly ahead of rear body end. Cirri of amphisiellid median cirral row and marginal rows about 10 µm long in life and composed of four cilia. Dorsal bristles about 3 µm long, invariably arranged in three kineties; kinety 1 composed of two or three bristles only. Bristles generally rather widely spaced within kineties. Caudal cirri lacking (Fig. 79c). Additional and/or deviating data from populations described by Hemberger (1985; Fig. 78a, b, Table 26) and Berger & Foissner (1989; Fig. 79f–l, Table 26): body size 115 × 15 µm in life; body very flexible, not flattened dorsoventrally, and more tapered than population described by Berger & Foissner (1987); cortical granules ellipsoidal, about 1 µm long, orange-yellow, arranged around cirri and dorsal bristles only (Fig. 79g); cytoplasm with many about 1 µm large, colourless fat granules; vivacious movement (Berger & Foissner 1989); body length:width ratio 7–8:1 (Hemberger 1985) and 6.7:1 on average (Berger & Foissner 1989). Contractile vacuole at end of first third of body, in specimen illustrated at 26% (Fig. 78a); transverse cirri about 15 µm, dorsal bristles 4 µm long (Hemberger 1985). According to Hemberger (1982, 1985) three dorsal kineties are present; unfortunately, no details about exact arrangement and length of kineties are provided, except for Fig. 78o showing an early divider. Accordingly, dorsal kinety 1 is roughly of body length, which is a distinct difference to our populations, where kinety 1 invariably terminates at or ahead of mid-body. Description of Namibian population (Fig. 80a–h, 80a–h, Table 26; from Foissner et al. 2002; see remarks): body size 80–160 × 10–15 µm in life, usually about 120 × 15 µm and thus slightly larger than the other populations. Body length:width ratio near 8:1 on average in protargol preparations (Table 26), that is, body slender with posterior half tapering and rear end narrowly rounded or bluntly pointed. Body often more or less distinctly sigmoidal and slightly twisted about main body axis; dorsoventrally only slightly flattened (about 1.5:1), acontractile, but very flexible. Narrowed posterior body portion usually stouter in preparations (Fig. 80g). On average 28 ellipsoidal macronuclear nodules scattered between buccal vertex and posterior fifth of body; individual nodules 5 × 3 µm in life, contain minute chromatin bodies. Usually three inconspicuous micronuclei at irregular position, 2.0 × 1.5 µm in size in life (Fig. 80a, g, 81b, e, h). Contractile vacuole slightly ahead of mid-body at left cell margin (Fig. 80a, 81a). Cortex thin and flexible, contains a loose lattice of fibres
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and conspicuous granules between cirral bases and around dorsal bristles. Individual granules brilliant yellowish, compact, ellipsoidal to ovoidal, 1.0–2.0 × 0.8–1.0 µm in size; rarely impregnate with the protargol method used (Fig. 80b, c, 81d–h). Cytoplasm colourless, contains some square crystals and small food vacuoles. Glides moderately rapidly on microscope slide and on and between soil particles, showing great flexibility. Adoral zone inconspicuous because occupying only 17% of body length on average (Table 26), indistinctly bipartite by small gap at left anterior corner of cell, producing three frontal (distal) and 10–11 ventral (proximal) membranelles of ordinary fine structure. Buccal field narrow and flat, buccal lip covers proximal third of adoral zone. Paroral and endoral difficult to distinguish, but show a characteristic, double curved pattern (Fig. 80g, 81b): a short, slightly curved anterior portion (paroral?) with 5 µm long cilia abuts on a longer and more distinctly curved posterior portion (endoral?); possibly, the short anterior portion overlaps the posterior to some extent. Pharyngeal fibres comparatively conspicuous. Cirral pattern and number of cirri of usual variability (Fig. 80a, d–h, Table 26). Cirri about 8 µm long in life, most composed of four cilia, buccal cirrus and some cirri of amphisiellid median cirral row consist of only two cilia. Frontal cirri not enlarged, arranged in transverse pseudorow, left cirrus (= cirrus I/1) frequently near gap between distal and proximal portion of adoral zone, right close to anterior frontoterminal cirrus. Buccal cirrus right of area where undulating membranes abut, inconspicuous because composed of two cilia only. Amphisiellid median cirral row composed of eight cirri on average, some of which composed of two cilia only; terminates at 23% of body length on average (Table 26); cirri slightly scattered, indicating that the row originates from several anlagen; however, ontogenetic data are needed for a correct interpretation and therefore classification (see remarks); anteriormost cirrus of row shifted slightly leftwards, indicating that it is cirrus III/2. Usually two frontoterminal cirri right of anterior portion of amphisiellid median cirral row (Fig. 80d, g). Two transverse cirri close to rear cell end, rarely they are lacking (Table 26). Right marginal row commences about at level of buccal vertex, terminates slightly ahead of rear cell end; left marginal row begins somewhat behind level of buccal vertex, terminates possibly slightly more posteriorly than right row. Dorsal bristles about 3 µm long in life, loosely arranged in invariably three kineties; kinety 1 composed of only 1–3 kinetids in anterior body half. Kinety 2 dis-
b Fig. 80a–h Hemisincirra inquieta (from Foissner et al. 2002. a–c, from life; d–h, protargol impregnation). a: Ventral view, 110 µm. b, c: Cortical granules are between cirri and around dorsal bristles. d: Dorsolateral view of anterior body portion showing relationship between frontoterminal cirri, marginal cirri, and amphisiellid median cirral row. e, f: A specimen likely having two caudal cirri. g, h: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 106 µm. Long arrow marks gap in adoral zone, short arrow denotes buccal cirrus. Frontal cirri connected by dotted line. ACR = amphisiellid median cirral row, AZM = distal end of adoral zone of membranelles, CC? = caudal cirri (see text), CV = contractile vacuole, FT = frontoterminal cirri, LMR = left marginal row, MA = macronuclear nodule, MI = micronucleus, RMR = right marginal row, TC = transverse cirri, 1–3 = dorsal kineties. Page 403.
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Fig. 81a–h Hemisincirra inquieta (from Foissner et al. 2002. a, e–h, from life; b–d, protargol impregnation). a: Dorsal view of a slightly squeezed specimen showing two rows of cortical granules associated with the dorsal kineties. b: The presumed paroral and endoral (posterior end marked by arrow head) form a characteristic, double curved pattern. Arrow marks left frontal cirrus (= cirrus I/1). c: The cortex contains a fibrillar lattice. d: Rarely impregnated granule clusters (arrowheads). e, f: Ventral and dorsal view of body portion behind mid-body. The brilliant cortical granules form clusters (arrowheads) between cirri and around dorsal bristles. g: Cortical granules (arrowheads) on dorsal side of rear body end. h: Optical section showing ellipsoidal shape of cortical granules (arrowheads). Explanation of original labelling: AM = ventral (proximal) adoral membranelles, BU = buccal cirrus, C = cirri, CV = contractile vacuole, DB = dorsal bristle, DK = dorsal kineties, F = fibres in buccal vertex, FC3 = right frontal cirrus, FV = anteriormost cirrus of amphisiellid median cirral row, LMR = left marginal row, MA = macronuclear nodules, TC = transverse cirri. Page 403.
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tinctly shortened anteriorly. 0–3, on average two cirri behind dorsal kineties; however, it cannot be excluded that these cirri are marginal cirri dislocated by the preparation procedures, which usually broaden the fragile posterior body portion (Fig. 80d, f –h, 81a, c, d, f). Cell division (Fig. 78c–o): Some sequences of ontogenesis are described for the type population by Hemberger (1982). The process commences with the proliferation of basal bodies at the posteriormost cirrus of the amphisiellid median cirral row. The primordium becomes relatively wide (Fig. 78c). Later, it is elongated and rather narrow (Fig. 78e). This oral primordium becomes larger and a narrow part of it extends anteriorly to the rear end of the parental undulating membranes (Fig. 78g). In the specimen shown in Fig. 78i, the parental undulating membranes have modified to a bipartite anlage (= anlage I). The anlage right of it is likely mainly formed by the narrow anlagen portion shown in Fig. 78g. The two rightmost primordia are modified cirri of the amphisiellid median cirral row. Right of the oral primordium is the undulating membrane anlage for the opisthe and two anlagen which either originated from the oral primordium or de novo. In the specimen shown in Fig. 78j, these two anlagen have elongated; likely the anterior portion of each anlage forms the rightmost anlagen of the proter. A middle to late divider with the fused macronucleus shows six frontal-ventral-transverse cirri anlagen, which correspond the ground pattern of the hypotrichs (Fig. 78k). Only the two rightmost anlagen form transverse cirri. Some of the primordial cirri are obviously absorbed in the final stages of cell division because the interphasic specimens have more or less invariably only 13 frontal-ventral-transverse cirri (Fig. 78a, m). Unfortunately, it is not possible to reconstruct the exact origin of the amphisiellid median cirral row, but very likely it is produced rather unspectacularly by cirri originating from the anlagen V and VI; perhaps anlage IV is also involved. The marginal rows divide in the ordinary manner, that is, each two anlagen originate in the parental rows to produce the new marginal cirri (Fig. 78k). The dorsal kinety pattern is formed as in many other hypotrichs, that is, within each parental kinety two anlagen originate; the anterior for the proter and the posterior for the opisthe (“Gonostomum pattern”, but without caudal cirri; see Berger 1999, p. 73). No caudal cirri are formed at the end of the dorsal kineties (Fig. 78o; Hemberger 1982). Kinety 1 is distinctly shortened in the populations described by Berger & Foissner (1987, 1989) and Foissner et al. (2002). Unfortunately, the length of this kinety in the type population is not described, but according to Fig. 78o it is of body length. For possible consequences resulting from this difference, see remarks. The division of the nuclear apparatus proceeds plesiomorphically manner, that is, the individual macronuclear nodules fuse to a single mass which later divides to the species-specific number (Fig. 78b, d, f, h, l, n, o). Occurrence and ecology: Hemisincirra inquieta is recorded from all main biogeographic regions, except Antarctica, and very likely confined to terrestrial habitats (Foissner 1998). Type locality not mentioned by Hemberger (1982, 1985); possibly
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it is mentioned on the type slides, which, however, I did not check. The population described by Berger & Foissner (1987) is from the upper soil layer (0–5 cm) of a spruce forest near the city of Ulm, Germany. The population studied by Berger & Foissner (1989; see nomenclature) was in a sample containing litter and the dark (volcanic) upper soil layer of a heath with dwarf shrubs (dominated by Betula nana, Empetrum nigrum, Vaccinium uliginosum, Arctostaphylos uliginosum) from Gooa Foss, Bardardålur, North Iceland. The Namibian population described by Foissner et al. (2002) is from a sample (humic litter/soil layer 0–3 cm; pH 6.7) in an Aloe dichotoma forest in the dwarf shrub savannah near the Gariganus Guest Farm, about 30 km northeast of the town of Keetmanshoop. It was frequent in Namibia, but saline habitats are less preferred (Foissner et al. 2002). Further records: alder stand at the Stubnerkogel, a mountain in the Gastein area, Austria (Berger & Foissner 1987, p. 211); soil of spruce forest in the Upper Austrian part of the Bohemian Forest treated with organically enriched magnesite fertilisers (Aescht & Foissner 1993, p. 328); soil from the Enns/Danube River floodplain, Upper Austria (Foissner 2004, p. 370); natural forest stands in Austria (Foissner et al. 2005, p. 628); soil from spruce forest in the city of Ulm, Germany (Lehle 1989, p. 141); soil contaminated by emissions from a textile factory in Nordhorn, Germany (Niebuhr 1989, p. 81); rice field near the city of Vercelli, Italy (Schwarz & Frenzel 2003, p. 247); dry mosses from near the city of Bratislava, Slovakia (Chrenková & Tirjaková 2000, p. 39); various sites (grassland, soil under leguminose tree, secondary pine forest) in Shimba Hills Nature Reserve, Kenya (Foissner 1999, p. 323); bark of Acacia tree and litter and soil layer of the tropical dry forest by the ranch house “La Casona” in the Santa Rosa National Park, Costa Rica (Foissner 1994a, p. 295; Foissner 1995, p. 39); various types of rain forest near the city of Manaus, Brazil (Foissner 1997, p. 322); Amazonian rain forest near the city of Iquitos, Peru (Foissner 1997, p. 322); soil and litter samples from the bush in Royal National Park south of Sydney, a pine forest and an Ecalyptus forest near Adelaide, and bark of trees from a pine forest and rain forest near Cairns, Australia (Blatterer & Foissner 1988, p. 8; Foissner 1997, p. 322). Hemisincirra inquieta feeds on heterotrophic flagellates (Polytomella) and bacteria (Foissner 1987, Foissner et al. 2002). Biomass of 106 specimens about 7 mg (Foissner 1987, p. 124).
Fig. 82a–p Hemisincirra namibiensis (from Foissner et al. 2002. a, from life; b–p, protargol impregnation). a: Ventral view of representative specimen, 80 µm. b, c: Infraciliature of ventral and dorsal side and nuclear apparatus of holotype specimen, 57 µm. Horizontal arrow marks gap in adoral zone, oblique arrow marks rear end of amphisiellid median cirral row. Pretransverse ventral cirri circled. d: Infraciliature of ventral side and nuclear apparatus of a specimen with six macronuclear nodules, 55 µm. Arrow marks buccal cirrus. e–p: Variability of nuclear apparatus (micronucleus black). AZM = distal end of adoral zone, FC = right frontal cirrus, FT = frontoterminal cirri, LMR = anterior end of left marginal row, MA = macronuclear nodule, MI = micronucleus, RMR = anterior end of right marginal row, TC = transverse cirri, 1, 2 = dorsal kineties. Page 418.
d
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Hemisincirra namibiensis Foissner, Agatha & Berger, 2002 (Fig. 82a–p, Table 26) 2002 Hemisincirra namibiensis nov. spec.1 – Foissner, Agatha & Berger, Denisia, 5: 857, Fig. 185a–p, Table 162 (Fig. 82a–p; original description; the holotype slide [accession number 2002/253] and a paratype slide [2002/254] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; Foissner et al. 2002, p. 39 and Aescht 2003, p. 392).
Nomenclature: The species-group name refers to the country (Namibia) where the type locality is (Foissner et al. 2002). Remarks: The cirral pattern of H. namibiensis is very similar to that of H. gellerti. However, in H. gellerti the amphisiellid median cirral row is distinctly longer than the adoral zone (vs. of about same length in present species) and the right marginal row is only slightly shortened anteriorly (vs. distinctly shortened). Further, Hemisincirra gellerti has cortical granules and more macronuclear nodules (8–9 on average vs. 4), adoral membranelles (13–17 vs. 12), and dorsal kineties (4 vs. 2) than H. namibiensis. Hemisincirra quadrinucleata has the same nuclear pattern, but is larger (110–130 µm vs. 60–80 µm) and has more adoral membranelles (15 or 16 vs. 12) and dorsal kineties (3 vs. 2); in addition, its undulating membranes are very differently arranged, namely as in Terricirra (see there). In life, Hemisincirra namibiensis is identified by the following combination of features (Foissner et al. 2002): 60–80 µm long; body elongate rectangular; four macronuclear nodules with usually two prominent micronuclei; right marginal row anteriorly strongly shortened, commencing at level of buccal vertex. Morphology: Body size in life 60–80 × 10–15 µm, on average 65 × 13 µm; body length:width ratio about 5:1 both in life and protargol preparations (Table 26). Body dorsoventrally flattened up to 2:1, flexible, but acontractile. Body outline elongate rectangular because of straight margins and evenly rounded ends. Usually four macronuclear nodules forming strand left of cell’s midline, nodules sometimes paired with one micronucleus in between (Fig. 82a, l); individual nodules usually ellipsoidal, rarely globular or irregular, contain many minute chromatin bodies. Usually two bright, globular micronuclei attached to macronuclear nodules at variable positions, prominent both in life and protargol preparations (Fig. 82a, c–p). Movement and presence or absence of contractile vacuole not observed. No cortical granules. Cytoplasm colourless, with some crystals and food vacuoles. Adoral zone occupies about 23% of body length, invariably composed of 12 membranelles of usual shape and structure in the 16 specimens investigated; distal four membranelles separated from remaining membranelles by an about one membranelle-wide gap at left anterior body corner. Buccal cavity small and flat. Exact structure and arrangement of undulating membranes not clearly recognisable; 1
Foissner et al. (2002) provided the following diagnosis: Size about 65 × 13 µm in vivo; elongate rectangular. On average 4 macronuclear nodules in series left of midline, 2 micronuclei, 12 adoral membranelles, 20 cirri in right and 24 cirri in left marginal row, 5 cirri in ventral cirral row, 2 frontoterminal cirri, 1 buccal cirrus, 4 transverse cirri, and 2 dorsal kineties.
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both membranes almost straight and usually one over the other, forming rather thick line. Pharyngeal fibres of ordinary length and structure after protargol impregnation, extend obliquely backwards (Fig. 82a, b, d). Cirral pattern rather constant, number of cirri of usual variability (Fig. 82a, b, d; Table 26). Most cirri about 10 µm long in life, fine because usually composed of two or four cilia only. Frontal cirri, frontoterminal cirri, and transverse cirri each usually composed of four cilia. Right frontal cirrus, as is usual, behind distal end of adoral zone. Buccal cirrus slightly behind anterior end of undulating membranes. Amphisiellid median cirral row usually composed of five cirri, terminates at 22% of body length, that is, at level of buccal vertex (23%). Invariably two frontoterminal cirri right of anterior end/half of amphisiellid median cirral row. One or two pretransverse ventral cirri, each composed of two cilia only, ahead of transverse cirri, which are near rear cell end. Both marginal rows commence about at level of buccal vertex and terminate distinctly ahead of rear cell end, that is, rows widely separated posteriorly. Dorsal bristles about 3 µm long, arranged in two kineties almost as long as body; bristles in kinety 2 rather widely spaced. No caudal cirri (Fig. 82c). Occurrence and ecology: Very likely confined to terrestrial habitats (Foissner et al. 2002, p. 53). Type locality of H. namibiensis is a sand dune at the escarpment of the southern Namib Desert between the towns of Aus and Helmeringshausen (26°05'S 16°35'E). This area is a semi-desert and savannah transition and the dune is overgrown by the dune grass Stipagrostis sp. and populated by a variety of insects, especially tenebrionids. The sample was composed of plant remnants and roots sieved off the sand around Stipagrostis shrubs at a depth of 0–10 cm (pH 5.4; Foissner et al. 2002, p. 18; site 17). Food not known.
Hemisincirra quadrinucleata Hemberger, 1985 (Fig. 83a, b, Table 26) 1982 Perisincirra quadrinucleata n. spec.1 – Hemberger, Dissertation2, p. 217, Abb. 38 (Fig. 83a, b; description of morphology). 1985 Hemisincirra quadrinucleata n. spec. – Hemberger, Arch. Protistenk., 130: 410, Abb. 16 (Fig. 83a, b; original description; no formal diagnosis provided; type slides are deposited in the Institut für landwirtschaftliche Zoologie und Bienenkunde, University of Bonn, Germany). 2001 Hemisincirra quadrinucleata Hemberger, 1985 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 30 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs).
Nomenclature: No derivation of the name is given in the original description. The species-group name quadrinucleata is a composite of the Latin numeral quattuor (quadr- in compositions; four), the thematic vowel ·i-, and nucleat·us, -a, -um (Latin adjective [m; f; n]; [nut]nucleus-like) and refers to the four macronuclear nodules. 1 2
This name is disclaimed for nomenclatural purposes (ICZN 1999, Article 8.3). See same footnote at Uroleptoides binucleatus.
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Remarks: According to the unusual arrangement of the undulating membranes, Hemisincirra quadrinucleata could belong to Terricirra (Berger & Foissner 1989, p. 36). However, since Hemberger (1982, 1985) give no information about the food vacuoles (spindle-shaped and with parallel-arranged bacteria in Terricirra) and the cortical granulation (distinct green to blue-green granules in Terricirra), Berger & Foissner (1989) did not transfer the present species to Terricirra (p. 447). Since no further data about H. quadrinucleata has become available I also retain the original classification. Hemisincirra quadrinucleata differs from Terricirra matsusakai (Fig. 93a–h), which also has four macronuclear nodules, by the number of dorsal kineties (3 vs. 4) and the adoral zone (with gap vs. without). Terricirra viridis has, like the present species, three dorsal kineties. However, it has eight macronuclear nodules (usually 4 in H. quadrinucleata [however, some specimens also have eight nodules!]) and lacks a gap in the adoral zone (Fig. 92a–j). In addition, it has conspicuous, dark-green cortical granules, a feature not described for H. quadrinucleata. Description of further populations of present species is recommended. The illustration of H. quadrinucleata in Hemberger (1982) shows the individual basal bodies of the cirri and adoral membranelles. Thus, this illustration, and not that in Hemberger (1985), which is identical in all other respects, is given in the present review. Morphology: Body size (in life?) 110–130 × 15–20 µm, that is, body length:width ratio 5–6:1. Body outline elongate elliptical, that is, margins almost parallel-sided and ends broadly rounded. Body very flexible. Usually four, rarely eight macronuclear nodules; position not indicated, but likely, as is usual, left of body midline behind adoral zone of membranelles. Two micronuclei in position shown in Fig. 83b, that is, near (not exactly in between) contact area of first and second, respectively, third and fourth macronuclear nodule. Contractile vacuole near
Fig. 83a, b Hemisincirra quadrinucleata (from Hemberger 1982. Protargol impregnation). a: Infracilature of ventral side, 123 µm. Body outline from live specimens. Arrowhead marks gap in adoral zone, short arrow denotes buccal cirrus, long arrow marks amphisiellid median cirral row whose anteriormost two cirri (circled) are very likely the frontoterminal cirri. Frontal cirri connected by dotted line. Cirri likely originating from anlage III connected by broken line. Note the conspicuous arrangement of the undulating membranes which is reminiscent of Terricirra. b: Nuclear apparatus (micronuclei black). AZM = distal end of adoral zone, CV = contractile vacuole, LMR = posterior end of left marginal row, RMR = right marginal row, TC = transverse cirri. Page 419.
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left cell margin behind buccal vertex, at 27% of body length in specimen illustrated (Fig. 83a). Presence/absence of cortical granules and details about cytoplasm (colour, food vacuoles, other inclusions) and movement not described. Adoral zone occupies 20–25% of body length, composed of 15–16 membranelles of usual fine structure; 4–5 frontal membranelles separated from ventral portion by distinct gap. Buccal field conspicuously narrow. Undulating membranes short, arranged as in Terricirra (see remarks), that is, in acute angle (Fig. 83a). Cirral pattern as shown in Fig. 83a. Three slightly enlarged frontal cirri arranged in almost transverse pseudorow with right cirrus behind distal end of adoral zone. Buccal cirrus right of anterior portion of anterior undulating membrane. Specimen illustrated (10 frontoventral cirri) with two cirri behind right frontal cirrus; in the more common specimens with only nine frontoventral cirri possibly only one cirrus behind. Amphisiellid median cirral row composed of four rather widely spaced cirri in specimen illustrated (Fig. 83a); possibly the anteriormost two are frontoterminal cirri (ontogenetic data needed for confirmation). Two, rarely three, about 10 µm long transverse cirri near rear cell end. Right marginal row commences about at level of buccal cirrus, terminates at rear cell end in specimen illustrated. Left marginal row commences left of buccal vertex, ends subterminal, that is, marginal rows separated posteriorly. Marginal cirri 8–10 µm long. Dorsal bristles 3–4 µm long, arranged in three kineties; exact arrangement and length of kineties not mentioned. Caudal cirri lacking. Occurrence and ecology: Very likely confined to terrestrial habitats (Foissner 1987, 1998). Type locality is the soil of a forest from the Puerto Maldonada area (about 69°12'W 12°36'S), Peru (Hemberger 1982, p. 2; 1985). Possibly restricted to the Neotropis because no further records published (Foissner 1998). We found it, for example, neither during a detailed survey of Namibian soil ciliates, nor in several Austrian forests (Foissner et al. 2002, 2005). Food not known. Biomass of 106 specimens about 26 mg (Foissner 1987, p. 124).
Hemisincirra octonucleata Hemberger, 1985 (Fig. 84a, b, Table 26) 1982 Perisincirra octonucleata n. spec.1 – Hemberger, Dissertation2, p. 219, Abb. 40 (Fig. 84a, b; description of morphology). 1985 Hemisincirra octonucleata n. spec. – Hemberger, Arch. Protistenk., 130: 411, Abb. 18 (Fig. 84a, b; original description; no formal diagnosis provided; type slides are deposited in the Institut für landwirtschaftliche Zoologie und Bienenkunde, University of Bonn, Germany). 2001 Hemisincirra octonucleata Hemberger, 1985 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 30 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs).
1 2
This name is disclaimed for nomenclatural purposes (ICZN 1999, Article 8.3). See same footnote at Uroleptoides binucleatus.
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Nomenclature: No derivation of the name is given in the original description. The species-group name octonucleata is a composite of the Latin numeral octo (eight) and nucleat·us, -a, -um (Latin adjective [m; f; n]; meaning [nut]nucleus-like) and refers to the eight macronuclear nodules. Remarks: According to Hemberger (1985) the assignment to Hemisincirra is not quite certain because the cirral pattern deviates somewhat from that of the other Hemisincirra species and ontogenetic data are lacking. Basically it would be possibly to transfer it to Lamtostyla because the amphisiellid median cirral row is longer than the adoral zone. However, since neither Lamtostyla nor Hemisincirra are very well defined (none of the two type species is characterised ontogenetically) I retain the original classification of the present species in Hemisincirra. Likely, only ontogenetic and/or molecular data on reliably determined populations will Fig. 84a, b Hemisincirra octonucleata (from Hemberger 1985. Protargol impregnation). a: Infraciliaimprove the classification significantly. ture of ventral side, 110 µm. Body outline from live Thus, detailed characterisation (includspecimen. Short arrow marks buccal cirrus, long aring, inter alia, live morphology, cell dirow denotes rear end of amphisiellid median cirral vision, relevant molecular data) of furrow. Supposed frontoterminal cirri circled. b: Nuther populations is recommended. clear apparatus. AZM = bipartite adoral zone of membranelles, CV = contractile vacuole, LMR = Terricirra viridis (Fig. 92a–j), which left marginal row, MA = macronuclear nodule, MI also has eight macronuclear nodules, = micronucleus, P = paroral, RMR = right marginal has, inter alia, green cortical granules row, TC = transverse cirri. Page 421. (presence/absence not known in Hemisincirra octonucleata), undulating membranes arranged at acute angle (vs. of ordinary arrangement), spindle-shaped food vacuoles with parallel-arranged bacteria (feature not known for present species), and a continuous adoral zone (vs. with distinct gap). Morphology: Body size (in life?) 95–120 × 18–25 µm; body length:width ratio about 5:1. Body outline parallel-sided with both ends broadly rounded. Consistency of body (flexible or rigid) not described; likely flexible as in congeners. Invariably eight macronuclear nodules arranged in two groups of four nodules each in speci-
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men illustrated; individual nodules usually ellipsoidal. One globular to ellipsoidal micronucleus attached to each group of macronuclear nodules (Fig. 84b); position of nuclear apparatus not described, but very likely, as is usual, slightly left of midline behind buccal vertex. Contractile vacuole near left cell margin behind adoral zone, in specimen illustrated at 32% of body length (Fig. 84a). Presence/absence of cortical granules and details about cytoplasm (colour, food vacuoles, other inclusions) and movement not described. Adoral zone occupies about 25% of body length, composed of 13–15 membranelles of usual fine structure, separated into frontal (= distal) portion (four membranelles) and ventral (= proximal) portion (9–11 membranelles) by distinct gap. Undulating membranes curved and optically slightly intersecting; paroral behind endoral (Fig. 84a). Cirral pattern as shown in Fig. 84a. In total 13–15 cirri in frontal area; detailed arrangement not described by Hemberger (1985). Frontal cirri slightly enlarged and arranged in transverse pseudorow behind frontal adoral membranelles. Buccal cirrus right ahead of anterior end of paroral. Two cirral pairs near distal end of adoral zone, right pair are likely the frontoterminal cirri. Amphisiellid median cirral row rather irregular, terminates at 39% of body length in specimen illustrated, that is, extends distinctly behind level of buccal vertex, which is at about 25% (Fig. 84a). Usually two transverse cirri near rear body end. Right marginal row commences at 19% of body length in specimen illustrated (Fig. 84a), terminates – like left row – at level of transverse cirri; marginal rows thus distinctly separated posteriorly. Left row begins left of buccal vertex. Dorsal bristles 3 µm long, arranged in three kineties; exact arrangement and length of kineties not described. Caudal cirri lacking. Occurrence and ecology: Very likely Hemisincirra octonucleata is confined to terrestrial habitats (Foissner 1987, 1998). Type locality is not given by Hemberger (1985), who mentioned only “suspension of mull rendzina soil”. However, this soil sample is not from Peru, the sole country mentioned in the material and methods section, but from Germany (see Hemberger 1982, p. 2). According to a personal communication by Norbert Wilbert (University of Bonn, Germany), the site is near the village of Mechernich (50°36'N 6°39'E), Germany. It is a beech forest on the “Kakushöhle Nordhang” and the outcrops are a middle devonian dolomite and a Pleistocene tufaceous limestone. No further records published. Thus, the neotropic distribution given by Foissner (1998, p. 204) and the lack of the present species in Germany according to Foissner (2000) are incorrect. Food not known. Biomass of 106 specimens about 24 mg (Foissner 1987, p. 124; 1998, p. 204).
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Hemisincirra gellerti (Foissner, 1982) Foissner in Berger, 2001 (Fig. 85a–h, Table 26) 1982 Perisincirra gellerti nov. spec.1 – Foissner, Arch. Protistenk., 126: 90, Abb. 22a–h, 68, Tabelle 20; not Abb. 22i (Fig. 85a–h; original description; the holotype slide [accession number 1981/91] is deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; Aescht 2003, p. 386). 1984 Hemisincirra gellerti (Foissner, 1982) nov. comb. – Foissner, Stapfia, 12: 119 (combination with Hemisincirra, see nomenclature). 1986 Hemisincirra gellerti (Foissner) – Foissner, Kataloge des OÖ. Landesmuseums, 5: 45, Abb. 7 (Fig. 85a; record from lichen sample). 2001 Hemisincirra gellerti (Foissner, 1982) comb. nov. – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 71 (combination with Hemisincirra, see nomenclature; nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs).
Nomenclature: Foissner (1982) dedicated this species to J. Gellért (Hungary) who made the first detailed studies about soil ciliates (e.g., Gellért 1956a, b; further references, see Berger 2006a). Foissner (1984) transferred this species to Hemisincirra before this genus was established by Hemberger (1985). I corrected this obviously invalid nomenclatural act and suggested citing the combining author as “Foissner in Berger, 2001” (Berger 2001). “Perisincira gellerti Foissner” in Tirjaková (1988, p. 500) is an incorrect subsequent spelling of Perisincirra. “Amphisiella gellerti Foissner ” in Foissner (1981, p. 18) is a nomen nudum because it does not, inter alia, satisfy the requirements of the ICZN (1964, Article 13). Remarks: Foissner (2000) described the subspecies Hemisincirra gellerti verrucosa, which differs, according to him, from H. gellerti gellerti only in the arrangement of the cortical granules. I consider both taxa as valid species because of the different cortical granulation and arrangement of the cirri in the frontoventral row, namely more or less distinctly paired/zigzagging in H. gellerti verrucosa (Fig. 134a–k), but continuous in H. gellerti gellerti (Fig. 85f). It is unlikely that a species shows such a high variation in two important features. Because of the zigzagging cirri in H. gellerti verrucosa I transfer it to the urostyloid taxon Anteholosticha in the present book. The original description of H. gellerti contains not only data from the type population (Großglockner area), but also from the Tullner Feld population (Foissner 1982, his Abb. 22i) which is, however, likely a synonym of Anteholosticha verrucosa. I have tried to eliminate the Tullner Feld data from the description below. Hemisincirra gellerti differs from H. wenzeli, inter alia, by the lack of caudal cirri and the longer amphisiellid median cirral row. For separation from other Hemisincirra species, see key. Morphology: Body size in life 65–110 × 12–20 µm; body length:width ratio of protargol-impregnated specimens 5.1:1 on average (Table 26), body outline thus 1
Foissner (1982) provided the following diagnosis: In vivo etwa 65–110 × 12–20 µm große, sehr zarte, hinten auffallend quer abgestutzte Perisincirra mit knapp halbkörperlanger Frontalreihe. Cirren der Frontalreihe besonders im oral Abschnitt häufig mehr oder minder deutlich zick-zackförmig angeordnet. Etwa 30 Längsreihen sehr kleiner, farbloser, subpelliculärer Granula. Makronucleus kettenförmig, aus 6–13 (meist 8) ellipsoiden Nodien aufgebaut. Adorale Membranellenzone zweigeteilt.
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Fig. 85a–h Hemisincirra gellerti (from Foissner 1982. a–c, e, from life; d, methyl green-pyronin staining; f–h, protargol impregnation). a: Ventral view of representative specimen, 108 µm. b: Left lateral view. c: Cortical granules on ventral side in mid-body. d: Ejected cortical granules. e: Dorsal view showing contractile vacuole during diastole. f–h: Infraciliature of ventral (f, h) and dorsal side (g) of different specimens, f = 80 µm. Arrow in (f) marks gap in adoral zone, broken lines connect cirri originating from anlagen I, II, and III. Arrow in (g) denotes short fifth kinety (note, however, that according to the morphometry, the present species has invariable only four kineties). Arrow in (h) marks, very likely, a pretransverse ventral cirrus. ACR = rear end of amphisiellid median cirral row, AZM = distal end of adoral zone of membranelles, CV = contractile vacuole, FC = right frontal cirrus, LMR = left marginal row, RMR = right marginal row, TC = transverse cirri, III/3 = cirrus behind right frontal cirrus, 1–4 = dorsal kineties. Page 424.
slender elliptical; anterior end narrowly rounded, rear end conspicuously truncated transversely (Fig. 85a, e). Body distinctly flattened dorsoventrally (Fig. 85b). Usually eight macronuclear nodules one after the other left of midline; individual nodules about 5 × 3 µm in life, chromatin bodies small. Several globular micronuclei (specimen illustrated with two; Fig. 85g) near macronuclear apparatus; individual micronuclei about 2 µm across. Contractile vacuole near left cell margin about in mid-body, during diastole with two long collecting canals (Fig. 85e). Pellicle and cytoplasm colourless. Cortical granules (termed “subpelliculäre” granules in original description) arranged in about 30 longitudinal rows, colourless; when methyl green-
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pyronin is added they are ejected and swell to ellipsoidal to bean-shaped, 1.5 µmsized bodies, forming a thin coat around cell (Fig. 85c, d). Cytoplasm with moderately many, very small, strongly refractive granules, some greasily shining, about 3 µm-sized inclusions, and few, colourless crystals about 3 µm long. Food vacuoles about 6 µm across. Movement moderately rapidly gliding, swims hastily to and fro under cover glass. Adoral zone of membranelles occupies 24% of body length and composed of 15 membranelles o average (Table 26). Bases of largest membranelles about 4 µm wide in life. Three distal membranelles distinctly set of from proximal portion of adoral zone (Fig. 85a, f). Buccal field flat, very small; exact arrangement of undulating membranes not described for type population (Fig. 87a, f). Pharyngeal fibres extend longitudinally backwards. All cirri of about same length, rather fine (Fig. 85a, f). Three frontal cirri behind distal adoral membranelles, not distinctly enlarged. Buccal cirrus composed of 2–3 basal bodies only, right of anterior portion of anterior undulating membrane (paroral?). One cirrus (= cirrus III/2) behind right frontal cirrus and thus left of anterior end of amphisiellid median cirral row, which is more or less continuous (Fig. 85f), that is, no distinct cirral pairs recognisable as, for example, in Anteholosticha verrucosa (Fig. 85c, i, k); row composed of 10 cirri, terminates at 43% of body length on average (Fig. 85f, Table 26). 2–5, on average three, transverse cirri (possibly one pretransverse ventral cirrus included) near rear cell end, project distinctly beyond body proper. Right marginal row commences at 14% of body length in specimen illustrated, terminates – like left row – slightly ahead of level of transverse cirri; marginal rows thus distinctly separated posteriorly; left marginal row commences near proximal end of adoral zone (Fig. 85f, g). Dorsal bristles about 3 µm long according to Fig. 85g. Number of kineties variable according to text of original description; according to Table 20 in Foissner (1982), however, invariable four kineties are present in type population. Specimen shown in Fig. 85g with five kineties: kineties 1 and 2 distinctly shortened anteriorly, kinety 5 composed of two basal body pairs only. Bristles rather widely spaced within kineties. Caudal cirri lacking. Occurrence and ecology: Very likely confined to terrestrial habitats and possibly of cosmopolitan distribution (Foissner 1987, 1998). Type locality is an alpine grassland near the Wallackhaus hotel (altitude 2310 m) at the Großglockner Hochalpenstrasse, a famous alpine road in Salzburg and Carinthia, Austria (Foissner 1982; site “SO8” in Foissner 1981). I suppose that Hemisincirra gellerti was sometimes confused with Anteholosticha verrucosa in life and even in protargol preparations (Fig. 134a–k) so that possibly not all records below refer to the present species: various habitats near the type locality in Salzburg, Austria (Foissner 1981, p. 18; as Amphisiella gellerti); common in alpine soils of the Gastein region (Salzburg, Austria), inter alia, in untreated, compacted, and fertilised sites (NPK, thomas-phosphate, crushed limestone) of a grazed alpine pasture with dwarf shrubs in the Schloßalm area (Foissner et al. 1982, p. 52; Berger et al. 1985, p. 106; 1986, p. 268; Foissner &
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Peer 1985, p. 42, with autecological data); spruce forest in northern Upper Austria (Petz et al. 1988, p. 82); soil of spruce forest in the Upper Austrian part of the Bohemian Forest treated with organically enriched magnesite fertilisers (Aescht & Foissner 1993, p. 328); spruce-fir-beech forest in Lower Austria (Foissner et al. 2005, p. 628); subalpine grassland in northern Styria, Austria (Foissner et al. 1990, p. 18); lichen from soil collected from an alpine pasture near the village of Obergurgl, Tyrol, Austria (Foissner 1986a, p. 44); soil from spruce forest in the city of Ulm and a beech forest near the village of Ermingen, Germany (Lehle 1989, p. 141); spruce stands in the Black Forest, Southern Germany (Funke 1986, p. 72; Lehle 1993, p. 17; 1994, p. 115; Lehle et al. 1992, p. 279); soil contaminated by emissions from a textile factory in Nordhorn, Germany (Niebuhr 1989, p. 81); coastal dunes of the North Sea island Norderney, Germany (Verhoeven 2002, p. 189); soil from the Sourhope Research Station near Kelso in Southern Scotland (Finlay et al. 2001, p. 362); agricultural soil from the village of Ostrov, Slovakia (Tirjaková 1988, p. 500; 1991, p. 41; Foissner 1994b, p. 158); soil and litter from Malé Karpaty Mountains, Slovakia (Tirjaková & Vdacny 2004, p. 28); soil under moss, city of Bratislava, Slovakia (Andelová & Tirjaková 2000, p. 35); riverside of Danube in Slovakia (Tirjaková 1992, p. 77; further record from Slovakia: Tirjaková 2005, p. 21); desert soils of the Grand Canyon (between Lees Ferry and Whitemore Wash) in northern Arizona, USA (Bamforth 2004, p. 417); soil from dry forest in the Santa Rosa National Park (Costa Rica) near the ranch house “La Casona” (Foissner 1995, p. 39; as Hemisincirra cf. gellerti, that is, identification uncertain); Amazonian rain forest in the vicinity of Iquitos, Peru (Foissner 1997, p. 322); soil samples from three sites (autochthonous pine forests, dry swamp) from Australia (Blatterer & Foissner 1988, p. 8); arid Australian soils (Robinson et al. 2002, p. 452); mire vegetation, that is, almost pure moss with very few soil particles from the Tafelkop area on Gough Island, South Atlantic ocean (Foissner 1996a, p. 284); Andreaea moss from Signy Island, Antarctic Region (Foissner 1996b, p. 97, 100). The record by Foissner et al. (1985, p. 108) obviously refers to Anteholosticha verrucosa (see remarks). Records from limnetic habitats (partially due to terrestrial influence, for example, due to flooding; some of them are possibly misidentifications): River Trattnach, Upper Austria (AOÖLR 1995, p. 100; determined by H. Blatterer); Guadarama River in the region of Villalba, Madrid, Spain (Fernandez-Leborans et al. 1990, p. 512); various brooks in the alpine zone of the High Tatra Mountains, Slovakia (Tirjaková 2001, p. 15; 2004, p. 7); in debris of the spring area below the Velky Javorník (the Mále Karpaty Mountains) near the city of Bratislava, Slovakia (Tirjaková 1997, p. 15; Tirjaková & Stloukal 2004, p. 16); Sumsoi region, Ukraine (Babko & Kovalchuk 1992, p. 124). Hemisincirra gellerti feeds on bacteria (Foissner 1982; 1987, p. 124). Biomass of 6 10 specimens about 10 mg (Foissner 1987, p. 124). Survived the application of 50 g l-1 NPK fertiliser (Tirjaková 1991, p. 41).
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Hemisincirra rariseta Foissner, Agatha & Berger, 2002 (Fig. 86a–g, Table 26) 2002 Hemisincirra rariseta nov. spec.1 – Foissner, Agatha & Berger, Denisia, 5: 865, Fig. 187a–g, Table 165 (Fig. 86a–g; original description; the holotype slide [accession number 2002/431] and two paratype slides [2002/429, 430] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; Foissner et al. 2002, p. 39 and Aescht 2003, p. 395).
Nomenclature: The species-group name rariseta is a composite of the Greek words rara (few) and saeta (bristle; cirrus in present case) and refers to the sparse frontoventral ciliature (Foissner et al. 2002). Remarks: This species very likely lacks transverse cirri and therefore the assignment to Hemisincirra is not quite certain. According to the lack of the transverse cirri and the habitus it could belong to Vermioxytricha, which, however, has only five frontal-ventral-transverse cirri anlagen (Foissner et al. 2002). The amphisiellid median cirral row of H. rariseta is composed of seven, very likely paired, cirri indicating that in total six (perhaps even seven) anlagen are present (Fig. 86f). In addition, Vermioxytricha arenicola (type species; Fig. 126a–m) forms a dorsomarginal kinety, a feature obviously lacking in the present species, which has two bipolar kineties. Thus, the original classification in Hemisincirra is (preliminarily) retained until more data (ontogenetic, molecular) enable a better classification. Hemisincirra rariseta differs from Vermioxytricha arenicola and V. muelleri, which have a similar body size and shape as well as number of macronuclear nodules and adoral membranelles, in the buccal cirrus (absent vs. present in Vermioxytricha) and dorsal kinety 2 (roughly of body length vs. composed of two bristles only). There are also several distinct differences in some morphometric features, like number of right and left marginal cirri, and the total number of cirri (frontal, buccal, frontoterminal, amphisiellid median row) in the frontal field (Table 26). Furthermore, Vermioxytricha arenicola has cortical granules, which are lacking in H. rariseta. Other vermiform species differ in the number of dorsal kineties: Hemisincirra buitkampi and Circinella vettersi have three and H. interrupta, H. vermicularis, and C. filiformis have only one. These species differ from the present species also in other features, but the number of dorsal kineties, which is usually very constant in hypotrichs, is – in the present case – the easiest characteristic recognisable also on careful live observation. However, all vermiform species are difficult to identify in life and therefore determinations should be checked in protargol preparations (Foissner et al. 2002). Morphology: Body size in life 120–200 × 10–20 µm, on average 170 × 14 µm; body length:width ratio invariably m8:1, on average 10:1 (Table 26). Body vermi1
Foissner et al. (2002) provided the following diagnosis: Size about 170 × 14 µm in vivo; vermiform. Usually 16 macronuclear nodules in series left of midline. On average 26 right and 25 left marginal cirri. 2 frontoterminal cirri, and 7 frontoventral cirri in a row extending near level of buccal vertex. Invariably 2 bipolar dorsal kineties. Adoral zone bipartited into 3 frontal and 13 ventral membranelles, occupies about 16% of body length.
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Fig. 86a–c Hemisincirra rariseta (from Foissner et al. 2002. a, from life; b, c, protargol impregnation). a: Ventral view of a twisted specimen, 165 µm. Short arrow marks gap in adoral zone, long arrow denotes three spread cirri at rear cell end, likely caudal cirri. b, c: Infraciliature of ventral and dorsal side and nuclear apparatus of holotype specimen, 165 µm. MA = rearmost macronuclear nodule, MI = micronucleus, 1, 2 = dorsal kineties. Page 428.
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Fig. 86d–g Hemisincirra rariseta (from Foissner et al. 2002. Protargol impregnation). d, f, g: Details of infraciliature in ventral (d), dorsolateral (f), and ventrolateral (g) view. Short arrow in (d) marks frontal (distal) adoral membranelles. Long arrow in (d) and arrow in (f) denote rear end of amphisiellid median cirral row. Arrow in (g) marks enlarged granule at anterior end of paroral, possibly a vestigial buccal cirrus. Dotted lines in (d, f) connect frontal cirri, broken lines connect cirri likely originating from same anlage during cell division (lines denoted by question mark have to be checked by ontogenetic data). e: Posterior body portion of specimen shown in (d). The marginal cirri are composed of four cilia only. Arrow marks three spread cirri at rear cell end (possibly caudal cirri). AZM = bipartite adoral zone of membranelles, E = endoral, FC = left frontal cirrus in gap of adoral zone, FT = frontoterminal cirri, LMR = left marginal row, P = paroral, PF = pharyngeal fibres, RMR = right marginal row, 2 = right dorsal kinety. Page 428.
form with anterior portion slightly and posterior portion conspicuously narrowed; however, a distinct tail is not present; frequently twisted about main body axis and serpentine with posterior portion usually straight or slightly curved; flattened only in oral area, acontractile (Fig. 86a, b). Nuclear apparatus in central quarters of cell, macronuclear nodules in series left of midline, rarely in two indistinct overlapping strands; individual nodules globular to elongate ellipsoidal, on average ellipsoidal, chromatin bodies minute. Usually one broadly ellipsoidal micronucleus each near anterior and posterior quarter of macronuclear strand. Contractile vacuole slightly ahead of mid-body. Cortex thin and very flexible; specific granules lacking. Cytoplasm colourless, contains many granules and food vacuoles with bacteria and fungal spores. Moves moderately rapidly and serpentinely on microscope slide and between soil particles, showing great flexibility.
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Adoral zone inconspicuous because occupying only 16% of body length on average, bipartite into three frontal (distal) and 13 ventral (proximal) membranelles of ordinary fine structure by a distinct gap at left anterior body corner (Fig. 86a, b, d, f, g). Anteriormost proximal membranelles composed of three basal body rows only (Fig. 86d, g). Buccal field narrow and flat, buccal lip covers posteriormost adoral membranelles. Paroral and endoral close together and staggered, paroral likely composed of dikinetids. Pharyngeal fibres conspicuous. Cirral pattern and number of cirri of usual variability, except for the strongly varying number of cirri in the amphisiellid median cirral row (Fig. 86a, b, d–f; Table 26). Cirri about 12 µm long and fine, consist of two rows with only two (posterior third of marginal rows, frontoterminal cirri, rear cirri of amphisiellid median cirral row) or three (anterior and middle third of marginal rows and amphisiellid median cirral row) cilia each. Frontal cirri arranged in oblique pseudorow, left cirrus invariably in gap of adoral zone, right cirrus behind distal end of adoral zone. Buccal cirrus very likely lacking or very close to anterior end of paroral, where a slightly enlarged granule is recognisable in some specimens (Fig. 86g, arrow). One cirrus (= cirrus III/2) behind right frontal cirrus. Amphisiellid median cirral row composed of an average of seven cirri forming indistinct pairs and pseudopairs, terminates at 14% of body length, that is, slightly ahead of level of buccal vertex; ontogenetic data are needed to know the number of anlagen involved in row formation. Usually two frontoterminal cirri right of anterior end of amphisiellid median cirral row. Postperistomial cirrus and transverse cirri lacking (see remarks). Right marginal row commences at 11% of body length on average (Table 26), that is, slightly ahead of level of buccal vertex; by contrast, left row commences slightly behind level of buccal vertex. Both marginal rows extend to posterior body end bearing 2–3 spread cirri; however, it cannot be excluded that these are caudal cirri. Dorsal bristles about 5 µm long in life, arranged in two kineties extending to rear body end; kinety 1 slightly shortened anteriorly. Bristles of kinety 2 more loosely spaced than those of kinety 1, except in anterior portion (Fig. 86c; Table 26). Likely 2–3 spread caudal cirri present (Fig. 86a, e; ontogenetic data needed for confirmation). Occurrence and ecology: Very likely Hemisincirra rariseta is confined to terrestrial habitats (Foissner et al. 2002, p. 53). Type locality is an area near the Gariganus Guest Farm (26°30'S 18°25'E), Namibia (Foissner et al. 2002, p. 16, 865; site 5). This site is an Aloe dichotoma forest in the dwarf shrub savannah. The sample was composed of litter and humus layer from under some Aloe trees (pH 6.7). Foissner et al. (2002) found it only at the type locality where it was, however, rather abundant. No further records published. Hemisincirra rariseta is well adapted to the sandy habitat by its vermiform body. Food not known.
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Hemisincirra interrupta (Foissner, 1982) Foissner in Berger, 2001 (Fig. 87a–f, Table 26) 1982 Perisincirra interrupta nov. spec.1 – Foissner, Arch. Protistenk., 126: 99, Abb. 26a–e, Tabelle 23 (Fig. 87a–f; original description; type slides possibly deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria, although not listed in catalogue by Aescht 2003, p. 388). 1984 Hemisincirra interrupta (Foissner, 1982) nov. comb. – Foissner, Stapfia, 12: 119 (combination with Hemisincirra; see nomenclature). 2001 Hemisincirra interrupta (Foissner, 1982) comb. nov. – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 72 (combination with Hemisincirra, see nomenclature; nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs).
Nomenclature: No derivation of the name is given in the original description. The species-group name interrupt·us, -a, -um (Latin adjective [m; f; n]; interrupted) obviously refers to the distinct gap (interruption) in the adoral zone of membranelles. Foissner (1984) transferred P. interrupta to Hemisincirra before Hemisincirra Hemberger, 1985 was available. I corrected this obviously invalid nomenclatural act in the catalogue of ciliates names (Berger 2001) and suggested to cite the name as shown in the heading. “Paruroleptus (?) interrupta Foissner” in Foissner (1981, p. 19, 21) is a nomen nudum because published without description. “Perisincira interrupta Foissner” in Tirjaková (1988, p. 500) is an incorrect subsequent spelling of Perisincirra. Remarks: Hemisincirra interrupta closely resembles Circinella filiformis (Foissner, 1982) Foissner, 1994 which, however, has a shorter adoral zone and a longer amphisiellid median cirral row. Because of this difference Foissner (1994) did not transfer the present species to Circinella. I preliminarily retain the classification in Hemisincirra because I have, at the present state of knowledge (ontogenetic and molecular data lacking), no better idea. Morphology: Body size in life 80–130 × 8–15 µm; body length:width ratio 11:1 in protargol preparations (Table 26). Body outline very slender, spindle-shaped to wedge-shaped, often slightly S-shaped and slightly twisted about main body axis; anterior and posterior end narrowly rounded (Fig. 87a, b). Anterior body portion distinctly flattened dorsoventrally, that is, remaining portion not flattened. On average 30 macronuclear nodules arranged chain-like in middle body portion slightly left of cell midline (Fig. 87a, c); individual nodules globular or ellipsoidal with many, tiny chromatin bodies. Micronuclei not known. Contractile vacuole near left cell margin slightly ahead of mid-body, at 42% of body length in specimen illustrated (Fig. 87b). Pellicle colourless, very soft and flexible. No specific cortical granules. Cytoplasm colourless, in rear body portion with some small, yellowish crystals. Food vacuoles 1
Foissner (1982) provided the following diagnosis: In vivo etwa 80–130 × 8–15 µm große, sehr schlank spindelförmige Perisincirra, deren Frontalreihe ungefähr so lang ist wie die adorale Membranellenzone. Durchschnittlich 14 adorale Membranellen, von denen die vorderen 3–4 durch eine sehr breite Lücke von den hinteren getrennt sind. Macronucleus kettenförmig, aus durchschnittlich 30 kugelförmigen bis ellipsoiden Nodien aufgebaut. 1 Dorsalkinete leicht links der Medianen.
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Fig. 87a–f Hemisincirra interrupta (from Foissner 1982. a, b, from life; c–f, protargol impregnation). a: Ventral view of representative specimen, 114 µm. b: Dorsal view showing contractile vacuole, 98 µm. c, d: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 105 µm. e, f: Infraciliature of anterior and posterior body portion. Note large gap in adoral zone (double-sided arrow in e). Long arrow in (e) denotes rear end of amphisiellid median cirral row, short arrows mark two cirri (frontoterminal cirri? anterior end of right marginal row?) right of anterior end of amphisiellid median cirral row. Cirri originating from frontal-ventral cirri anlage I and III connected by broken lines (has to be checked by ontogenetic data). Frontal cirri connected by dotted line. Arrows in (f) mark three cirri at rear cell end whose correct designation (transverse? caudal? marginal?) is not yet known because ontogenetic data are lacking. AZM = distal end of adoral zone of membranelles, CV = contractile vacuole, LMR = left marginal row, P = paroral, PF = pharyngeal fibres, RMR = right marginal row, I, III = frontal-ventral cirri anlagen, 1 = dorsal kinety. Page 432.
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about 5 µm across and with indefinable content. Movement moderately rapid, creeps hastily to and fro. Adoral zone occupies 15% of body length and composed of 14 membranelles on average; 3–4 distalmost membranelles separated from remaining proximal portion by wide gap (Fig. 87a, c, e). Paroral two-rowed and slightly curved, arranged right of rear portion of adoral zone; endoral not seen. Buccal field very small, almost flat. Cirral pattern and number of cirri of usual variability (Fig. 87c, e, f, Table 26). All cirri very fine and almost all about 10 µm long. Three slightly enlarged frontal cirri with middle cirrus about at level of right cirrus so that the pseudorow is slightly concave. Buccal cirrus not seen, indicating that it is lacking. Amphisiellid median cirral row commences behind middle and right frontal cirrus; cirri slightly irregularly arranged (indicating that it originates from more than one anlage); terminates at 12% of body length on average, that is, slightly shorter than adoral zone (Table 26). Whether frontoterminal cirri are present or not is not known (Fig. 87e). Postperistomial cirri lacking. Rear cell end with several (three according to Fig. 87f) about 15 µm long transverse or caudal cirri, which are distinctly spread (see dorsal ciliature). Beginning of right marginal row not clearly recognisable in specimens illustrated because it is not known whether the cirri right of the anterior portion of the amphisiellid median cirral row are marginal or frontoterminal cirri (Fig. 87c, e). Marginal rows separated posteriorly. Left row commences left of level of buccal vertex; distance between individual cirri in posterior portion about three times as high as in anterior portion. Cirri in right row roughly equally spaced. Length of dorsal bristles not mentioned, according to Fig. 87d about 2 µm. Bristles arranged in one kinety of body length slightly right of midline (Fig. 87d). Whether the cirri at the rear cell end are transverse or caudal cirri cannot be decided without ontogenetic data (Fig. 87c, f). Perhaps the middle cirrus is a caudal cirrus produced by the single dorsal kinety and the two lateral cirri are the rear end of the marginal rows. Occurrence and ecology: Very likely confined to terrestrial habitats (Foissner 1987) and reliably recorded from Holarctis, Palaeotropis, and Australis (Foissner 1998, p. 204). Type locality of H. interrupta is the area near the Wallackhaus hotel (2290 m above sea-level), at the Großglockner Hochalpenstrasse, an alpine road in the Central Alps in Austria; Hemisincirra interrupta occurred in a soil sample from a eutrophic area (Foissner 1982; see also Krainer 1999, p. 668). For further records along this road, see Foissner (1981). Further records: alpine soils in the Gastein area, Salzburg, Austria (Foissner & Peer 1985, p. 42; including autecological data); Pruno-Fraxinetum from a floodplain (calcaric fluvisol; typical mull) in eastern Austria (Foissner et al. 2005, p. 628; site “Müllerboden”); agricultural soils in Slovakia (Tirjaková 1988, p. 500); soil samples from the riverside zone of the Danube river (km 1858) in Slovakia (Tirjaková 1992, p. 77); soil under moss in the city of Bratislava, Slovakia (Andelová & Tirjaková 2000, p. 35; further record from Slovakia: Tirjaková 2005, p. 21); spruce forest and beech forest near the city of Ulm, Germany (Lehle 1989, p. 141; see also Foissner 2000, p. 258); spruce stands in the Black For-
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est, Southern Germany (Lehle 1993, p. 17; 1994, p. 115; Lehle et al. 1992, p. 279); upper soil layer (0–5 cm) with litter of a secondary pine forest near the city of Adelaide, Australia (Blatterer & Foissner 1988, p. 8). Food not known. Biomass of 106 specimens about 8 mg (Foissner 1987, p. 124).
Hemisincirra vermicularis Hemberger, 1985 (Fig. 88a–c, Table 26) 1982 Perisincirra vermiculare n. spec.1 – Hemberger, Dissertation2, p. 217, Abb. 39 (Fig. 88a, b; description of morphology). 1985 Hemisincirra vermiculare n. spec. – Hemberger, Arch. Protistenk., 130: 411, Abb. 17 (Fig. 88a, b; original description; no formal diagnosis provided; type slides are deposited in the Institut für landwirtschaftliche Zoologie und Bienenkunde, University of Bonn, Germany). 2001 Hemisincirra vermiculare Hemberger, 1985 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 30 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs).
Nomenclature: The species-group name vermicular·is, -is, -e (Latin adjective [m; f; n]; similar to a small worm; Hentschel & Wagner 1996, p. 606) alludes to the wormlike body (Hemberger 1985). Since Hemisincirra is feminine, the ending of the species-group name (adjective) has to be corrected from vermiculare (neuter) to vermicularis. Remarks: Hemisincirra vermicularis has, like several other Hemisincirra species, a slender, worm-shaped body making the identification rather difficult because the cirral pattern is very difficult to recognise in life in such slender specimens. However, it has a unique combination of characters (e.g., four contractile vacuoles, one dorsal kinety) not known from an other species so that the validity is beyond reasonable doubt. Hemberger (1982, 1985) did not study the cell division of H. vermicularis and therefore he could not decide whether it has transverse or caudal cirri. Consequently, the generic assignment is uncertain, that is, further data (ontogenetic, molecular) are needed for a well founded classification. In addition, a redescription is recommended because the description does not contain information about the presence/absence of cortical granules. For problems with the type locality, see occurrence and ecology section. Morphology: Body size (in life?) about 200 × 12 µm resulting in a body length:width ratio of 16:1, that is, body vermiform. About 10 macronuclear nodules arranged one after the other, likely slightly left of body midline. Individual nodules ellipsoidal (Fig. 88b). One micronucleus each near anterior and posterior end of macronuclear apparatus; whether the position shown in Fig. 88b is constant or not is not described. Four contractile vacuoles at left body margin, during diastole with collecting canals; in specimen illustrated the individual vacuoles are at about 13%, 21%, 51%, and 76% of body length (Fig. 88a). Presence/absence of cortical granules 1 2
This name is disclaimed for nomenclatural purposes (ICZN 1999, Article 8.3). See same footnote at Uroleptoides binucleatus.
436
SYSTEMATIC SECTION and details about cytoplasm (colour, food vacuoles, other inclusions) and movement not described. Adoral zone occupies only about 10% of body length, bipartite, that is, ventral (= proximal) portion (about eight membranelles) separated from frontal (= distal), transversely arranged portion (four membranelles) by distinct gap. Buccal field likely very small and difficult to observe. Undulating membranes small, that is, composed of only few cilia (Fig. 88a, c). Cirral pattern as shown in Fig. 88a, c, that is, as in the other vermiform species included in Hemisincirra. Three frontal cirri in almost transverse pseudorow; according to Fig. 88c middle and right frontal cirrus composed of four cilia, left cirrus – like all other cirri – composed of two cilia only. Buccal cirrus likely lacking because according to Hemberger (1985) the frontal ciliature is composed of 10 cirri (Fig. 88a, c), namely three frontal cirri, two cirri behind right frontal cirrus, and five cirri in the continuous amphisiellid median cirral row which terminates about at level of buccal vertex. Postperistomial cirri lacking; whether the (three?) cirri at the rear cell end are transverse or caudal cirri could not be decided. Right marginal row commences about at same level as amphisiellid median cirral row, terminates distinctly ahead of cell end (at about 93% of body length in specimen illustrated in Fig. 88a). Left marginal row commences close to buccal vertex, terminates about at same level as right row, that is, marginal rows widely separated posteriorly.
Fig. 88a–c Hemisincirra vermicularis (from Hemberger 1985. Protargol impregnation). Infraciliature of ventral side and nuclear apparatus, 200 µm. Body outline from live specimen. Short arrows in (a) mark contractile vacuoles, long arrow denotes (transverse? caudal?) cirri at rear cell end. Asterisks mark rear end of marginal rows. Arrow in (b) marks micronucleus. Arrow in (c; detail of a) marks rear end of amphisiellid median cirral row. Dotted line connects frontal cirri, broken line connects cirri likely originating from anlage III. Tiny undulating membranes circled. AZM = bipartite adoral zone with four frontal (= distal) and eight ventral (= proximal) membranelles, CV = anteriormost contractile vacuole, LMR = anterior end of left marginal row, MA = macronuclear nodule, RMR = anterior end of right marginal row. Page 435.
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Dorsal bristles about 4 µm long; arranged in one kinety (details neither described nor illustrated). Correct designation (transverse or caudal) of cirri at rear cell end not possible without ontogenetic data. Occurrence and ecology: Likely confined to terrestrial habitats (Foissner 1987, 1998). Type locality not mentioned in original description and in Hemberger (1982); according to a personal communication, he discovered it in the Puerto Maldonada region in Peru, as already suggested by Foissner (1998, p. 204). Food not known. Biomass of 106 specimens about 24 mg (Foissner 1987, p. 124).
Incertae sedis in Hemisincirra Hemisincirra wenzeli Foissner, 1987 (Fig. 89a–h, Table 29) 1987 Hemisincirra wenzeli nov. spec.1 – Foissner, Zool. Beitr. (N.F.), 31: 216, Abb. 10a–h, Tabelle 2 (Fig. 89a–h; original description; the holotype side [accession number 1988/149] and a paratype slide [1988/150] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; Aescht 2003, p. 400). 2001 Hemisincirra wenzeli Foissner, 1987 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 31 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs).
Nomenclature: Foissner (1987a) dedicated this species to Fritz Wenzel (Hamburg, Germany), who made basic studies on moss ciliates (e.g., Wenzel 1953). Remarks: The generic assignment of this species is uncertain because H. wenzeli has caudal cirri, whereas such cirri are lacking in the type species H. buitkampi and likely in many (all?) other Hemisincirra species. According to the Urosoma-like arrangement of the cirri of the amphisiellid median cirral row it could belong to Hemiurosoma Foissner, Agatha & Berger, 2002, whose members have caudal and transverse cirri (p. 614). However, all species included in Hemiurosoma have a dorsomarginal kinety. Such a row is obviously lacking in H. wenzeli (Fig. 89g), strongly indicating that a classification in Hemiurosoma is not correct. Caudiamphisiella also has caudal cirri, but a rather long amphisiellid median cirral row. In addition, the type species C. antarctica is marine and has more than three bipolar dorsal kineties so that the assignment of the present species to this genus also seems inappropriate. Because of these uncertainties I retain the present species in Hemisincirra; ontogenetic and/or relevant molecular data are needed for a more proper classification. The most similar species in Hemisincirra are H. gellerti and H. inquieta, which, however, lack caudal cirri (details see these species). 1 Foissner (1987a) provided the following diagnosis: In vivo etwa 70–110 × 12–18 µm große Hemisincirra mit ellipsoiden farblosen subpelliculären Granula. Durchschnittlich 12 adorale Membranellen und 29 ellipsoide Makronucleus-Teile, die im mittleren Abschnitt des Ciliaten regellos angeordnet sind. 2 Transversal- und 3 Caudalcirren. 3 Dorsalkineten.
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Fig. 89a–h Hemisincirra wenzeli (from Foissner 1987a. a–e, from life; f–h, protargol impregnation). a: Ventral view of a representative specimen, 110 µm. b: Contracted specimen. c, d: Cortical granules in lateral view and top view. The granules are 1.2 × 0.8 µm, colourless, and arranged along cirral rows and dorsal kineties. e: Dorsal view showing contractile vacuole. f, g: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 69 µm. h: Infraciliature of anterior body portion, 33 µm. Arrow marks gap in adoral zone. Frontal cirri connected by dotted line; broken lines connect cirri very likely originating from same anlage (only shown for anlagen I–III; has to be checked by ontogenetic data). Cirri of amphisiellid median cirral row (circled) arranged in Urosoma-pattern. AZM = distal end of adoral zone of membranelles, CC = caudal cirri, CG = cortical granules, FC = right frontal cirrus, LMR = left marginal row, PF = pharyngeal fibres, RMR = right marginal row, TC = transverse cirri, 1–3 = dorsal kineties. Page 437.
Morphology: Body size in life 70–110 × 12–18 µm; body length:width ratio 6.5:1 on average in protargol preparations (Table 26). Body outline oblong, more or less sigmoidal, slightly to distinctly narrowed posteriorly (Fig. 89a, b). Body slightly contractile, contracted specimens S-shaped (Fig. 89e). Body not flattened dorsoventrally, except for oral region which is slightly flattened. On average 29 macronuclear nodules irregularly arranged in middle portion of cell; most nodules of specimen illustrated, however, as is usual, left of midline (Fig. 89g). Individual nodules ellipsoidal, chromatin bodies of ordinary size. 2–4 micronuclei, which do not impregnate with protargol sometimes. Contractile vacuole near left cell margin slightly ahead of mid-body (Fig. 89e). Pellicle fine and flexible. Cortical granules (designated as sub-
Hemisincirra
439
pellicular granules in original description) along cirral rows and dorsal kineties; individual granules colourless, about 1.2 × 0.8 µm, do not impregnate with protargol. Cytoplasm colourless, stuffed more or less distinctly (depending on nutritional state) with food vacuoles about 8 µm across and 1–2 µm-sized, colourless crystals mainly in posterior body portion. Locomotion moderately rapid, often moves rapidly to and fro while slightly contracting and bending sigmoidally (Fig. 89e). Adoral zone occupies about 15–20% of body length, composed of 12 membranelles on average (Table 26); distalmost three membranelles set off from proximal portion of adoral zone by small gap (Fig. 89a, f, h). Buccal field narrow, but rather deep; right margin bordered by the two narrowly spaced undulating membranes. Cirral pattern and number of cirri of usual variability (Fig. 89a, f, h, Table 26). Frontal cirri slightly enlarged, arranged in oblique pseudorow with middle cirrus behind distal end of adoral zone. Buccal cirrus right of anterior end of paroral. Amphisiellid median cirral row usually composed of four cirri arranged in Urosomapattern, that is, cirrus III/2 slightly left and ahead of remaining cirri of row, which is close to anterior portion of right marginal row. Postperistomial cirri lacking. Two transverse cirri close to rear cell end, about 15 µm long. Right marginal row commences about at level of buccal cirrus, terminates – like left row – slightly subterminal so that marginal rows distinctly separated posteriorly. Left row commences close to buccal vertex. Marginal cirri about 10 µm long. Dorsal bristles about 3 µm long, arranged in three kineties; kinety 1 composed of two basal body pairs only, that is, row distinctly shortened anteriorly and posteriorly (this kinety is certainly not a dorsomarginal kinety because it is on the left margin and, very likely, produces a caudal cirrus); kineties 2 and 3 roughly of body length. Three caudal cirri, indicating that at the end of each kinety one cirrus is produced; however, ontogenetic data are needed for correct interpretation. Caudal cirri about 15 µm long (Fig. 89a, g). Occurrence and ecology: Very likely confined to terrestrial habitats (Foissner 1987, 1998). Type locality of Hemisincirra wenzeli is the Danish island of Bornholm in the Baltic Sea where it occurred in mosses collected by F. Wenzel (Foissner 1987a); the abundance in the raw cultures was moderately high. Further records: beech forest (Luzulo-Fagenion) near the village of Dürnstein, Lower Austria (Foissner et al. 2005, p. 628); plant debris from the Coral Pink Sand Dunes about 25 km west of Rockville, Zion National Park, Utah, USA (Foissner 1994, p. 164); litter and roots under moss in an autochthonous pine forest (pH 7.4) near Adelaide, Australia (Blatterer & Foissner 1988, p. 8); upper (0–2 cm) leaf litter and soil layer from a rain forest near Cairns, Australia (Foissner 1997, p. 322). Hemisincirra wenzeli feeds on heterotrophic flagellates and very likely also on bacteria (Foissner 1987a). Biomass of 106 specimens about 14 mg (Foissner 1998, p. 204).
440
SYSTEMATIC SECTION
Table 28 Morphometric data on Mucotrichidium hospes (from Foissner et al. 1990a) Characteristics a Body, length Body, width Adoral zone of membranelles, length Adoral membranelles, number Macronuclear nodule, length Macronuclear nodule, width Macronuclear nodules, distance in between Macronuclear nodules, number Micronucleus, largest diameter Micronuclei, number Frontal cirri, number Buccal cirri, number Cirri behind right frontal cirrus, number Amphisiellid median cirral row, number of cirri Left marginal cirri, number Right marginal cirri, number Dorsal kineties, number Caudal cirri, number
mean
M
SD
SE
CV
Min
Max
n
67.4 29.0 29.5 49.3 12.4 7.1 2.8 2.0 3.9 1.3 3.0 5.5 7.2 34.7 39.3 37.9 3.0 2.7
63.0 28.0 29.0 51.0 13.0 7.0 3.0 2.0 4.0 1.0 3.0 5.0 7.0 34.0 39.0 37.0 3.0 3.0
9.8 5.1 4.3 3.1 2.1 1.0 1.4 0.0 0.7 0.5 0.0 0.9 0.9 2.0 3.9 4.4 0.0 0.5
– – – – – – – – – – – – – – – – – –
14.6 17.7 14.7 6.3 16.9 13.7 48.9 0.0 17.9 36.0 0.0 17.1 12.5 5.7 10.0 11.6 0.0 17.5
54.0 22.0 23.0 44.0 8.0 6.0 1.0 2.0 3.0 1.0 3.0 4.0 6.0 31.0 34.0 30.0 3.0 2.0
83.0 38.0 36.0 54.0 17.0 10.0 7.0 2.0 6.0 2.0 3.0 7.0 9.0 37.0 48.0 49.0 3.0 3.0
17 17 17 18 17 17 17 17 17 27 16 17 17 17 21 18 21 17
a All measurements in µm. Data based on mounted, protargol-impregnated (Foissner’s protocol), and randomly selected specimens. CV = coefficient of variation in %, M = median, Max = maximum, mean = arithmetic mean, Min = minimum, n = number of individuals investigated, SD = standard deviation, SE = standard error of arithmetic mean.
Mucotrichidium Foissner, Oleksiv & Müller, 1990 1990 Mucotrichidium nov. gen.1 – Foissner, Oleksiv & Müller, Arch. Protistenk., 138: 196 (original description). Type species (by original designation): Uroleptus hospes Ehrenberg, 1831. 1999 Mucotrichidium Foissner, 1990 – Shi, Song & Shi, Progress in Protozoology, p. 104 (revision of hypotrichs; incorrect authorship). 1999 Mucotrichidium Foissner, 1990 – Shi, Acta Zootax. sin., 24: 255 (revision of hypotrichs; incorrect authorship). 2001 Mucotrichidium Foissner, Oleksiv & Müller 1990 – Aescht, Denisia, 1: 103 (catalogue of generic names of ciliates). 2001 Mucotrichidium Foissner, Oleksiv and Müller, 1990 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 48 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Mucotrichidium Foisner, Oleksiv and Müller, 1990 – Lynn & Small, Phylum Ciliophora, p. 448 (guide to ciliate genera; incorrect spelling of Foissner).
Nomenclature: Mucotrichidium is a composite of the Latin nouns mucor (slime) and trichos (hair), and the diminutive suffix -idium (for genus-group names of small animals; Werner 1972, p. 47) and refers to the fact that Foissner et al. (1990a) found 1 Foissner et al. (1990a) provided the following diagnosis: Spirofilidae (?) mit 1 kurzen und 1 langen Ventralreihe. Hinter dem Peristom 1 isolierter Cirrus zwischen der Ventralreihe und der linken Marginalreihe. Caudal- und Transversalcirren vorhanden.
Mucotrichidium
441
M. hospes in the mucilaginous loricae of Chironomus-larvae. Neuter gender (Foissner et al. 1990a). Characterisation (A = supposed apomorphy): Amphisiellidae(?) with continuous adoral zone of membranelles. Three frontal cirri. Buccal cirri present. More than one cirrus left of anterior portion of amphisiellid median cirral row, originate from anlage III only. Frontal-ventral-transverse cirri (likely) originate from six anlagen. Amphisiellid median cirral row likely originates from anlagen IV (middle portion), V (rear portion), and VI (front portion). Postperistomial cirrus, transverse (?) cirri, and caudal cirri present. One left and one right marginal row. Dorsal kineties originate by within row formation only. Lives in spawn of evertebrates (A). Remarks: See single species. Species included in Mucotrichidium (basionym given): (1) Uroleptus hospes Ehrenberg, 1831.
Single species Mucotrichidium hospes (Ehrenberg, 1831) Foissner, Oleksiv & Müller, 1990 (Fig. 90a–k, 91a–j, Table 28) 1831 Uroleptus hospes E. 1 – Ehrenberg, Abh. preuss. Akad. Wiss., year 1831: 116 (original description; no type material available). 1838 Uroleptus hospes – Ehrenberg, Infusionsthierchen, p. 359, Tafel XL, Fig. III (Fig. 90a–k; revision, first illustration). 1850 Uroleptus hospes Ehrenberg – Diesing, Systema Helminthum, 1: 153 (revision). 1990 Mucotrichidium hospes (Ehrenberg, 1831) nov. comb.2 – Foissner, Oleksiv & Müller, Arch. Protistenk., 138: 198, Abb. 4a–j, Tabelle 2 (Fig. 91a–j; detailed redescription, neotypification, and combination with Mucotrichidium; 2 neotype slides have been deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; slides, however, not mentioned by Aescht 2003). 1999 Mucotrichidium hospes (Ehrenberg, 1831) – Shi, Song & Shi, Progress in Protozoology, p. 104, Fig. 24 (schematic redrawing of Fig. 91i?; revision of hypotrichs). 1999 Mucotrichidium hospes (Ehrenberg, 1831) – Shi, Acta Zootax. sin., 24: 255, Fig. 23 (schematic redrawing of Fig. 91i?; revision of hypotrichs).
1
Ehrenberg (1831) provided the following diagnosis: Körperdurchmesser 1/20´´´ (about 105 µm). Körper grünlich, behaart, walzenförmig, vorn abgerundet, nach hinten abnehmend und geschwänzt; Schwanz conisch, ¼ der Körperlänge; Gleicht einem geschwänzten Börsenthierchen. Berlin. In den Hüllen des Froschlaichs. 2 Foissner et al. (1990a) provided the following improved diagnosis: In vivo etwa 115 × 50 µm großes, stark kontraktiles, lanzettförmiges Mucotrichidium, dessen Vorderende auffallend nach dorsal gebogen ist. Adorale Membranellenzone und rechter Rand des Buccalfeldes gelb gefärbt. 1 Mikronucleus zwischen den 2 Makronucleus-Teilen. 1 lange, in der Mitte unterbrochene rechte und 1 kurze frontale linke Ventralreihe. Durchschnittlich 49 adorale Membranellen, 5 Buccalcirren, 5 Transversalcirren in 2 Gruppen, 3 Caudalcirren und 3 gleich lange Dorsalkineten.
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SYSTEMATIC SECTION
Fig. 90a–k Mucotrichidium hospes (from Ehrenberg 1838. From life). a–e, i: Freely motile specimens. Note that the body shape and habitat agree rather perfectly with that of the neotype population so that the identification by Foissner et al. (1990a) is beyond reasonable doubt. f: Empty snail spawn; each egg contains one specimen of M. hospes. g, h, j: Distorted specimens. k: Snail egg with a specimen of M. hospes. Page 441.
2001 Mucotrichidium hospes (Ehrenberg, 1831) Foissner, Oleksiv and Müller, 1990 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 98 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Mucotrichidium hospes – Lynn & Small, Phylum Ciliophora, p. 448, Fig. 23A, B (Fig. 91f, i; guide to ciliate genera; in the figure legends ventral and dorsal are confused).
Nomenclature: The species-group name hospes (Latin noun; guest) refers to the fact that this species was found in the coat of a frogspawn (Ehrenberg 1831). Type species of Mucotrichidium. Remarks: Foissner et al. (1990a) established Mucotrichidium for a somewhat uncommon hypotrich living mainly in empty spawn of evertebrates. They assigned it provisionally to the Spirofilidae because the cirral rows are distinctly helical, at least in prepared specimens (Fig. 91e–j). Shi (1999) and Shi et al. (1999) transferred Mucotrichidium to the Amphisiellidae. I have no translations of the Chinese papers, but likely they proposed this classification because of the “amphisiellid” cirral pattern. Indeed, the cirral pattern of M. hospes closely resembles that of Hemiamphisiella terricola, especially as concerns the presence of the postperistomial cirrus. Furthermore, the amphisiellid median cirral row is likely also composed of components of the three rightmost anlagen (Fig. 91i): (i) the anterior portion consists of cirri of anlage VI (homologous to the frontoterminal cirri); (ii) the middle portion is likely short, that is, composed of only one cirrus (Fig. 91j, long arrow) formed by anlage IV (this is the same anlage that also forms the postperistomial cirrus!); and (iii) the posterior portion is composed of the cirri of anlage V. Of course, this assumption has to be checked by ontogenetic data. Consequently, I preliminarily assign, like the Chinese workers, Mucotrichidium to the amphisiellids. The main difference between the interphasic infraciliature of Mucotrichidium and Hemiamphisiella (p. 288) is in the number of cirri left of the anterior portion of the
Mucotrichidium
443
Fig. 91a–d Mucotrichidium hospes (from Foissner et al. 1990a. From life). a, b: Ventral views of representative specimens, a = 105 µm. Note the single micronucleus between the two macronuclear nodules and the ingested diatoms. Three cirri are inserted on the fold at the rear tip of the cell; they form a characteristic, stiletto-like protrusion. This species usually inhabits empty spawns of frogs, snails, insects, etc. c: Right lateral view showing that the anterior cell portion is turned upwards. d: Resting cyst, 45 µm. Page 441.
amphisiellid median cirral row, namely, more than one against only one. However, ontogenesis may also reveal further differences between these taxa. Possibly, the cirri between the posterior portion of the marginal rows are true transverse cirri (that is, originate from different anlagen) in M. hospes. By contrast, all these cirri originate from anlage VI in Hemiamphisiella (Eigner & Foissner 1994). The present species was discovered rather early by Ehrenberg. In the 19th century it was recorded, although without providing morphological data, several times from spawns of various evertebrates (see occurrence and ecology section). In the last century, this species was never recorded until Foissner et al. (1990a) rediscovered it in a pond in the city of Salzburg. Kahl (1932) overlooked this species in his revision, and Borror (1972, p. 12) synonymised it with Trichoda musculus Müller, 1773, a rather problematic species (possibly a rotifer) discovered in a putrid infusion (for brief review, see Foissner et al. 1991, p. 244, 248).
444
SYSTEMATIC SECTION
Mucotrichidium
445
Strongylidium mucicola Kahl, 1932 has a similar habitus (Foissner et al. 1990a). However, the two narrowly spaced cirral rows indicate that this species has a midventral complex composed of cirral pairs, that is, it is possibly a urostyloid (see Fig. 9415, 16 in Kahl 1932). Anyhow, synonymy with M. hospes is very unlikely because it has only one buccal cirrus (vs. many in M. hospes) and lacks the postperistomial cirrus (vs. present), which is easily recognisable even in live specimens. Generally, Strongylidium Sterki, 1878 is a difficult, little known group, whose type species has two frontoventral rows (Kahl 1932, p. 551). Quennerstedt (1869, p. 34) discussed the present species likely because he found a population closely resembling M. hospes. Since he did not provide an illustration I did not translate the paper. For comparison with Spiroamphisiella see there. Morphology: The paragraphs below are based on the very detailed description of the neotype population by Foissner et al. (1990a). The data by Ehrenberg (1838) are presented in the last paragraph. Body size of neotype specimens in life about 115 × 50 µm; body length:width ratio of specimens shown in Fig. 91a, b about 2.6–2.8:1, ratio of prepared specimens 2.3:1 on average (Table 28). Body slender to wide lanceolate, anterior end broadly rounded, posterior end tapered and with a fold bearing some cirri (Fig. 91a, b). Body not dorsoventrally flattened. Lateral view highly characteristic, because anterior body third strongly curved dorsally (Fig. 91c). Body very contractile (up to 50%), thus specimens distinctly smaller in preparations than in life; contracted cells ellipsoidal. Macronuclear nodules slightly left of midline in middle body third; individual nodules ellipsoidal, in life about 20 × 15 µm, contain many chromatin bodies. Micronucleus spherical, very large, that is, in life 5–7 µm across; almost invariably arranged in between the two macronuclear nodules (Fig. 91a, e, j); rarely one micronucleus per macronuclear nodule. Contractile vacuole about in mid-body at left cell margin, without collecting canals. Pellicle very flexible, colourless. Cortical granules lacking. Cytoplasm packed with greasily shining, colourless globules 1–5 µm across, some small crystals, and many food vacuoles, partly containing diatoms making cells yellow at low magnification. Movement burrowing, in the insectloricae often resting, and slowly rotating when freely motile; body often distorted due to movement.
b
Fig. 91e–j Mucotrichidium hospes (from Foissner et al. 1990a. Protargol impregnation). e, f: Infraciliature of left and right side and nuclear apparatus of same specimen, 62 µm. g, h: Infraciliature of left and right side of posterior body portion of same specimen. i, j: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 60 µm. Frontal cirri connected by dotted line. Broken lines connect cirri which (very likely) originate from same anlage. Long arrow marks a cirrus slightly set off from anterior end of posterior portion of amphisiellid median cirral row; very likely, it originates from the same anlage (namely anlage IV) as the postperistomial cirrus (short arrow). ACR = amphisiellid median cirral row, AZM = distal end of adoral zone of membranelles, CC = caudal cirri, FC = right frontal cirrus (usually distinctly smaller than the middle and left one), LMR = left marginal row, MI = micronucleus, P = paroral, RMR = right marginal rows, TC = transverse cirri, I–VI = frontal-ventral-transverse cirral anlagen (proposal has to be checked by ontogenetic data), 1–3 = dorsal kineties. Page 441.
446
SYSTEMATIC SECTION
Adoral zone occupies about 50%(!) of body length (44% on average in protargol preparations; Table 28), composed of 49 membranelles on average; largest membranelles about 7 µm wide in life. Zone extends relatively far onto right body margin. Buccal field deep, anteriorly curved hook-like. Paroral and endoral of equal length, each likely composed of two kineties. Paroral convex (anteriorly not hookshaped), endoral conspicuously sigmoidal. The yellowish colour of the adoral zone and the right margin of the buccal field is very conspicuous and was observed in all neotype specimens; the colouring is diffuse and not due to granules recognisable in the light microscope. Cirral pattern and number of cirri of usual variability, except for the number of buccal and caudal cirri, which vary rather strongly (Table 28). Frontal cirri 15 µm long, marginal and remaining cirri about 10 µm long. Three frontal cirri; left and middle one of about same size as other cirri; right cirrus distinctly smaller than other frontal cirri. Buccal cirri right of anterior half of paroral. On average seven (parabuccal) cirri behind right frontal cirrus, row terminates about at same level as buccal row. One postperistomial ventral cirrus distinctly behind proximal end of adoral zone; specimen shown in Fig. 91i with somewhat isolated cirrus (long arrow) ahead of rear portion of amphisiellid median cirral row, indicating that this cirrus and the postperistomial cirrus are formed, as in Hemiamphisiella, from anlage IV. Amphisiellid median cirral row with more or less distinct break, indicating that it is composed of cirri formed by anlage VI (anterior portion) and anlage V (posterior portion); amphisiellid row distinctly spiral in prepared specimens. Between posterior portion of marginal rows five cirri arranged in two groups; exact designation (rear part of amphisiellid median cirral row? transverse cirri? ...) impossible without ontogenetic data; rear group, which is composed of three cirri, arranged on fold of posterior end (see above) and forming conspicuous stiletto-shaped protrusion. These cirri are 25 µm, 20 µm, and 15 µm long and frayed on left side (Fig. 91a, h). Right marginal row commences near distal end of adoral zone, extends spirally to rear body end. Left marginal row commences, in prepared and therefore contracted(!) specimens, left of middle portion of adoral zone, traverses more or less distinctly dorsally to posterior body end. Dorsal bristles in life about 3 µm long, arranged in three almost bipolar, distinctly spiralled kineties. At rear end of each kinety one fine caudal cirrus, proving that dorsomarginal kineties and/or fragmentation are/is lacking. Infraciliature of posterior body portion rather difficult to understand. Observations by Ehrenberg (1838; Fig. 90a–k): For Ehrenberg’s (1831) diagnosis, see corresponding footnote. Ehrenberg found one specimen per snail egg; when he released them from the egg, they extended slowly and began to swim. He recognised a large mouth as longitudinal gap in a wide pit behind the anterior body end, about 20 food vacuoles, and 8–10 longitudinal rows of cilia. The last feature must not be over-interpreted because at Ehrenberg’s time it was extremely difficult (basically impossible) to recognise the cirral pattern of hypotrichs more or less correctly.
Terricirra
447
However, all other features, including the unusual microhabitat (spawn of various evertebrates) agree perfectly with the population studied by Foissner et al. (1990a). Resting cysts (Fig. 91d): According to Foissner et al. (1990a), cysts are spherical with smooth surface, 40–50 µm across (44.6 µm on average; SD = 3.1 µm; CV = 6.9%; n = 9). Content finely granulated and colourless. Ectocyst 3–4 µm thick, composed of many fine layers. Mesocyst 1–2 µm thick and compact. Occurrence and ecology: Mucotrichidium hospes is very likely confined to limnetic habitats. The type locality is, due to the neotypification, the pond at the Salzburg University (Austria), where Foissner et al. (1990a) found it in mid-July in empty, old, mucilaginous loricae of Chironomus-larvae. Jersabek & Schabetsberger (1996, p. 11; identification by W. Foissner) found it in the slime of old Ophrydium colonies in the pelagial of a small alpine karst lake (Dreibrüdersee, altitude 1643 m; Totes Gebirge) in Austria. Ehrenberg (1831) discovered it with high abundance in the coat of an empty frogspawn during April and in snailspawn during August in the city of Berlin, Germany. Further records: Vienna, Austria (Riess 1840, p. 22); in empty gnat eggs and surrounding mucus from the city of St. Petersburg, Russia (Weisse 1848, p. 355; Weisse also discussed that he very likely had, in an earlier paper, misidentified the present species as Uroleptus piscis, which he found in the same area in eggs of Nais); Quennerstedt (1869, p. 34) found it (or a similar species on/in Lymnea spawn?) likely in Sweden (see remarks). Stein (1854, p. 205) discussed that a Swiss worker could have observed the cysts of the present species in spawn of Lymnea stagnalis. Mucotrichidium hospes feeds on diatoms (Navicula sp., Cocconeis sp.), cyanobacteria (oscillatorians), and coccal and colonial green algae (Foissner et al. 1990a).
Terricirra Berger & Foissner, 1989 1989 Terricirra nov. gen.1 – Berger & Foissner, Bull. Br. Mus. nat. Hist. (Zool.), 55: 35 (original description). Type species (by original designation): Perisincirra viridis Foissner, 1982. 1997 Terricirra – Berger & Foissner, Arch. Protistenk., 148: 129 (brief note on the systematic position). 1999 Terricirra Berger & Foissner, 1989 – Berger, Monographiae biol., 78: 894 (brief note on systematic position). 2001 Terricirra Berger & Foissner 1989 – Aescht, Denisia, 1: 159 (catalogue of generic names of ciliates; see nomenclature). 2001 Terricirra Berger and Foissner, 1989 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 88 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Terricirra Berger and Foissner, 1989 – Lynn & Small, Phylum Ciliophora, p. 458 (guide to ciliate genera).
Nomenclature: No derivation of the name is given in the original description. Terricirra is a composite of terr·a (Latin noun; the earth, the land), the thematic vowel 1
Berger & Foissner (1989) provided the following diagnosis: Vermiform to long Oxytrichidae with green to blue subpellicular granules and spindle-shaped food vacuoles containing parallel arranged bacteria. Undulating membranes short, forming an acute angle. 1 short frontoventral row.
448
SYSTEMATIC SECTION
·i-, the Latin noun cirr·us (curl; cirrus, an important feature of the hypotrichs), and the inflectional ending -a; it refers to the fact that the members of this genus live in soil. Feminine gender (Aescht 2001, p. 302). Aescht (2001) incorrectly assumed that the original description of the present genus also contains a redescription of the type species Perisincirra viridis. Tericirra mutsusakai in Lynn & Small (2002, p. 458) is an incorrect subsequent spelling. Characterisation (A = supposed apomorphy): Amphisiellidae with continuous or interrupted (T. livida) adoral zone of membranelles. Undulating membranes short, form acute angle (A?). Three more or less distinctly enlarged frontal cirri. One buccal cirrus. One cirrus left of anterior portion of amphisiellid median cirral row. Amphisiellid median cirral row only slightly longer than adoral zone of membranelles. Postperistomial cirrus lacking. Pretransverse ventral cirri indistinct or lacking. Transverse cirri present. One right and one left marginal row. Three (type species), four, or one dorsal kinety(ies). Caudal cirri lacking (A?). Terrestrial. Additional characters: The three species included have some further features in common, namely, body very flexible, but not or only slightly contractile; macronuclear nodules in midline or slightly left of it; contractile vacuole at or somewhat ahead of mid-body, at left cell margin. Remarks: We assigned Terricirra to the Oxytrichidae, however, without providing a foundation (Berger & Foissner 1989). Later, Berger & Foissner (1997) transferred it from the Oxytrichidae to the Amphisiellidae because of the similarity with Lamtostyla, which was assigned to the amphisiellids by Petz & Foissner (1996) based on ontogenetic data. Lynn & Small (2002) classified Terricirra in the Trachelostylidae (p. 471), together with Hemisincirra, Trachelostyla, Lamtostyla, and Gonostomum. However, Terricirra lacks the trachelostylid oral apparatus and therefore the position supposed by Berger & Foissner (1997) is retained. Perisincirra viridis, which was fixed as type species of Terricirra, has three dorsal kineties of body length (Fig. 92e). By contrast, Terricirra matsusakai has four kineties and kinety 4 terminates ahead of mid-body, strongly indicating that this is a dorsomarginal row (Fig. 92e). Thus, the generic assignment of T. matsusakai is uncertain and therefore ontogenetic and molecular data are needed for a more detailed discussion about the monophyly of Terricirra. Shi (1999, p. 255) and Shi et al. (1999, p. 103) put Terricirra into the synonymy of Lamtostyla, which they assigned to the amphisiellids. Since these papers are written in Chinese I do not know the foundation for this step. Indeed, the cirral pattern of Lamtostyla, Lamtostylides, and Terricirra is very similar so that it is somewhat a matter of taste to accept a group or to submerge it in an older, similar one. In the present book I establish Lamtostylides for Lamtostyla species which have only one cirrus (usually III/2) left of the anterior portion of the amphisiellid median cirral row. This pattern is due to the lack of the cirral anlage IV. Terricirra species also have only one cirrus left of the anterior portion of the frontoventral row, indicating that they also lack anlage IV. However, in T. viridis (type species) the ordinary six anlagen can be present as indicated by the rather irregular amphisiellid median cirral row. Unfortu-
Terricirra
449
nately, ontogenetic data are not available for any of the three species of Terricirra so that a more detailed discussion is impossible and a thorough comparison of Terricirra and Lamtostylides cannot be made. However, Terricirra was established, inter alia, because of its curious, spindle-shaped food vacuoles containing parallel-arranged bacteria and the distinct, green or blue cortical granules (Berger & Foissner 1989). Further, Terricirra species have a special undulating membrane-pattern. Thus, the synonymy of Lamtostyla and Terricirra proposed by the Chinese authors is likely not justified. Relevant molecular data have to be awaited for a final decision. Spindle-shaped food vacuoles and distinctly coloured cortical granules are lacking in Lamtostylides species. Only L. halophila has the same type of food vacuoles (Fig. 66a), but colourless cortical granules and more or less ordinary undulating membranes; it is therefore mentioned in the key to Terricirra species. In both Terricirra and Lamtostylides, the undulating membranes are rather short. However, in Lamtostylides they are arranged roughly in parallel, whereas they form an acute angle in Terricirra. Perhaps these two genera are the sister group to Lamtostyla. A further group likely closely related to Terricirra, Lamtostylides, and Lamtostyla is Hemisincirra, a rather heterogeneous group comprising slender, terrestrial species with a short to long frontoventral row (see there for details). Species included in Terricirra (alphabetically arranged basionyms are given): (1) Hemisincirra livida Berger & Foissner, 1987; (2) Perisincirra viridis Foissner, 1982; (3) Terricirra matsusakai Berger & Foissner, 1989.
Key to Terricirra species and similar species If you know that your specimen/population belongs to Terricirra, species identification is rather simple because the species can be distinguished by body shape and the number of macronuclear nodules. If you are unsuccessful with the key below, see also Lamtostylides (p. 322), Lamtostyla (p. 161), or Hemisincirra (p. 387). Lamtostylides halophilus is also included in the key below because it has, like Terricirra spp., spindle-shaped food vacuoles. 1 Body vermiform; adoral zone with distinct gap (Fig. 94a, g). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Terricirra livida (p. 458) - Body wide to elongate elliptical; adoral zone without gap (Fig. 92a, 93a). . . . . . 2 2 Eight macronuclear nodules (Fig. 92a, d). . . . . . . . . . . Terricirra viridis (p. 450) - Four or two macronuclear nodules (Fig. 66a, c, 93a, e,). . . . . . . . . . . . . . . . . . . . 3 3 Four macronuclear nodules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 - Two macronuclear nodules; cortical granules colourless (Fig. 66a, e). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lamtostylides halophilus (p. 336) 4 Adoral zone of membranelles without gap; four dorsal kineties (Fig. 93a–h). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Terricirra matsusakai (p. 454) - Adoral zone of membranelles with gap; three dorsal kineties (Fig. 83a, b). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemisincirra quadrinucleata (p. 419)
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SYSTEMATIC SECTION
Terricirra viridis (Foissner, 1982) Berger & Foissner, 1989 (Fig. 92a–j, Table 29) 1982 Perisincirra viridis nov. spec.1 – Foissner, Arch. Protistenk., 126: 92, Abb. 23a–e, 71, 73, Tabelle 21 (Fig. 92a–f, i, j; original description. Type slides likely deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; see nomenclature). 1984 Hemisincirra viridis (Foissner, 1982) nov. comb. – Foissner, Stapfia, 12: 119 (combination with Hemisincirra; see nomenclature). 1989 Terricirra viridis (Foissner, 1982) nov. comb. – Berger & Foissner, Bull. Br. Mus. nat. Hist. (Zool.), 55: 36 (combination with Terricirra). 2001 Terricirra viridis (Foissner, 1982) Berger and Foissner, 1989 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 72 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs).
Nomenclature: No derivation of the name is given in the original description. The species-group name virid·is, -is, -e (Latin adjective [m; f; n]; green) refers to the colour of the cortical granules making the cell green at low magnification. Perisincirra viridis is type species of Terricirra. Aescht (2003, p. 399) does not mention this species in the list of type specimens deposited in the museum in Linz. However, other species described by Foissner (1982) are mentioned, indicating that it was overlooked in this list. “Perisincira viridis Foissner” in Tirjaková (1988, p. 500) is an incorrect subsequent spelling of Perisincirra. Foissner (1984) transferred the present species to Hemisincirra before this genus was established by Hemberger (1985). Since this generic assignment is not relevant at present I do not correct this obviously invalid act. Remarks: Foissner (1982) provisionally assigned this species to Perisincirra Jankowski, 1978, a little-known genus based on the type species Uroleptus kahli Grolière, 1975. Somewhat later, Foissner (1984) transferred it to Hemisincirra (for nomenclatural problems see previous chapter), because Perisincirra and Hemisincirra are quite different taxa (for details on the complicated systematics, see Hemisincirra). In 1989, we fixed P. viridis as type species of Terricirra, which is mainly characterised by the spindle-shaped food vacuoles, the prominent, green cortical granules, and the short undulating membranes forming an acute angle (details see genus section). Terricirra matsusakai has only four macronuclear nodules (vs. eight in T. viridis), but four dorsal kineties (vs. three). Hemisincirra octonucleata has more or less the same nuclear apparatus, but only half the number of marginal cirri (e.g., 12–15 right marginal cirri vs. 24–32) and a distinct break in the adoral zone (vs. absent) (Fig. 84a, b). Morphology: The following description is based on the original description, supplemented by two micrographs from the population found by Foissner et al. 1 Foissner (1982) provided the following diagnosis: In vivo etwa 140–190 × 30–45 µm große, durch ungefähr 15 Reihen dunkelgrüner subpelliculärer Granula bereits bei kleiner Vergrößerung auffallend grün gefärbte Perisincirra mit kettenförmigem Makronucleus, der aus 8 ellipsoiden Nodien besteht. Adorale Membranellenzone etwa 1/6 körperlang, im unteren Abschnitt S-förmig gebogen. 3 Dorsalkineten.
Terricirra
451
Fig. 92a–f Terricirra viridis (from Foissner 1982. a–c, from life; d–f, protargol impregnation). a: Ventral view of representative specimen, 167 µm. b: Left-lateral view. c: Dorsal view showing contractile vacuole and cortical granules, 140 µm. d–f: Infraciliature of ventral (d, f; same specimen) and dorsal side and nuclear apparatus, d = 82 µm, e = 86 µm. Arrow in (e) marks the single dorsal bristle. Broken lines in (f) connect cirri which (very likely) originate from the same anlage. Frontal cirri connected by dotted line, cirri within rectangle likely homologous to the four frontoventral cirri and cirri within ellipse likely homologous to postoral ventral cirri of the 18-cirri hypotrichs (details see text). Accordingly, the amphisiellid median cirral row of T. viridis is possibly composed of cirri from anlage VI (anterior portion), IV (middle portion), and V (posterior portion). However, ontogenetic data are needed to confirm this interpretation. CV = contractile vacuole, MA = macronuclear nodule, MI = micronucleus, RMR = right marginal row, TC = transverse cirri (possibly, pretransverse ventral cirri are also present), I–VI = frontal-ventral-transverse cirri anlagen, 1–3 = dorsal kineties. Page 450.
(2005; Fig. 92g, h). Body size in life 140–190 × 30–45 µm, body length:width ratio 4.7:1 on average in protargol preparations (Table 29). Body outline elongate elliptical, both ends rounded (Fig. 92a, c, h). Body very flexible and slightly contractile
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SYSTEMATIC SECTION
Fig. 92g–j Terricirra viridis (g, h, original micrographs of the population found by Foissner et al. 2005, kindly supplied by W. Foissner; i, j, from Foissner 1982. g, h, differential interference contrast; i, bright field; j, protargol impregnation). g, i: The cortical granules are dark-green, about 0.8 µm across, and arranged in short, usually longitudinal rows. h: Nuclear apparatus and contractile vacuole. Arrows mark micronuclei. j: Infraciliature of oral region. Right frontal cirrus and cirrus III/2 connected by broken line. AZM = distal end of adoral zone, BC = buccal cirrus, CV = contractile vacuole, MA = macronuclear nodule. Page 450.
under cover glass; about 2:1 flattened dorsoventrally (Fig. 92b). Eight macronuclear nodules form row slightly left of midline in central body portion; individual nodules connected by fine strand; chromatin bodies large, cloddy. One micronucleus each near anterior and posterior end of macronuclear figure; individual micronuclei about 4 µm across in life (Fig. 92d, h, j). Contractile vacuole slightly ahead of mid-body near left cell margin, without collecting canals (Fig. 92c, h). Pellicle and cytoplasm colourless. Cortical granules (termed “subpellicular granules” in original description) arranged in longitudinal rows on ventral side; on the dorsal side they are arranged in short, oblique rows which form indistinct longitudinal stripes; individual granules about 0.8 µm across, dark-green. Granules occur mainly along cirral rows and dorsal kineties (Fig. 92c, g, i); after cells die granules bleach out and dissolve.
Terricirra
453
Cytoplasm packed with 8–15 × 3–6 µm-sized, invariably spindle-shaped food vacuoles containing exclusively bacteria; food vacuoles squeezed out from cell become globular. Cytoplasm also contains 1–3 µm-sized, colourless, brilliant globules and tiny, yellowish crystals. Movement slow, gliding, stands still for some time. Adoral zone occupies only about 17% of body length and composed of 18 membranelles on average; zone always characteristically curved as shown in Fig. 92d, f. Bases of largest membranelles about 4 µm wide. Buccal field very small, rather flat. Paroral slightly curved and endoral straight, form acute angle. Pharyngeal fibres long, very conspicuous in protargol preparations. Cirral pattern and number of cirri of usual variability (Fig. 92d, f, Table 29). All cirri relatively fine and short, bases of about same size. Frontal cirri form slightly oblique pseudorow with right cirrus behind distal end of adoral zone. Buccal cirrus right of endoral and slightly ahead of paroral. One cirrus (= cirrus III/2) left of anterior portion of amphisiellid median cirral row, which terminates at 25% of body length on average (Table 29), composed of 5.4 cirri on average which are somewhat irregularly arranged, so that one cannot exclude that the row is composed of two or three portions. Specimen illustrated, like 18-cirri hypotrichs, with 11 frontal-ventral cirri, that is, one cannot exclude that the frontal-ventral-transverse cirri of T. viridis originate from six anlagen although only one cirrus clearly arranged left of amphisiellid median cirral row which, if the designation in Fig. 92f is correct, is made of three portions (Fig. 92d, f). Transverse cirri arranged in V- or hook-shaped, slightly subterminal row, about 12 µm long in life, project, like 10-µm-long marginal cirri, slightly beyond body margin. Whether pretransverse ventral cirri are present or not can be decided only after studying cell division. Right marginal row commences slightly ahead of level of anterior end of amphisiellid median cirral row. Left marginal row commences left of proximal end of adoral zone, terminates, like right row, about at level of transverse cirri; marginal rows thus widely separated posteriorly. Distance between individual cirri almost equal within each row (Fig. 92d). Dorsal bristles about 5 µm long in life, arranged in three kineties (Fig. 92e); kineties 1 and 2 (incorrectly designated as “right kineties” in original description) slightly shortened anteriorly, kinety 3 roughly bipolar; bristles originate from small, square bases; bases in rearmost sixth of kineties sometimes without bristles. Left of anterior end of kinety 3 (invariable?) a single bristle (Fig. 92e, arrow). Caudal cirri lacking. Occurrence and ecology: Very likely confined to terrestrial habitats and possibly of cosmopolitan distribution because recorded from Holarctis, Paleaotropis, and Australis (Foissner 1987, 1998). Type locality of T. viridis is a bottom land (15°45'E 48°23'N) near the village of Grafenwörth, Lower Austria, where Foissner (1982) discovered it in soil from a dry side arm of the Danube river (detailed description of this locality, see Foissner et al. 1985, p. 88, site “AV”). Abundance was very low. Further records: soil from a beech forest near the city of Salzburg, Austria (Foissner
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SYSTEMATIC SECTION
et al. 2005, p. 632; Fig. 92g, h); agricultural soil from a farm near the village of Ostrov, Slovakia (Tirjaková 1988, p. 500; see also Matis et al. 1996, p. 19). Feeds on bacteria, which are arranged in parallel in spindle-shaped food vacuoles (Fig. 92a; Foissner 1982). Biomass of 106 specimens about 33 mg (Foissner 1987, p. 124; 1998, p. 210).
Terricirra matsusakai Berger & Foissner, 1989 (Fig. 93a–h, Table 29) 1989 Terricirra matsusakai nov. spec.1 – Berger & Foissner, Bull. Br. Mus. nat. Hist. (Zool.), 55: 36, Fig. 56–60, Table 9 (Fig. 93a–e; original description; the holotype slide [reference number 1988:4:10:1] and the paratype side [1988:4:10:2] have been deposited in the British Museum of Natural History in London, England). 2001 Terricirra matsusakai Berger and Foissner, 1989 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 88 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Tericirra mutsusakai – Lynn & Small, Phylum Ciliophora, p. 458, Fig. 54A, B (Fig. 93d, e; guide to ciliate genera; incorrect subsequent spelling of both genus and species-group name). 2002 Terricirra matsusakai Berger & Foissner, 1989 – Foissner, Agatha & Berger, Denisia, 5: 868, Fig. 381v, 388f, g (Fig. 94f–h; record from Namibia and micrographs of live specimens).
Nomenclature: We dedicated this species to Tadao Matsusaka, University of Kumamoto (Japan), who collected the soil sample harbouring the species (Berger & Foissner 1989). Remarks: For separation from congeners see key. Hemisincirra quadrinucleata is also very similar, inter alia, in the arrangement of the undulating membranes (Fig. 83a, b). However, it has fewer dorsal kineties than T. matsusakai (3 vs. 4), a discontinuous adoral zone (vs. continuous), and a more anteriorly arranged contractile vacuole. Since Hemberger (1982, 1985) gave no information about the food vacuoles and the cortical granulation, Hemisincirra quadrinucleata was not transferred to Terricirra by Berger & Foissner (1989). The amphisiellid median cirral row of T. matsusakai is usually composed of four cirri only. I suppose that these cirri are homologous to the frontoterminal cirri (= cirri VI/4 and VI/3) and the anteriorly displaced postoral ventral cirri V/4 and V/3 of the 18-cirri oxytrichids. The presence of only one cirrus (likely III/2) left of the amphisiellid median cirral row indicates that, as in Lamtostylides, anlage IV is lacking. Dorsal kinety 4 of T. matsusakai is distinctly shortened posteriorly, strongly indicating that it is a dorsomarginal row. However, ontogenetic data are needed for a correct interpretation of the cirral and dorsal kinety pattern and therefore for a more proper classification. In the Namibian specimens we could confirm two main features of Terricirra matsusakai and Terricirra in general, namely, the dark-green cortical granules and 1
Berger & Foissner (1989) provided the following diagnosis: In vivo c. 125–135 × 27–30 µm. Subpellicular granules dark green, spherical, c. 1 µm in diameter. 18 adoral membranelles on average, 4 macronuclear segments, 4 dorsal kineties.
Terricirra
455
Fig. 93a–e Terricirra matsusakai (from Berger & Foissner 1989. a–c, from life; d, e, protargol impregnation). a: Ventral view of representative specimen, 135 µm. b: Right lateral view. c: Dorsal view showing arrangement of dark-green, spherical cortical granules and contractile vacuole with distinct collecting canals during diastole. Arrow marks frontal scutum. d, e: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 83 µm. Arrow in (d) marks buccal cirrus. Note the undulating membranes which form an acute angle. ACR = anterior end of amphisiellid median cirral row, AZM = adoral zone of membranelles, CG = cortical granules, LMR = left marginal row, MA = macronuclear nodule, Mi = micronucleus, TC = transverse cirri, 1–4 = dorsal kineties. Page 454.
the ellipsoidal food vacuoles (Fig. 93f–h). Both are very conspicuous, but require observation at magnifications of 100× or more. Morphology: This chapter is based on the original description supplemented by some data from a Namibian population studied by Foissner et al. (2002; Fig. 93f–h). Body size in life about 125–135 × 27–30 µm; body length:width ratio 3.9:1 on average in protargol preparations (Table 29). Body margin parallel, in area of adoral zone distinctly converging anteriorly; both ends rounded. Body very flexible, slightly contractile, flattened about 1.5–2.0:1 dorsoventrally (Fig. 93b). Four macronuclear nodules roughly arranged in pairs in cell midline or slightly left of it; indi-
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SYSTEMATIC SECTION
Fig. 93f–h Terricirra matsusakai (from Foissner et al. 2002. From life). f: Namibian specimen showing Terricirraspecific, ellipsoidal to fusiform food vacuoles. g, h: Cortical granulation of dorsal side. The granules are arranged in loose rows and very conspicuous because dark-green (thus black in bright field micrograph) and rather compact (thus refractive under interference contrast). CG = cortical granules, FV = spindle-shaped food vacuoles. Page 454.
vidual nodules about 15 × 11 µm in life, with small chromatin bodies. 1–3 micronuclei attached to macronuclear nodules (Fig. 93e). Contractile vacuole about in midbody near left cell margin, during diastole with distinct collecting canals (Fig. 93c). About six rows of widely spaced cortical granules (termed “subpellicular granules” in original description) on ventral and dorsal surface, respectively; individual granules dark green, spherical, and about 1 µm across (Fig. 93c, g, h). Cytoplasm colourless, with many, about 7–9 µm long, spindle-shaped food vacuoles, 1–2 µm large fat globules, and cytoplasmic crystals. Movement without peculiarities.
Terricirra
457
Adoral zone occupies about 19% of body length, composed of 18 membranelles on average of ordinary fine structure. Buccal field flat and narrow, posterior area covered by hyaline buccal lip. Undulating membranes short and inconspicuous, form acute angle. Pharyngeal fibres clearly recognisable in life and in protargol preparations, extend longitudinally backwards (Fig. 93a, d). Cirral pattern and number of cirri of usual variability (Fig. 93d, Table 29). Frontal cirri not distinctly larger than remaining cirri, arranged in oblique pseudorow with right cirrus, as is usual, behind distal end of adoral zone. Buccal cirrus right of anterior (endoral?) undulating membrane. Usually one cirrus (= cirrus III/2) left of anterior portion of amphisiellid median cirral row, respectively, behind right frontal cirrus. Amphisiellid median cirral row composed of 3–5, usually four cirri, terminates somewhat behind level of buccal vertex (at 25% of body length in specimen shown in Fig. 93d). Distinct pretransverse ventral cirri lacking. Transverse cirri about 15 µm long in life, arranged in short, longitudinal, slightly subterminal pseudorow. Right marginal row commences about at level of buccal cirrus, terminates at rear cell end. Left marginal row begins left of proximal end of adoral zone, terminates somewhat ahead of rear cell end, marginal rows thus slightly separated posteriorly. Marginal cirri about 10 µm long in life (Fig. 93d). Dorsal bristles about 2 µm long in life, arranged in four kineties. Kinety 1 shortened at both ends, kineties 2 and 3 roughly of body length; kinety 4 terminates slightly ahead of mid-body, indicating that it is a dorsomarginal row. Caudal cirri lacking (Fig. 93e). Cell division: Ontogenesis commences with the apokinetal formation of the oral primordium in about the middle of the cell (Berger & Foissner 1989). Occurrence and ecology: Very likely confined to terrestrial habitats and of cosmopolitan distribution because recorded from Holarctis, Paleaotropis, Australis, and Neotropis (Foissner 1998, p. 210). Type locality of T. matsusakai is the village of Kyokushi, Kumamoto Prefecture (Japan), where we discovered it in a soil sample (pH = 4.5) from a rice field collected in March 1985. Further records: forest soil (Pruno-Fraxinetum; calcaric fluvisol; typical mull) from eastern Austria and from a beech forest near the city of Salzburg (Foissner et al. 2005, p. 632); soil sample from a tropical dry forest about 5 km east of the ranch house “La Casona” in the Santa Rosa National Park in Costa Rica (Foissner 1995, p. 39); three soil samples from the Shimba Hills Nature Reserve, about 40 km south of the city of Mombasa and about 20 km west of the Indian Ocean coast (Foissner 1999, p. 324); mud from flat rockpools without grass (pH 6; not saline) from the escarpment of the Central Namib Desert, Spitzkoppe area about 120 km north-east of the city of Swakopmund, Namibia (Foissner et al. 2002, p. 63b). Feeds on bacteria, which are arranged in parallel in the spindle-shaped food vacuoles (Fig. 93a, f; Berger & Foissner 1989, p. 35; Foissner et al. 2002). Biomass of 106 specimens about 45 mg (Foissner 1998, p. 210).
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SYSTEMATIC SECTION
Terricirra livida (Berger & Foissner, 1987) Berger & Foissner, 1989 (Fig. 94a-i, Table 29) 1987 Hemisincirra livida nov. spec.1 – Berger & Foissner, Zool. Jb. Syst., 114: 211, Fig. 40–48, Table 7 (Fig. 94a–i; original description; the holotype slide [accession number 1986/63] and one paratype slide [1986/64] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; Aescht 2003, p. 389). 1989 Terricirra livida (Berger & Foissner, 1987) nov. comb. – Berger & Foissner, Bull. Br. Mus. nat. Hist. (Zool.), 55: 36 (combination with Terricirra). 2001 Terricirra livida (Berger and Foissner, 1987) Berger and Foissner, 1989 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 30 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs).
Nomenclature: No derivation of the name is given in the original description. The species-group name livid·us, -a, -um (Latin adjective [m; f; n]; livid) refers to the livid colour of the cortical granules. Remarks: In the original description we assigned this species to Hemisincirra because of the slender body and the cirral pattern which is similar to that of Uroleptus kahli Buitkamp, 1977, type of Hemisincirra Hemberger, 1985 (Berger & Foissner 1987). Somewhat later, we transferred it to Terricirra because of the agreement in the cortical granulation and the specific, spindle-shaped food vacuoles (Berger & Foissner 1989). The undulating membranes of T. livida are too small to be studied in detail in protargol preparations; in T. viridis and T. matsusakai they form an acute angle. Morphology: Body size in life 110–155 × 10–25 µm (n = 4), body length:width ratio 7.2:1 on average in protargol preparations (Table 29). Body vermiform, often strongly twisted about main body axis; body margins converging anteriad and posteriad, sometimes nearly parallel; anterior end rounded, posterior end tapered and usually bent to the left in ventral view (Fig. 94a); anterior fifth very thin, remaining body not flattened dorsoventrally, but constantly tapered posteriorly; flexible but not contractile. 13–18, on average 16 macronuclear nodules in central body portion and, as is usual, slightly left of midline; individual nodules spherical, ellipsoidal, or Fig. 94a–i Terricirra livida (from Berger & Foissner 1987. a–d, from life; e–i, protargol impregnation). a: Ventral view of representative specimen, 118 µm. b–d: Shape variants in dorsal view showing contractile vacuole (c) and cortical granules (d). e, f: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 124 µm. g–i: Infraciliature of the anterior body portion in ventral (g, i) and dorsal (h) view, g, h = 50 µm, i = 32 µm. Arrow in (g) denotes tiny buccal cirrus. Dotted line in (i) connects frontal cirri (see text for details). The undulating membranes are too small to be recognised in detail. ACR = amphisiellid median cirral row, AZM = bipartite adoral zone of membranelles, FC = right frontal cirrus, LMR = left marginal row, MA = macronuclear nodule, MI = micronucleus, RMR = right marginal row, 1 = dorsal kinety. Page 458. 1
Berger & Foissner (1987) provided the following diagnosis: In vivo about 110–155 × 10–25 µm (n = 4), vermicular, often strongly twisted, blue-green to livid subpellicular granules and spindle-shaped food vacuoles. About 16 adoral membranelles. 1 dorsal kinety. Remarks: “16 adoral membranelles” in this diagnosis is incorrect; on average, Terricirra livida has only about 11 membranelles (Table 29).
d
Terricirra
459
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SYSTEMATIC SECTION
Table 29 Morphometric data on Terricirra livida (liv, from Berger & Foissner 1987), Terricirra matsusakai (mat, from Berger & Foissner 1989), and Terricirra viridis (vir, from Foissner 1982) Characteristics a Body, length
Body, width
Adoral zone of membranelles, length
Posterior macronuclear nodule, length
Posterior macronuclear nodule, width
Posterior micronucleus, length
Posterior micronucleus, width
Species mean liv mat vir liv mat vir liv mat vir liv mat vir d liv mat vir d liv mat vir d liv mat vir liv liv
Nuclear apparatus, length Anterior body end to anterior end of nuclear apparatus, distance Anterior body end to posterior end of liv amphisiellid median cirral row, distance vir Adoral membranelles, number liv mat vir Macronuclear nodules, number liv mat vir Micronuclei, number liv mat vir Frontal cirri, number liv mat vir e Buccal cirri, number liv mat vir Amphisiellid median cirral row, number liv of cirri mat vir Cirri left of amphisiellid median cirral mat row, number Left marginal row, number of cirri liv mat
M
SD
SE
CV
Min
Max
n
106.9 93.2 86.9 14.8 23.9 18.4 11.1 17.5 14.4 6.6 12.0 7.6 3.1 5.7 5.1 1.8 1.6 2.5 1.8 1.6 2.5 65.4 14.1
105.0 95.0 80.0 14.0 23.0 18.0 11.0 17.0 14.0 7.0 13.0 7.0 3.0 6.0 5.0 1.7 1.6 2.5 1.7 1.6 2.5 70.0 14.0
8.6 11.2 13.1 1.5 5.3 1.8 1.7 1.2 1.2 0.5 2.1 1.0 0.5 1.0 0.7 0.2 0.0 0.1 0.2 0.0 0.1 18.8 1.9
2.6 3.4 4.6 0.4 1.6 0.6 0.5 0.4 0.4 0.2 0.6 0.4 0.2 0.3 0.2 0.1 0.0 0.0 0.1 0.0 0.3 6.7 0.6
8.0 12.0 15.0 9.9 22.1 9.6 15.3 6.9 8.2 7.6 17.9 13.4 17.5 17.6 12.8 8.9 2.6 3.4 8.9 2.6 3.4 28.7 13.3
98.0 122.0 78.0 112.0 67.0 107 13.0 17.0b 18.0 35.0 16.0 22.0 8.0 14.0 15.0 20.0 13.0 16.0 6.0 7.0 8.0 14.0 6.6 9.0 2.0 4.0 4.0 7.0 4.1 6.0 1.6 2.0 1.5 1.6 2.3 2.6 1.6 2.0 1.5 1.6 2.3 2.6 13.0 84.0 10.0 17.0
11 11 7 11 11 7 11 11 7 11 11 7 11 11 7 11 11 7 11 11 7 11 11
16.7 22.0 10.8 18.2 18.3 15.9 4.0 8.0 2.1 1.9 2.0 3.0 3.0 4.0 1.0 1.0 1.0 8.4 4.0 5.4 1.1
16.0 23.0 11.0 18.0 18.0 16.0 4.0 8.0 2.0 2.0 2.0 3.0 3.0 4.0 1.0 1.0 1.0 8.0 4.0 6.0 1.0
2.1 3.2 0.6 1.3 0.7 1.5 0.0 0.0 0.3 0.8 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1.0 0.5 0.9 0.3
0.6 1.2 0.2 0.4 0.3 0.4 0.0 0.0 0.1 0.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.3 0.1 0.3 0.1
12.3 14.6 5.6 6.9 3.8 9.1 0.0 0.0 14.4 43.5 0.0 0.0 0.0 0.0 0.0 0.0 0.0 12.3 11.2 16.6 27.6
15.0 17.0 10.0 16.0 17.0 13.0 4.0 8.0 2.0 1.0 2.0 3.0 3.0 4.0 1.0 1.0 1.0 6.0 3.0 4.0 1.0
20.0 28.0 12.0 20.0 19.0 18.0 4.0 8.0 3.0 3.0 2.0 3.0 3.0 4.0 1.0 1.0 1.0 10.0 5.0 6.0 2.0
11 7 11 11 7 11 11 7 11 11 7 11 11 7 11 11 7 11 11 7 11
53.1 30.3
51.0 29.0
5.8 3.7
1.8 1.1
10.9 12.1
42.0 23.0
63.0 36.0
11 11
Terricirra
461
Table 29 Continued Characteristics a Left marginal row, number of cirri Right marginal row, number of cirri
Transverse cirri, number
Dorsal kineties, number
Dorsal kinety 1, number of basal bodies
Species mean vir liv mat vir liv c mat vir liv mat vir liv
36.4 69.4 32.2 34.7 2.0 3.2 5.0 1.0 4.0 3.0 19.9
M
SD
SE
CV
Min
37.0 69.0 32.0 35.0 2.0 3.0 5.0 1.0 4.0 3.0 20.0
2.7 6.0 4.4 1.0 0.0 0.6 0.9 0.0 0.0 0.0 1.9
1.0 1.8 1.3 0.4 0.0 0.2 0.3 0.0 0.0 0.0 0.6
7.5 8.6 13.7 3.0 0.0 19.0 18.5 0.0 0.0 0.0 9.4
33.0 56.0 26.0 33.0 2.0 3.0 3.0 1.0 4.0 3.0 17.0
Max 42.0 77.0 41.0 36.0 2.0 5.0 6.0 1.0 4.0 3.0 23.0
n 7 11 11 7 10 11 7 11 11 7 11
a All measurements in µm. CV = coefficient of variation in %, M = median, Max = maximum value, mean = arithmetic mean, Min = minimum value, n = number of individuals investigated, SD = standard deviation, SE = standard error of arithmetic mean. Data based on protargol-impregnated specimens. b
In the original description 7.0 is incorrectly given.
c
Designated as caudal cirri in original description (see description for details).
d
Macronuclear nodule or micronucleus measured (anterior, posterior?) not indicated.
e
Cirrus left of anterior portion of amphisiellid median cirral row (= cirrus III/2) included.
dumbbell-shaped, with medium-sized chromatin bodies. On average two micronuclei in variable position near macronuclear apparatus (Fig. 94e, h). Contractile vacuole, as is usual, near left body margin, distinctly ahead of mid-body (at about 30% of body length in specimen illustrated; Fig. 94c); during diastole with distinct collecting canals. Cortical granules (termed “subpellicular granules” in original description) arranged in short rows; individual granules about 1.5 µm across and blue-green to livid so that cells have a bluish shimmer at low magnification (Fig. 94d). Cytoplasm colourless, with numerous 1–3 µm-sized, colourless globules and many 7–9 × 2–4 µm-sized, spindle-shaped food vacuoles with parallel arranged bacteria. Movement very slow, worm-like. Adoral zone short, that is, only about 10% of body length on average in protargol preparations (Table 29); composed of about 11 membranelles on average, three distal membranelles distinctly separated from proximal, nearly perpendicular portion of zone (Fig. 94a, e, g, i). Buccal area very small. Undulating membranes short, too small to be studied in detail with the light microscope. Pharyngeal fibres long, that is, extend to near mid-body. Cirral pattern and number of cirri of usual variability (Fig. 94e–i, Table 29). Three enlarged cirri near anterior end; designation uncertain because it is not known whether the rear cirrus is the right frontal cirrus or cirrus III/3; on the other hand, one can also not exclude that both the right frontal cirrus and cirrus III/2 are present, and, for example, the left frontal cirrus is lacking. I preliminarily designate the three
462
SYSTEMATIC SECTION
enlarged cirri as frontal cirri; ontogenetic data are needed for a final interpretation of this pattern. Left frontal cirrus close to middle of three distalmost membranelles, middle cirrus ahead of right frontal cirrus which is close to anterior end of amphisiellid median cirral row. Frontal cirri and anteriormost cirrus of right marginal row composed of 4–6 cilia, remaining cirri made of two cilia only, that is, very fine. Amphisiellid median cirral row composed of about eight cirri on average, commences close to right frontal cirrus and terminates at 16% of body length on average, that is, distinctly longer than adoral zone (Table 29). Transverse cirri lacking and caudal cirri present according to original description; however, since the posterior body portion is so narrow it is basically impossible to decide unequivocally whether these cirri are transverse or caudal cirri; ontogenetic data are needed for a correct interpretation. Right marginal row commences at anterior cell end, terminates at tip of body; anteriormost cirrus slightly enlarged (Fig. 94h). Left marginal row commences at proximal end of adoral zone and makes, like right row, nearly one turn to posterior end of cell (Fig. 94e, f). Dorsal bristles 2–3 µm long in life, arranged in only one kinety of body length. According to original description two caudal cirri present; however, whether these are caudal cirri or transverse cirri cannot be decided without ontogenetic data; since only one dorsal kinety is present, two cirri would originate from this kinety, a hint that these are not caudal cirri, but transverse cirri (Fig. 94a, e). Left of anterior end of right marginal row usually two dorsal bristles (Fig. 94f, h). Their origin (remnant of parental row? anterior end of long dorsal kinety 1? short dorsomarginal kinety?) is not known. A similar pattern is present in Vermioxytricha (Fig. 126m, p). Occurrence and ecology: Very likely confined to terrestrial habitats and possibly of cosmopolitan distribution because recorded from Holarctis, Palaeotropis, and Australis (Foissner 1987, 1998). Type locality of T. livida is a goat pasture (about 1000 m altitude) between the cities of Nauplion and Tripolis, Peloponnese (Greece), where we discovered it in the litter and upper soil layer (Berger & Foissner 1987). About two months after the discovery of the type population we found a second population in an arable soil near the city of Vienna, Austria (Berger & Foissner 1987, p. 213). Further records: soil from the Enns/Danube River floodplain, Upper Austria (Foissner 2004, p. 371); forest soil (Pruno-Fraxinetum; calcaric fluvisol; typical mull) from eastern Austria (Foissner et al. 2005, p. 632); blackish compost soil from a municipal compost plant of Munich, Bavaria, Germany (Foissner 2000a, p. 260); mud from deep, dry rock-pools (pH 5.8) from the escarpment of the Central Namib Desert, Spitzkoppe area about 120 km north-east of the city of Swakopmund and from dark, very humus soil from an alluvial grassland near the Bambatsi Guest Farm between the towns of Khorixas and Outjo, Namibia (Foissner et al. 2002, p. 63b). Feeds on bacteria (Berger & Foissner 1987). Biomass of 106 specimens about 24 mg (Foissner 1987, p. 124; 1998, p. 210).
Tetrastyla
463
Tetrastyla Schewiakoff, 1892 1892 Tetrastyla oblonga n. g. et sp. – Schewiakoff, Verh. naturk.-med. Ver. Heidelb., 4: 561 (original description; no formal diagnosis provided). Type species (by original designation): Tetrastyla oblonga Schewiakoff, 1892. 1909 Tetrastyla Schewiakoff, 1892 – Calkins, Protozoölogy, p. 55 (classification of Protozoa). 1979 Tetrastyla Schewiakoff, 1892 – Jankowski, Trudy zool. Inst., 86: 67 (catalogue of generic names of euplotids and hypotrichs). 2001 Tetrastyla Schewiakoff 1893 – Aescht, Denisia, 1: 160 (catalogue of generic names of ciliates). 2001 Tetrastyla Schewiakoff, 1892 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 89 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs).
Nomenclature: Tetrastyla is a composite of the Greek quantifier tetra (four) and the Greek noun ho stylos (style, cirrus in the present case) and alludes to the presence of four frontal and four transverse cirri (Schewiakoff 1893). According to Aescht (2001), Tetrastyla Schewiakoff, 1892 is a nomen nudum because published without description (ICZN 1999, Article 12.1). However, this is incorrect because Schewiakoff (1892) provided a rather detailed description. Corliss (1979, p. 208) mentioned Tetrastyla in his table with nomina oblita (= forgotten names). Characterisation (A = supposed apomorphy): Amphisiellidae(?) with continuous adoral zone of membranelles. Four frontal cirri (likely three frontal cirri and either one buccal cirrus or one parabuccal cirrus). Postperistomial cirrus lacking. Amphisiellid median cirral row almost of body length. Transverse cirri present. One right and one left marginal row. Remarks: See single species. Species included in Tetrastyla: (1) Tetrastyla oblonga Schewiakoff, 1892.
Single species
Tetrastyla oblonga Schewiakoff, 1892 (Fig. 95a–c) 1892 Tetrastyla oblonga n. g. et sp. – Schewiakoff, Verh. naturk.-med. Ver. Heidelb., 4: 561 (original description; no formal diagnosis provided and no type material available). 1893 Tetrastyla oblonga nov. gen. et sp. – Schewiakoff, Zap. imp. Akad. Nauk, 41: 66, Tafel IV, Fig. 57 (Fig. 95a; again described as new genus and species; first illustration). 1932 Amphisiella (Tetrastyla) oblonga Schewiakoff, 1893 – Kahl, Tierwelt Dtl., 25: 591, Fig. 106 28 (Fig. 95b; revision; combination with Holosticha, see nomenclature). 1960 Amphisiella (Tetrastyla) oblonga Schewiakoff – Dragesco, Trav. Stn biol. Roscoff, 122: 314 (combination with Amphisiella; guide to psammophilic ciliates). 1972 Tetrastyla oblonga Schewiakoff, 1892 – Borror, J. Protozool., 19: 19 (revision). 1974 Amphisiella oblonga Schewiakoff – Stiller, Fauna Hung., 115: 96, Fig. 58B (redrawing of Fig. 95a; revision of hypotrichs). 2001 Tetrastyla oblonga Schewiakoff, 1892 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 89 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs).
464
SYSTEMATIC SECTION
Nomenclature: No derivation of the name is given in the original description. The species-group name oblong·us, -a, -um (Latin adjective [m; f; n]; elongate) alludes to the elongate body outline. Type species of Tetrastyla. Kahl (1932) classified Amphisiella as subgenus of Holosticha. Thus, the correct name in his revision is Holosticha (Amphisiella) oblonga (Schewiakoff, 1892) Kahl, 1932. Consequently, Kahl (1932) is not the combining author for the classification in Amphisiella, as supposed by Carey (1992, p. 179), because subgenera are nomenclaturally irrelevant in this respect. The transfer to Amphisiella was made by Dragesco (1960). Remarks: Schewiakoff (1892, 1893) described this species from freshwater in New Zealand, a rather isolated area. The ventral row indicates, as suggested by Kahl (1932), a synonymy with Amphisiella. However, it is not recommendable to synonymise the limnetic Tetrastyla with Amphisiella, which contains – according to Kahl’s and my opinion – mainly/only marine species. Detailed studies on freshwater habitats in New Zealand have to be awaited before the legitimacy of Tetrastyla and T. oblonga can be discussed seriously. Consequently, the characterisation of Tetrastyla is not very specific. The four frontal cirri comprise possibly the ordinary three frontal cirri and a buccal cirrus or the frontal cirri and cirrus III/2. Of course, other patterns are also possible. Kahl (1932) supposed that the four transverse cirri are an abnormal pattern of the ordinary five transverse cirri. However, other species, which more or less invariably have four transverse cirri are known, for example, Cyrtohymena tetracirrata (for review, see Berger 1999, p. 317). Perhaps anlage II or anlage IV – which usually forms cirri arranged between the anterior portion of the amphisiellid median cirral row and cirrus III/2 – does not form a transverse cirrus. Dragesco (1960) found a similar species in the sand of Lake Geneva, France. Since he observed some deviations, he considered the European population preliminarily as mesopsammic form of T. oblonga. Corliss (1961, p. 169) considered, like Kahl (1932), Tetrastyla as synonym of Amphisiella. Borror (1972) classified it as species of questionable systematic position, and Carey (1992, p. 179) listed this limnetic species, likely by mistake, in his guide to marine interstitial ciliates. In addition, Carey (1992) incorrectly wrote that it has two ventral rows (Fig. 95c). Detailed redescription of T. oblonga is needed, preferably from a population from near the type locality, that is, limnetic habitats in New Zealand. Morphology: The following paragraphs are based only on the descriptions by Schewiakoff (1892, 1893). Body size 160 × 65 µm. Body outline elongate to oval, anterior portion indistinctly narrowed, posterior portion slightly widened, both ends rounded. Ventral side flat, dorsal side moderately vaulted. Body flexible. Two macronuclear nodules left of midline distinctly behind buccal vertex, connected by fine strand; individual nodules ovoid, caryoplasm finely meshed, with some distinct chromatin bodies. Micronucleus small, ellipsoidal, and homogenous, usually laterally attached to anterior
Tetrastyla
465 Fig. 95a–c Tetrastyla oblonga (a, from Schewiakoff 1893; b, after Schewiakoff 1893 from Kahl 1932; c, after Schewiakoff or Kahl from Carey 1992. From life). Ventral view of a representative specimen, 160 µm. Schewiakoff (1892, 1893) discovered this species in puddles in the jungle of New Zealand. Possibly this species has a local distribution and has therefore not yet been rediscovered. Note that in Carey’s (1992) illustration the number of cirri per row is distinctly higher than in the original drawing by Schewiakoff. In addition, he mentioned this limnetic species in a guide to marine ciliates. Page 463.
macronuclear nodule. Contractile vacuole, as is usual, about in mid-body and at left cell margin, vaults cortex distinctly at end of diastole, empties, as is usual, via dorsal side. Cytoplasm finely granulated, hyaline, colourless, “ectoplasm” forms distinct seam (Fig. 95a), possibly caused by cortical granules although the presence/absence of such organelles is not discussed by Schewiakoff (1892, 1893). Swims rapidly or creeps on algae. Adoral zone of membranelles occupies about 33% of body length. Buccal cavity of ordinary size, anterior margin distinctly curved leftwards. Paroral distinct. Four frontal cirri (see remarks for possible pattern). Three cirral rows, that is, one right and one left marginal row, each in ordinary position. Amphisiellid median cirral row extends from near frontal cirri to near transverse cirri; number of cirri (about 17) possibly underestimated because this is a rather low number for such a large species (details must not be over-interpreted). Four moderately thick transverse cirri, longer than frontal cirri. Dorsal bristles “short”, that is, likely around 3 µm. Number and arrangement of kineties and presence/absence of caudal cirri not known. Population studied by Dragesco (1960): body length 100 µm; five frontal cirri; three transverse cirri. Occurrence and ecology: Tetrastyla oblonga is very likely confined to limnetic habitats. The type locality is the jungle near the Waitakeri Falls (New Zealand), where Schewiakoff (1892) discovered T. oblonga in puddles. Dragesco (1960) found this species, or a very similar one, in the sand of Lake Geneva near the village of Excenevex. Record not substantiated by morphological data: river Maritsa, Bulgaria (Detcheva 1981, p. 23; 1983a, p. 204). The record from the litter of a forest soil in Louisiana by Bamforth (1968, p. 14) is likely a misidentification. Tetrastyla oblonga feeds on algae (Schewiakoff 1893).
466
SYSTEMATIC SECTION
Taxa not Considered in the Amphisiellidae The following taxa, included in the amphisiellids by some workers (Tables 4–14), lack a more or less distinct amphisiellid median cirral row originating from the two or three rightmost anlagen. They are therefore excluded from the amphisiellids here. Amphisiellides Foissner, 1988, Stapfia, 17: 120. Type species (by original designation): Uroleptoides atypica Hemberger, 1985. Remarks: Classified in the Amphisiellidae by Foissner (1988), Eigner & Foissner (1994), Petz & Foissner (1996), Shi (1999) and Shi et al. (1999), and Lynn & Small (2002). In the type species the frontoventral row is formed from a single anlage and a dorsal kinety fragmentation is very likely present. Thus, Amphisiellides is removed from the amphisiellids and classified in the oxytrichids (p. 651). Balladyna Kowalewskiego, 1882, Pam. fizyogr., 2: 408. Type species (by original designation and monotypy): Balladyna parvula Kowalewskiego, 1882. Remarks: Classified in the Amphisiellidae by Hemberger (1982; as incertae sedis) and Tuffrau & Fleury (1994). Balladyna parvula is a very small hypotrich with about seven frontoventral cirri, five prominent transverse cirri, and relatively long marginal cirri and dorsal bristles. The habitus closely resembles that of species of the Oxytricha setigera group, a relationship not discussed by Berger (1999). Since there is no evidence for an amphisiellid median cirral row, Balladyna parvula is not reviewed in the present book. Balladynella Stiller, 1974, Annls hist.-nat. Mus. natn. hung., 66: 129, 130. Remarks: Stiller (1974a) overlooked to fix a type species. Consequently, Balladynella Stiller, 1974 is not available (ICZN 1964, Article 13b). According to Aescht (2001, p. 30), now available as Balladynella Jankowski, 1979 (p. 51) with Balladyna fusiformis Kahl, 1932 as type species by original designation. Classified in the Amphisiellidae by Tuffrau & Fleury (1994). The habitus of the type species is more reminiscent of an oxytrichid, and since there is no evidence for an amphisiellid median cirral row, it is not reviewed in the present book. Banyulsella Dragesco, 1954, Vie Milieu, 4: 637. Type species (by monotypy): Banyulsella viridis Dragesco, 1954. Remarks: First mentioned, but not described (thus a nomen nudum) in a species list by Dragesco (1953, p. 629). Described in detail by Dragesco (1960, p. 316). Classified as incertae sedis in the Amphisiellidae by Hemberger (1982). The single species, Banyulsella viridis, is very small (about 50 µm), has a very large (more than 50% of body length), U-shaped adoral zone, three enlarged frontal cirri, two short and two long “fronto-ventral” cirral rows, and a long row of transverse(?) cirri. Marginal cirri obviously lacking (which would be an apomorphy). One row of fine cirri on rear portion of dorsal side. Six small macronuclear nodules. Mesopsammon of Banyuls-sur-Mer, Mediterranean Sea, France. The habitus
Taxa not Considered in the Amphisiellidae
467
is not reminiscent of an amphisiellid. Thus, it is not discussed in the present book. Jankowski (1975, p. 27) established the Banyulsellidae (incorrectly spelled Banylsellidae) for this species, which will be treated in a later volume of the monograph of hypotrichs. Circinella Foissner, 1994, Europ. J. Protistol., 30: 157. Type species (by original designation): Circinella arenicola Foissner, 1994. Remarks: Classified in the Amphisiellidae by Shi et al. (1999), Shi (1999), and Lynn & Small (2002). The ventral row of C. arenicola is formed from a single anlage. Thus, it is not included in the amphisiellids because the amphisiellid median cirral row originates from two or three anlagen. Circinella is reviewed in a later volume of the monograph of hypotrichs. Cladotricha Gaievskaia, 1925, Russk. Arkh. Protist., 4: 259, 281. Type species (by original designation): Cladotricha koltzowii Gaievskaia, 1925. Remarks: Classified in the Amphisiellidae by Hemberger (1982). According to Eigner (1997) it probably belongs to the Orthoamphisiellidae Eigner, 1997. It will be discussed in a later volume of the monographic series. Coniculostomum Njiné, 1979, Protistologica, 15: 353. Type species (by monotypy): Laurentia monilata Dragesco & Njiné, 1971. Remarks: Classified in the Amphisiellidae by Tuffrau & Fleury (1994). Because of the fragmenting dorsal kinety and the rigid body we assigned it to the Stylonychinae (Berger & Foissner 1997). For review, see Berger (1999, p. 606). Epiclintes Stein, 1863, Amtliche Berichte Deutscher Naturforscher und Ærzte in Karlsbad, 37: 162. Type species (by subsequent designation by Stein 1864, p. 444): Oxytricha auricularis Claparède & Lachmann, 1858. Remarks: Classified in the Amphisiellidae by Hemberger (1982). Epiclintes is a urostyloid which has a midventral complex composed of midventral rows only. For review, see Berger (2006, p. 1116). Eschaneustyla Stokes, 1886, Proc. Am. phil. Soc., 23: 28. Type species (by monotypy): Eschaneustyla brachytona Stokes, 1886. Remarks: Classified in the Amphisiellidae by Small & Lynn (1985). Eschaneustyla is a urostyloid likely closely related to Epiclintes (see previous entry) because the midventral complex is composed of midventral rows only. For review, see Berger (2006, p. 1146). Gastrostyla Engelmann, 1862, Z. wiss. Zool., 11: 383. Type species (by monotypy): Gastrostyla steinii Engelmann, 1862. Remarks: Classified in the Amphisiellidae by Eigner & Foissner (1994), Petz & Foissner (1996), and Lynn & Small (2002). Gastrostyla has a fragmenting dorsal kinety and was therefore assigned to the Oxytrichidae by Berger (1999, p. 789). Later, it was split into subgenera by Foissner et al. (2002, p. 720), and recently Foissner et al. (2004) showed – via a molecular analysis
468
SYSTEMATIC SECTION
– that the type species (G. steinii) belongs to the Stylonychinae. This position is confirmed by the rigid body and the lack of cortical granules (own observations and pers. comm. by W. Foissner). By contrast, Gastrostyla (Spetastyla) mystacea (Stein, 1859) Sterki, 1878 has a very flexible body and cortical granules, strongly indicating (de facto proving) that Gastrostyla (Gastrostyla) and Gastrostyla (Spetastyla) are not closely related. For taxonomic and nomenclatural consequences see Maregastrostyla (p. 136). Kahliella Corliss, 1960, J. Protozool., 7: 275. Type species (by original designation): Kahlia acrobates Horváth, 1932. Remarks: Classified in the Amphisiellidae by Hemberger (1982) and Small & Lynn (1985). Kahliella species lack an amphisiellid median cirral row (e.g., Fleury et al. 1985, Berger & Foissner 1987, Eigner 1995) and are therefore not reviewed in the present volume. Tuffrau (1979) established a new family for Kahliella and its relatives, which will be reviewed in a later volume of the monograph of hypotrichs. Lacazea Dragesco, 1960, Trav. Stn. biol. Roscoff, 122: 330. Type species (by original designation): Lacazea ovalis Dragesco, 1960. Remarks: Classified as incertae sedis in the Amphisiellidae by Hemberger (1982). Lacazea ovalis is almost circular in outline, has a single macronucleus (indicating that it is a euplotid), and a difficult-tointerpret cirral pattern, which is not reminiscent of an amphisiellid. It will be reviewed in a later volume of the monograph of hypotrichs. Onychodromopsis Stokes, 1887, Ann. Mag. nat. Hist., 20: 107. Type species (by monotypy): Onychodromopsis flexilis Stokes, 1887. Remarks: Classified in the Amphisiellidae by Small & Lynn (1985) and Tuffrau & Fleury (1994). Onychodromopsis is an oxytrichid reviewed by Berger (1999, p. 475). Note the problems with Allotricha (see also Petz & Foissner 1996). Onychodromus Stein, 1859, Lotos, 9: 4. Type species (by original designation): Onychodromus grandis Stein, 1859. Remarks: Classified in the Amphisiellidae by Small & Lynn (1985). Onychodromus is an oxytrichid (dorsal kinety fragmentation present) with an increased number of frontal-ventral-transverse cirri anlagen and a rigid body. For the latter reason assigned to the Stylonychinae by Berger (1999, p. 722). Later, we transferred the second species (O. quadricornutus Foissner, Schlegel & Prescott, 1987) to Styxophrya Foissner, Moon-van der Staay, van der Staay, Hackstein, Krautgartner & Berger, 2004 because of morphological and molecular differences to the type species (Foissner et al. 2004). Orthoamphisiella Eigner & Foissner, 1991, Acta Protozool., 30: 129. Type species (by original designation): Orthoamphisiella stramenticola Eigner & Foissner, 1991. Remarks: Classified in the Amphisiellidae by Shi et al. (1999), Shi (1999), and Lynn & Small (2002). The long ventral row is formed from a single anlage (Eigner & Foissner 1993), whereas the amphisiellid median cirral row originates from two or
Taxa not Considered in the Amphisiellidae
469
three anlagen. Thus, Orthoamphisiella is not included in the present monograph. Eigner (1997) established the Orthoamphisiellidae to include this and some other genera. Orthoamphisiella breviseries Foissner, Agatha & Berger, 2002 is closely related to Gonostomum according to molecular data (Foissner et al. 2004, Gong et al. 2006). Paragastrostyla Hemberger, 1985, Arch. Protistenk., 130: 407. Type species (by original designation): Paragastrostyla lanceolata Hemberger, 1985. Remarks: Classified in the Amphisiellidae by Eigner & Foissner (1994), Petz & Foissner (1996), Shi et al. (1999), Shi (1999), and Lynn & Small (2002). However, Paragastrostyla is very likely a urostyloid group closely related to Holostichides Foissner, 1987a. Thus, it was reviewed by Berger (2006, p. 613). Paraurostyla Borror, 1972, J. Protozool., 19: 9. Type species (by original designation): Urostyla weissei Stein, 1859. Remarks: Classified in the Amphisiellidae by Hemberger (1982), Small & Lynn (1985), and Tuffrau & Fleury (1994). Species of this group lack an amphisiellid median cirral row and – more important – have a fragmenting dorsal kinety 3. Thus, Paraurostyla belongs to the oxytrichids (for review, see Berger 1999, p. 841), a position already suggested by Borror (1979) and Wirnsberger et al. (1985). Later, this relationship was confirmed by molecular data (e.g., Bernhard et al. 2001, Hewitt et al. 2003). Parurosoma Gelei, 1954, Acta biol. hung., 5: 332. Type species (by monotypy): Holosticha (Parurosoma) dubium Gelei, 1954. Remarks: Classified in the Amphisiellidae by Tuffrau & Fleury (1994). The ventral cirral pattern is strongly reminiscent of 18-cirri oxytrichids. Thus, Berger & Foissner (1997) and Berger (1999, p. 491) assigned it to the oxytrichids. Periholosticha Hemberger, 1985, Arch. Protistenk., 130: 403. Type species (by original designation): Periholosticha lanceolata Hemberger, 1985. Remarks: Classified in the Amphisiellidae by Lynn & Small (2002). Periholosticha lanceolata feigns an amphisiellid median cirral row, which is, however, composed of several midventral cirral pairs. For review of this urostyloid, see Berger (2006, p. 498). Psammomitra Borror, 1972, J. Protozool., 19: 8, 15. Type species (by monotypy): Mitra radiosa Quennerstedt, 1867. Remarks: Classified in the Amphisiellidae by Tuffrau & Fleury (1994) and Lynn & Small (2002). Psammomitra retractilis (Claparède & Lachmann, 1858) Borror, 1972, the correct name of the type species, is a urostyloid reviewed in detail by Berger (2006, p. 221). Pseudouroleptus Hemberger, 1985, Arch. Protistenk., 130: 398. Type species (by original designation): Pseudouroleptus caudatus Hemberger, 1985. Remarks: Classified in the Amphisiellidae by Hemberger (1982), Eigner & Foissner (1994), Petz & Foissner (1996), and Lynn & Small (2002). Hemberger (1982, 1985) described and
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SYSTEMATIC SECTION
illustrated a dorsal kinety fragmentation strongly indicating that Pseudouroleptus belongs to the Oxytrichidae (p. 658). Territricha Berger & Foissner, 1988, Zool. Anz., 220: 127. Type species (by original designation): Territricha stramenticola Berger & Foissner, 1988. Remarks: Classified in the Amphisiellidae by Tuffrau & Fleury (1994, p. 141). When we described this species we assumed that it is closely related to the urostyloids because of the zigzagging cirral pattern. Thus, the classification in the amphisiellids is hardly comprehensible. However, the fragmenting dorsal kinety 3 assigns it unequivocally to the oxytrichids (for review, see Berger 1999, p. 884). Trachelochaeta Šrámek-Hušek, 1954, Arch. Protistenk., 100: 265. Type species (by original designation): Trachelochaeta bryophila Šrámek-Hušek, 1954. Remarks: Classified in the Amphisiellidae by Hemberger (1982) and Tuffrau & Fleury (1994). The cirral pattern and oral apparatus does not indicate an amphisiellid relationship. Thus, it is not reviewed in here. Trachelostyla Kahl, 1932, Tierwelt Dtl., 25: 596. Remarks: Classified in the Amphisiellidae by Tuffrau & Fleury (1994). Stichochaeta pediculiformis Cohn, 1866, type of the genus Trachelostyla Borror, 1972 (note that Trachelostyla Kahl, 1932 is invalid), has a modified 18-cirri pattern and a special type of oral apparatus indicating that it does not belong to the amphisiellids. It is the type of the Trachelostylidae reviewed in the present book (p. 471). Wallackia Foissner, 1976, Acta Protozool., 15: 387, 390. Type species (by original designation): Wallackia schiffmanni Foissner, 1976. Remarks: Classified in the Amphisiellidae by Tuffrau & Fleury (1994). Wallackia species lack an amphisiellid median cirral row (Foissner et al. 2002, p. 635ff) and are therefore not reviewed here. This genus will be reviewed in a later volume of the monograph of hypotrichs.
Trachelostylidae Small & Lynn, 1985 1985 Trachelostylidae n. fam. 1 – Small & Lynn, Phylum Ciliophora, p. 460 (original description; guide to ciliate genera; see remarks). Nominotypical genus: Trachelostyla Borror, 1972. 2001 Trachelostylidae Small and Lynn, 1985 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 113 (nomenclator containing all basionyms, combinations, and higher taxa of euplotids and hypotrichs). 2002 Trachelostylidae Small & Lynn, 1985 2 – Lynn & Small, Phylum Ciliophora, p. 457 (guide to ciliate genera; see remarks).
Nomenclature: Small & Lynn (1985) established the family Trachelostylidae containing Trachelostyla, Psammomitra, Urosoma, and Urosomoida. Simultaneously, they established the Gonostomatidae (Small & Lynn 1985, p. 455), however, without including the type species of the type genus, Gonostomum affine, which they assigned to Trachelostyla (for details about this rather complex situation, see remarks at Trachelostyla). If Gonostomum Sterki, 1878 and Trachelostyla Borror, 1972 have to be unified in a higher taxon, it is recommended to use the Gonostomatidae simply because this group is based on the older genus. Characterisation (A = supposed apomorphy): Primarily 18-cirri hypotrichs with body distinctly cephalised (A), that is, anterior body portion narrowed head-like and thus set off from body proper. Adoral zone along anterior and left margin of head, undulating membranes straight and relatively short. Three “postoral” ventral cirri right of adoral zone of membranelles. One right and one left marginal row. Caudal cirri present. Dorsal kinety formation in Trachelostyla pattern (A). Dorsomarginal kineties lacking. Marine and inland saltwater. The ground pattern of the Trachelostylidae: In the paragraphs below the supposed ground pattern3 of the Trachelostylidae is discussed. Apomorphy of the Trachelostylidae. It is difficult to estimate which features are apomorphies for the trachelostylids because the nearest relative is uncertain. The phylogenetic analyses based on molecular data are rather different. According to Gong et al. (2006) and Shao et al. (2007), Trachelostyla is closely related to Gonostomum, according to Schmidt et al. (2007) it is not. The Chinese proposal is supported by agreements in the shape of the adoral zone and the anteriorly displaced postoral ventral cirri (for review of Gonostomum, see Berger 1999, p. 367). By contrast, the very different dorsal kinety formation in Gonostomum (each of the three kineties divides individually; Hemberger 1982, Song 1990) and Trachelostyla (multiple fragmentation of kinety 1, two kineties formed by kinety 6; Shao et al. 2007; details see Addenda) support the distinct separation found by Schmidt et al. (2007). In
1
Small & Lynn (1985) provided the following diagnosis: Frontoventral cirri scattered on forward part, sometimes on rear part, never on mid area of ventrum. 2 Lynn & Small (2002) provided the following characterisation: Frontoventral cirri scattered on anterior peristomal region, sometimes in posterior, never on mid-area of ventral surface; kinetosomes for stomatogenesis arise de novo on ventral surface. 3 For a brief explanation of the term ground pattern see chapter 2.2 of the general section.
471
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SYSTEMATIC SECTION
all trees both Trachelostyla and Gonostomum branch off near the base of the Hypotricha, likely because they lack dorsomarginal kineties. Body distinctly cephalised. This feature is rather conspicuous and easily recognisable in life. The outgroup comparison shows that this body outline has to be interpreted as apomorphy of the trachelostylids. However, other hypotrichs with a narrowed anterior end are known, for example, Hemigastrostyla Song & Wilbert, 1997 (for review, see Berger 1999, p. 933), Spiroamphisiella (p. 148), Cossothigma (p. 382), Biholosticha discocephalus (Kahl, 1932) Berger, 2003 (for review, see Berger 2006, p. 1178), and Stichotricha Perty, 1849 (for review, see Kahl 1932, p. 556; Foissner et al. 1991). Of these taxa, only Hemigastrostyla enigmatica (Dragesco & Dragesco-Kernéis, 1986) Song & Wilbert, 1997 has a cirral pattern which is reminiscent of that of Trachelostyla because 18 frontal-ventral-transverse cirri are present and the postoral ventral cirri are somewhat displaced anteriad. However, Hemigastrostyla enigmatica has fragmenting dorsal kineties and a (non-migrating) dorsomarginal row indicating that it belongs to the oxytrichids (Song & Hu 1999). By contrast, Gong et al. (2007) found a close relationship of H. enigmatica and Amphisiella annulata (details see Addenda). For the type species, Hemigastrostyla stenocephala (Borror, 1963) Song & Wilbert, 1997, the dorsal morphogenesis is not known (for review, see Berger 1999, p. 822) and H. szaboi Wilbert & Song, 2005 has only three bipolar dorsal kineties, each forming a caudal cirrus, that is, lacks a kinety fragmentation. These high variability indicates that Hemigastrostyla is nonmonophyletic. Dorsal kinety formation in Trachelostyla-pattern. Details see Addenda. Plesiomorphies of the Trachelostylidae. Most plesiomorphies of the Trachelostylidae are basically the same as for the Hypotricha. Thus, the reader is referred to chapter 2.2.2 of the general section where the ground pattern of the Hypotricha is discussed in detail. Here, only three important plesiomorphies are briefly discussed. Adoral zone of membranelles and undulating membranes in Gonostomumpattern. The oral apparatus of Gonostomum and Trachelostyla is rather similar because the adoral zone runs mainly along the left body margin and the undulating membranes are rather short and straight. The question is whether or not this type of oral apparatus evolved once or two (or more) times independently. Wallackia Foissner, 1976 and Kahliella Corliss, 1960 also have a “Gonostomum-like” oral apparatus; however, the dorsal infraciliature of Kahliella on the one hand and Gonostomum and Trachelostyla on the other is rather different because Kahliella has a dorsomarginal kinety, a feature lacking in the other two genera. Consequently, I suppose that Kahliella belongs to the non-oxytrichid Dorsomarginalia, whereas Trachelostyla and Gonostomum very likely branch off outside the Dorsomarginalia. This implies that the gonostomoid oral apparatus evolved twice. In spite of the similarities between Gonostomum and Trachelostyla I do not include Gonostomum in the Trachelostylidae because it lacks a head and does not occur in the sea. In addition, Gonostomum and Trachelostyla form the dorsal kineties rather different (see Addenda). Possibly, the Trachelostylidae and the Gonostomatidae – also established by Small & Lynn
Trachelostylidae
473
(1985) and comprising Gonostomum, Paragonostomum Foissner, Agatha & Berger, 2002, and perhaps even Wallackia – are sister groups. Postoral ventral cirri displaced anteriad right of proximal portion of adoral zone. The postoral ventral cirri (cirri IV/2, V/3, V/4) of the 18-cirri hypotrichs are usually arranged behind the buccal vertex. In Gonostomum, Hemigastrostyla enigmatica, and Trachelostyla this cirral group is displaced anteriad, that is, they are right of the proximal portion of the adoral zone of membranelles. At least two explanations are possible: (i) the postoral ventral cirri do not migrate so far posteriorly as in ordinary 18-cirri hypotrichs during cell division; or (ii) the postoral ventral cirri are basically at their ordinary position, but due to the rather long adoral zone (40–50% of body length) they are not behind the buccal vertex, but right of the rear portion of the adoral zone. In addition to the special shape of the adoral zone, the undulating membranes are comparatively short and straight both in Gonostomum and the trachelostylids. Anteriorly displaced postoral ventral cirri are very likely also present in other 18-cirri hypotrichs, for example, Hemisincirra inquieta (Fig. 79l) and Lamtostyla granulifera (Fig. 40n). Remarks: Small & Lynn (1985) placed the trachelostylids in the Sporadotrichina, together with the Oxytrichidae. Besides the type genus, Psammomitra, Urosoma, and Urosomoida have been included. According to the characterisation by Lynn & Small (2002), seven genera are included in the trachelostylids. However, in the key and main text they listed only five. I include only two marine genera in the trachelostylids, namely, Trachelostyla and Spirotrachelostyla. For a foundation of the exclusion of most genera from the trachelostylids, see Taxa not Considered chapter. As discussed above, gonostomatids are possibly the sister group of the trachelostylids. The classification of Gonostomum in the oxytrichids by Berger & Foissner (1997) and Berger (1999, p. 367) is based on the presence of the 18-cirri pattern and a different interpretation of the dorsal kinety pattern. Earlier I assumed that the 18cirri pattern is an apomorphy of the oxytrichids and the simple dorsal kinety pattern of Gonostomum evolved from the complicated Oxytricha-pattern. Genera included in Trachelostylidae: Trachelostyla Borror, 1972 (type); Spirotrachelostyla Gong, Song, Li, Shao & Chen, 2002.
Key to the genera of the Trachelostylidae See also Spiroamphisiella (p. 148) and Hemigastrostyla (Berger 1999, p. 822, 933) if you cannot identify your population with this and the following keys. 1 Body spirally twisted and spindle-shaped. . . . . . . . . . Spirotrachelostyla (p. 502) - Body not twisted. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trachelostyla (p. 474)
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SYSTEMATIC SECTION
Trachelostyla Borror, 1972 1932 Trachelostyla gen. n. – Kahl, Tierwelt Dtl., 25: 596 (nomen nudum; see nomenclature). 1972 Trachelostyla Kahl, 19321 – Borror, J. Protozool., 19: 15 (original description). Type species (by original designation 2): Stichochaeta pediculiformis Cohn, 1866 (see nomenclature). 1979 Trachelostyla Kahl, 1932 – Jankowski, Trudy zool. Inst., 86: 67 (catalogue of generic names of hypotrichs). 1979 Trachelostyla Kahl, 1932 – Tuffrau, Trans. Am. microsc. Soc., 98: 526 (generic classification of hypotrichs). 1979 Trachelostyla Kahl, 1932 – Corliss, Ciliated Protozoa, p. 310 (revision; see nomenclature). 1982 Trachelostyla Kahl, 1932 3 – Hemberger, Dissertation, p. 261 (revision of hypotrichs). 1984 Trachelostyla Kahl, 1932 – Maeda & Carey, Bull. Br. Mus. nat. Hist. (Zool.), 47: 5 (revision, see remarks). 1985 Trachelostyla – Small & Lynn, Phylum Ciliophora, p. 461 (guide to ciliate genera; see remarks). 1987 Trachelostyla Kahl, 1932 – Tuffrau, Annls Sci. nat. (Zool.), 8: 115 (generic classification of hypotrichs). 1992 Trachelostyla (Kahl, 1935–5) Maeda and Carey, 1984 – Carey, Marine interstitial ciliates, p. 185 (guide). 1994 Trachelostyla Kahl 1932 – Tuffrau & Fleury, Traite de Zoologie, 2: 141 (revision of hypotrichs). 1999 Trachelostyla Borror, 1972 – Berger, Monographiae biol., 78: 894 (brief note; see nomenclature and remarks). 1999 Trachelostyla Kahl, 1932 – Shi, Song & Shi, Progress in Protozoology, p. 130 (generic revision of hypotrichs). 2000 Trachelostyla Kahl, 1932 – Shi, Acta Zootax. sinica, 25: 17 (generic revision of hypotrichs). 2001 Trachelostyla Borror 1972 – Aescht, Denisia, 1: 165 (catalogue of generic names of ciliates). 2001 Trachelostyla Borror, 1972 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 89 (nomenclator containing all basionyms, combinations, and higher taxa of euplotids and hypotrichs). 2002 Trachelostyla Kahl, 1932 4 – Hu & Song, Hydrobiologia, 481: 174 (improved diagnosis). 2002 Trachelostyla Kahl, 1932 – Lynn & Small, Phylum Ciliophora, p. 458 (guide to ciliate genera; see remarks). 2003 Trachelostyla Kahl, 1932 – Hu, Gong & Song, Pathogenic protozoa, p. 173 (guide to pathogenic ciliates). 2006 Trachelostyla Kahl, 1932 – Berger, Monographiae biol., 85: 1213 (brief note). 2006 Trachelostyla Borror, 19725 – Gong, Song, Li, Shao & Chen, Europ. J. Protistol., 42: 72 (redefinition and redescription of type species). 1
Borror (1972) provided the following diagnosis: One row each of right and left marginal cirri, not confluent posteriorly. No postbuccal frontoventral cirri except 2 just anterior of transverse cirri. Body narrowed, elongate, fragile. 2 Borror (1972) considered his type fixation as subsequent designation because he thought that Kahl’s original description was valid. 3 Hemberger (1982) provided the following diagnosis: Je 1 rechte und linke Marginalreihe, caudal getrennt; keine Ventralcirren, außer 2 unscheinbaren vor den Transversalcirren; 5 Transversalcirren; lange Dorsalcilien; während der Morphogenese wird auch die Proter-AMZ neu gebildet; Makronuklei zahlreich. 4 Hu & Song (2002) provided the following improved diagnosis: Body shape spindle-like, usually flexible and more or less twisted spirally, frontal area considerably narrowed and neck-like; frontoventral cirri well differentiated and mostly located in frontal area; typically with 5 transverse cirri, caudal cirri present. One marginal cirral row on each side. Gelatinous lorica commonly present. Usually found in marine water. 5 Gong et al. (2006) provided the following redefined diagnosis: Dorsoventrally flattened trachelostylids, body non-spirally twisted and elongate, with peristomal region conspicuously narrowed; ciliature generally in 11:2:5:3 pattern, i.e. 11 cirri in frontal region; two ventral cirri located anterior to five transverse cirri; three caudal cirri present. One left and one right row of marginal cirri not confluent posteriorly.
Trachelostyla
475
Nomenclature: No derivation of the name is given in the papers by Kahl (1932) and Borror (1972). Trachelostyla is a composite of the Greek nouns ho tráchelos (neck) and ho stýlos (handle, stalk, style; likely cirrus in present case). Possibly it refers to the fact that this hypotrich (indicated by the part -styla = cirrus) has a neck-like narrowed anterior body portion. Feminine gender (Aescht 2001, p. 303). The complicated nomenclature of Trachelostyla was already explained by Berger (1999). Kahl (1932) established this genus with two species, namely Stichochaeta pediculiformis Cohn, 1866 and “Trachelostyla caudata spec. n.”. Since he did not determine any as type species, the taxon is invalid, that is, not available according to Article 13(b) of the ICZN (1964; see also Article 13.3 of the ICZN 1999). This fact was overlooked by Corliss (1979) and later authors (e.g., Maeda & Carey 1984) who incorrectly assigned this genus to Kahl (1932). Since Kahl’s Trachelostyla is a nomen nudum, the diagnosis proposed by Kahl (1932) is not mentioned in a footnote. Berger (1999) considered Borror (1972) as the author of Trachelostyla – although Borror also did not realise that Kahl’s taxon was invalid – because he fixed the type species. This proposal was followed by Aescht (2001) and Gong et al. (2006). Since Trachelostyla Kahl, 1932 is not available, Trachelostyla Borror, 1972 is not a junior homonym. Incorrect subsequent spellings: Trachelostya (Gong et al. 2006, p. 71); Traxhelostyla pediculiformis (Czapik 1952, p. 4). Characterisation (A = supposed apomorphy): Trachelostylidae with non-twisted body. Additional characters: Only the type species is described in detail. Thus the list of additional features is short and uncertain. Body flexible; cortical granules likely lacking; dorsal bristles likely long (>5 µm). Remarks: See nomenclature before reading the remarks. I did not consider Trachelostyla in the monograph of the oxytrichids (Berger 1999) because I thought that neither the ventral nor the dorsal ciliature of this marine taxon is known in detail. Obviously I overlooked that Borror (1963, 1972), and even already Maupas (1883), illustrated the ventral cirral pattern in some detail (Fig. 97c, 98c, e), clearly showing that Trachelostyla is rather similar to Gonostomum because of the oral apparatus and the anteriorly displaced postoral ventral cirri. This pattern was confirmed recently by Xu & Song (1999; for Trachelostyla sp.; Fig. 99d, e) and by Gong et al. (2006) for the type species (Fig. 96d, e). The nucleotide sequence data provided by the latter workers also show that Trachelostyla is closely related to Gonostomum Sterki, 1878, a genus classified in the oxytrichids by Berger (1999, p. 367). An assignment of Trachelostyla to the oxytrichids was already proposed by Borror (1972) and even by Kahl (1932). However, Kahl’s assignment is worthless because he classified all genera in the Oxytrichidae, except for the euplotids and the aspidiscids. Borror (1972) included eight species in Trachelostyla, namely, the three species included in the present review (see next chapter), and (i) T. bryonicolum (Gellért, 1956b) Borror, 1972; (ii) T. ciliophorum (Gellért, 1956b) Borror, 1972; (iii) T. geleii (Gellért, 1957) Borror, 1972; (iv) T. macrostoma (Gellért, 1957) Borror, 1972; and (v) T. spirotrichoides (Gellért, 1956b) Borror, 1972. However, these five species de-
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SYSTEMATIC SECTION
scribed by Gellért are (very likely) synonyms of Gonostomum affine (Stein, 1859) Sterki, 1878, type of Gonostomum (for review, see Berger 1999, p. 369). Gonostomum affine was assigned to Gastrostyla Engelmann, 1862 by Borror (1972, p. 14), a classification which is certainly incorrect because of several morphological differences, for example, dorsal kinety fragmentation lacking in Gonostomum against present in Gastrostyla (for review, see Berger 1999, p. 789). This difference was later confirmed by molecular data (Foissner et al. 2004, Agatha et al. 2005, Gong et al. 2006). Buitkamp (1977a, p. 125) provided a brief revision of Trachelostyla and considered T. pediculiformis and T. caudata as valid species. Further, he transferred Gonostomum affine, type of Gonostomum, to Trachelostyla and therefore synonymised the older Gonostomum Sterki, 1878 with the younger Trachelostyla Borror, 1972 (or likely Trachelostyla Kahl, 1932 in Buitkamp’s opinion), that is, he ignored the principle of priority (ICZN 1964, Article 23e (i)). According to Corliss (1979), Tuffrau (1979), and Curds et al. (1983, p. 417), Stichochaeta Clàparede & Lachmann, 1858 is a synonym of Trachelostyla Kahl, 1932. Apart from the fact that they also ignored the principle of priority – Stichochaeta is older than Trachelostyla! – and overlooked the invalidity of Trachelostyla Kahl, 1932 (see nomenclature), this statement is incorrect because Stichochaeta is undoubtedly a junior synonym of Stichotricha Perty, 1849 (Stein 1859, p. 176; Bütschli 1889, p. 1743; Kahl 1932, p. 557, 559, 596; Borror 1972, p. 12; Foissner et al. 1991, p. 210). Corliss (1977, p. 137; 1979) and Tuffrau (1979) classified Trachelostyla in the Holostichidae; however, since a midventral complex, the sole morphological apomorphy of the urostyloids, is lacking it was not considered by Berger (2006) in the monograph on this group. Tuffrau & Fleury (1994) assigned it to the Amphisiellidae (Table 8), a classification which is very likely also incorrect because a distinct amphisiellid median cirral row is lacking. Recent data by Gong et al. (2007), however, suggest a close relationship of Amphisiella and Trachelostyla (details see Addenda). Maeda & Carey (1984) revised Gonostomum Sterki, 1878 and Trachelostyla Kahl, 1932, explaining in some detail the history of both taxa. Unfortunately, they did not mention the type species and therefore did not recognise that the establishment of Trachelostyla by Kahl (1932) was invalid because he did not fix a type species (see nomenclature). They accepted T. pediculiformis and T. caudata, but did not consider T. rostrata in detail because it is poorly described. Small & Lynn (1985) and Lynn & Small (2002) mixed two genera because they give Trachelostyla affine (= Gonostomum affine in Berger 1999) and T. pediculiformis1 as examples of Trachelostyla (possibly their classification is based mainly on Buitkamp’s suggestion). As already mentioned above, Gonostomum affine is the type species of Gonostomum, a valid, older(!) genus whose species occur predominantly in soil, rarely in freshwater. By contrast, Trachelostyla species are marine and 1
Note that the Trachelostyla pediculiformis illustrated by Small & Lynn (1985) and Lynn & Small (2002) is very likely T. rostrata (see Fig. 102b).
Trachelostyla
477
have a characteristically narrowed anterior body portion justifying the recognition of both Gonostomum and Trachelostyla, although morphological (shape of adoral zone; short undulating membranes; postoral ventral cirri displaced anteriad), ontogenetic (see type species), and molecular data (Gong et al. 2006) suggest a close relationship. However, the dorsal kinety formation of Gonostomum and Trachelostyla proceeds rather differently (Shao et al. 2007; details see Addenda). Kent (1882, p. 775) considered Stichochaeta pediculiformis as type of Stichochaeta Claparède & Lachmann, 1858. However, his assumption is irrelevant because the type species of Stichochaeta is S. cornuta Claparède & Lachmann, 1858 by monotypy (see Fromentel 1874, p. 161; Jankowski 1979, p. 65; Berger 2001, p. 80). Likely par lapsus, Aescht (2001, p. 154) accepted Kent’s incorrect opinion. Further, Stichochaeta was correctly synonymised with Stichotricha Perty, 1849 by Stein (1859; see also Foissner et al. 1991, p. 210). Cell division data on Trachelostyla are sparse (Kool 1970, Gong et al. 2006). The formation of the 18-cirri pattern proceeds as in other hypotrichs which have this plesiomorphic pattern, that is, from six (I–VI) anlagen. The formation of a new adoral zone for the proter is characteristic for the urostyloid taxon Pseudokeronopsidae Borror & Wicklow, 1983 (for review, see Berger 2006, p. 832). However, since a midventral complex is lacking, a classification of Trachelostyla in the urostyloids, as suggested, for example, by Corliss (1979), was avoided (Berger 2006, p. 1213). Thus, the formation of a new adoral zone for the proter has to be considered as convergence. Just recently, Shao et al. (2007) studied the cell division of T. pediculiformis in detail (see Addenda). The molecular data indicate different positions of Trachelostyla. According to Gong et al. (2006) it is closely related to Gonostomum and clusters in a group also containing an oxytrichid. According to Fig. 2 in Schmidt et al. (2007), it is the first species branching off within the Hypotricha; this non-dorsomarginalian position agrees with my estimation based on the dorsal ciliature (see also Addenda). Trachelostyla species mainly (exclusively?) occur in marine habitats and inland salt waters (e.g., Borror 1973, p. 50). Records of Trachelostyla from terrestrial habitats, for example, by Biczók (1956, p. 139; as Trachelostyla sp.) or limnetic habitats (e.g., Grispini 1938, p. 153) are thus very likely misidentifications. Some marine populations have not been identified to species level (e.g., Wiackowski 1981, p. 220; Sich 1990, p. 35; Santangelo & Lucchesi 1995, p. 51; Yang et al. 2005, p. 23). In the review on the oxytrichids I supposed that the terrestrial Oxytricha (Opisthotricha) elongata Grandori & Grandori, 1934 very likely belongs to Trachelostyla or Trachelochaeta (Berger 1999, p. 242). However, both proposals are likely incorrect because Trachelostyla is marine and has a narrowed anterior body portion (vs. terrestrial and not narrowed) and Trachelochaeta Šrámek-Hušek, 1954 is limnetic and has four cirral rows (vs. salt water and two cirral rows). Basically, Oxytricha elongata is reminiscent of Gonostomum. It will be briefly treated in a supplement to Gonostomum in a later monograph.
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SYSTEMATIC SECTION
Species included in Trachelostyla (alphabetically arranged basionyms are given): (1) Stichochaeta pediculiformis Cohn, 1866; (2) Trachelostyla caudata Kahl, 1932; (3) Trachelostyla rostrata Lepsi, 1962. Species misplaced in Trachelostyla: The following species – originally assigned to Trachelostyla – are now classified in other (related?) genera. Note that several other species (mainly synonyms of Gonostomum affine) have been transferred to Trachelostyla (see Berger 1999, 2001). Trachelostyla canadiensis Buitkamp & Wilbert, 1974. Remarks: A junior synonym of Gonostomum affine (Stein, 1859) Sterki, 1878. For review, see Berger (1999, p. 369). Trachelostyla dubia Dragesco, 1954. Remarks: Preliminary classified in Cossothigma Jankowski, 1978 (p. 383). Trachelostyla spiralis Dragesco & Dragesco-Kernéis, 1986. Remarks: Now classified in Spirotrachelostyla (p. 503). Trachelostyla tani Hu & Song, 2002. Remarks: Now classified in Spirotrachelostyla (p. 506).
Key to Trachelostyla species If you know that your specimen/population belongs to Trachelostyla, then the determination is relatively simple because body shape and nuclear apparatus are key features. However, note that similar forms exist, namely Spirotrachelostyla (body twisted; p. 502), Cossothigma (three cirral rows; p. 382), and Spiroamphisiella (body twisted, amphisiellid median cirral row; p. 148). 1 Posterior body portion narrowed tail-like (Fig. 101a–d). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trachelostyla caudata (p. 494) - Posterior body portion broadly rounded (Fig. 96a, j, 97a, c, l, 102a, b). . . . . . . . 2 2 Two macronuclear nodules (Fig. 102a, b). . . . . . . Trachelostyla rostrata (p. 498) - Many (9–64; neotype population with 9–17) macronuclear nodules (Fig. 96a, e, 97c, l). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trachelostyla pediculiformis (p. 478)
Trachelostyla pediculiformis (Cohn, 1866) Borror, 1972 (Fig. 96a–j, 97a–o, 98a–k, 99a–e, 100a, Table 30, Addenda) 1866 Stichochaeta pediculiformis n. sp.1 – Cohn, Z. wiss. Zool., 16: 285, 299, Tafel XV, Fig. 38a, b (Fig. 97a, b; original description; no type material available). 1
Cohn (1866, p. 299) provided the following diagnosis: Körper silbergrau, von dunklen Körperchen dicht erfüllt, flexil, nicht retractil, vom Rücken zusammengedrückt, linear, oblong, hinten abgerundet, vorn mit einem kürzeren und schmäleren, scharf abgesetzten Rüssel, welcher an der Spitze eine Anzahl (etwa sechs) langer wirbelnder Borsten, längs der Bauchseite einen Saum dichter, kurzer Wimpern, sowie eine Anzahl getrennter, langer (Spring-) Borsten trägt, während der eigentliche Körper auf der Bauchseite
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1882 Stichochœta pediculiformis, Cohn – Kent, Manual Infusoria II, p. 775, Plate XLIV, Fig. 13, 14 (redrawings of Fig. 97a, b; revision). 1883 Gonostomum pediculiforme – Maupas, Archs Zool. exp. gén., 1: 550, Planche XXIV, fig. 8–13 (Fig. 97c–h; redescription and combination with Gonostomum). 1888 Gonostomum pediculiforme – Gruber, Ber. naturf. Ges. Freiburg i. B., 3: 63, Fig. 11 (description of nuclear apparatus; see remarks). 1888 Stichochœta corsica nov. spec. – Gourret & Roeser, Archs Biol., 8: 187, Planche XIV, Fig. 6 (Fig. 97i; original description of synonym; no formal diagnosis provided and no type material available). 1889 Gonostomum pediculiformis – Bütschli, Protozoa, p. 1748 (generic revision). 1928 Gonostomum pediculiforme Cohn – Kahl, Arch. Hydrobiol., 19: 210, Abb. 44h (Fig. 97k; description of German population). 1929 Gonostomum pediculiforme Cohn – Hamburger & Buddenbrock, Nord. Plankt., 7: 93, Fig. 115 (redrawing of Fig. 97c; guide to planktonic ciliates). 1932 Trachelostyla (Stichochaeta) pediculiformis (Cohn, 1866) – Kahl, Tierwelt Dtl., 25: 596, Fig. 1065, 6, 11 (Fig. 97j, l, m; revision; see nomenclature). 1932 Gonostomum pediculiforme (Cohn) Maupas 1883 – Wang & Nie, Contr. biol. Lab. Sci. Soc. China, 8: 359, Fig. 67 (Fig. 97n; description of Chinese population). 1933 Trachelostyla pediculiformis (Cohn 1866) – Kahl, Tierwelt N.- u. Ostsee, 23: 112, Fig. 17 25 (Fig. 97o; guide to marine ciliates). 1936 Trachelostyla (Stichochaeta) pediculiformis (Cohn 1866) – Kiesselbach, Thalassia, 2: 20, 53, Abb. 46 (Fig. 98a; brief redescription). 1963 Trachelostyla pediculiformis (Cohn) 1866 – Biernacka, Polskie Archwm Hydrobiol., 11: 51, Abb. 100 (Fig. 98b; description of Polish population). 1963 Trachelostyla pediculiformis (Cohn, 1866) – Borror, Arch. Protistenk., 106: 511, Fig. 116 (Fig. 98c; description of American population; first silver impregnation?; likely no permanent slides available). 1970 Trachelostyla pediculiformis (Cohn, 1866) – Kattar, Zoologia e biologia marinha, 27: 188, Fig. 33 (Fig. 98d; description of Brazilian population). 1972 Trachelostyla pediculiformis (Cohn, 1866) Kahl, 1932 – Borror, J. Protozool., 19: 15, Fig. 43 (Fig. 98e; combination, with Trachelostyla, see nomenclature; generic revision of euplotids and hypotrichs). 1973 Trachelostyla pediculiformis – Hartwig, Mikrokosmos, 62: 330, Bild 1C (micrograph). 1974 Trachelostyla pediculiformis (Cohn, 1866) Kahl, 1932 – Jones, Univ. South Alabama Monogr., 1: 42, Plate XXIX, Fig. 6 (Fig. 98f; illustrated record). 1982 Trachelostyla pediculiformis (Cohn, 1866) Kahl, 1932 – Hemberger, Dissertation, p. 261 (revision of hypotrichs). 1984 Trachelostyla pediculiformis (Cohn, 1866) Kahl, 1932 – Maeda & Carey, Bull. Br. Mus. nat. Hist. (Zool.), 47: 5, Fig. 1 (Fig. 98g; redescription and revision). 1984 Trachelostyla pediculiformis (Cohn) – Burkovsky, Ecology of free-living ciliates, p. 164, Fig. 57 6 (Fig. 98h; ecology of ciliates). 1985 Trachelostyla pediculiformis (Cohn, 1866) – Aladro Lubel, An. Inst. Biol. Univ. Méx., 55: 27, Lámina 13, Fig. 2 (Fig. 98i; description of Mexican population). 1990 Trachelostyla pediculiformis (Cohn, 1866) – Aladro Lubel, Martínez Murillo & Mayén Estrada, Manual de ciliados, p. 135, figure on p. 135 (Fig. 98j; review). 1992 Trachelostyla pediculiformis (Kahl, 1930–5) Maeda and Carey, 1984 – Carey, Marine interstitial ciliates, p. 186, Fig. 737 (Fig. 98k, a redrawing of Fig. 98g; guide; incorrect author and incorrect combining authors). 1999 Trachelostyla pediculiformis (Cohn, 1866) Kahl, 1932 – Petz, Ciliophora, p. 299, Fig. 8.68 (Fig. 98c, e, g; review of South Atlantic plankton ciliates). mit drei Reihen kürzerer Wimpern besetzt ist. Mundöffnung an der Basis des Rüssels, mit einigen längeren, herausragenden präoralen Borsten. Am hintern Ende längere Schwanz- und Afterborsten. Bewegung abwechselnd kriechend und rückwärts springend.
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1999 Trachelostyla sp. – Xu & Song, Progress in Protozoology, p. 258, Fig. 34A–E (Fig. 99a–e; description of Chinese population; see remarks). 2001 Trachelostyla pediculiformis (Cohn, 1866) Borror, 1972 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 80 (nomenclator containing all basionyms, combinations, and higher taxa of euplotids and hypotrichs). 2006 Trachelostyla pediculiformis (Cohn, 1866) Borror, 1972 – Gong, Song, Li, Shao & Chen, Europ. J. Protistol., 42: 64, Fig. 1–29, Table 1 (Fig. 96a–j; redescription, neotypification, and analysis of small subunit ribosomal RNA gene sequence; GenBank accession number DQ057346; the neotype slide [registration number 2005:3:24:15] is deposited in the Natural History Museum in London, UK).
Nomenclature: No derivation of the species-group name is given in the original description. The name pediculiform·is -is -e (m, f, n) is obviously a composite of the Latin substantive pedicul·us (small louse), the thematic vowel ·i-, and the Latin formis (a species looking like another species). I do not known to which feature the name refers. Type species of Trachelostyla Borror, 1972. Note that Kent (1882) and Aescht (2001) incorrectly considered the present species as type of Stichochaeta (see remarks in genus section). The species-group name corsica refers to the type locality (Corsica, France) where the junior synonym was discovered by Gourret & Roeser (1888). Kahl (1932) is not the author of Trachelostyla (see nomenclature in genus section). Thus, he cannot be the combining author for the present species. This act was done, although not intentionally, by Borror (1972), because he is the author of Trachelostyla (see genus section for details). The misleading spelling “Trachelostyla (Stichochaeta) pediculiformis” in Kahl (1932) does not mean that he considered Stichochaeta as subgenus of Trachelostyla, but should only indicate that this species was originally assigned to Stichochaeta. Incorrect subsequent spellings: Trachelostyla pediculariformis (Madoni 2006, p. 165, 169); Trachelostyla pediculiforme (Mazei et al. 2002, p. 389); Trachelostyla pendiculiformis (Hu & Song 2002, p. 178). Remarks: Cohn (1866) made his observations on populations from an aquarium. Unfortunately, this bottle was charged with marine material both from the island of Heligoland (North Sea, Germany) and from Dorsetshire (now Dorset; Atlantic Ocean; Southwest England) so that the type locality of the new species described in Cohn’s paper is unknown. The lack of the type locality and differences in some morphological features in the previous descriptions caused Gong et al. (2006) to neotypify T. pediculiformis. Cohn (1866) provided a rather detailed description showing only some more or less distinct differences to later descriptions. Unfortunately, he could not see the nuclear apparatus. Two bright areas in the first and second third of the body were interpreted as contractile vacuoles (although no contraction was observed) which is likely a misobservation because nobody else ever saw one or two contractile vacuoles unequivocally in this species (Fig. 97b). Gong et al. (2006, p. 68) correctly discussed that the two bright areas could be macronuclear nodules so that one cannot exclude that Cohn’s population was binucleate. Interestingly, such binucleate populations have been reliably described by later workers, namely, by
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Kahl (1932) who found a “Trachelostyla-like” form with three cirral rows (Fig. 103e) and by Small & Lynn (1985; Fig. 102b), whose population is now assigned to T. rostrata (see there for details). Cohn (1866) described three cirral rows (two marginal and one ventral? three ventral plus two marginal?) on the ventral side of the body proper (see footnote above). However, Gong et al. (2006) considered the redescription by Maupas (1883) – who first described circularly arranged macronuclear nodules and the oxytrichid (Gonostomum-like) cirral pattern (Fig. 97c) – as authoritative and therefore correctly fixed such a population as neotype of T. pediculiformis to stabilise the situation (Fig. 96a–j). Kent (1882), the first revisor, did not change the generic classification proposed by Cohn (1866), but made a serious nomenclatural mistake (see nomenclature). Maupas (1883) provided a very detailed redescription. As already mentioned above, he recognised the cirral pattern more or less perfectly, that is, he observed – like Gong et al. (2006) in the neotype material – 11 cirri on the frontal area, two pretransverse ventral cirri, and five transverse cirri (cp. Fig. 96d with Fig. 97c). Further, he was the first to describe the nuclear apparatus, which consists of about 15 roughly globular macronuclear nodules peripherally arranged in body proper (Fig. 97c, e; however, see previous paragraph for the problem with the nuclear apparatus). Maupas (1883) transferred it to Gonostomum because of similarities in the oral apparatus. Gruber (1888) was uncertain about his identification. Indeed, due to the posteriorly narrowed body portion one cannot exclude that he observed Holosticha heterofoissneri Hu & Song, 2001, which has a very similar nuclear apparatus, but a different cirral pattern (for review, see Berger 2006, p. 152). Bütschli (1889) accepted the classification in Gonostomum proposed by Maupas (1883). Kahl (1928b) provided a detailed redescription which matches the data by Maupas (1883) and post-Kahlian workers (e.g., Borror 1972, Gong et al. 2006) very well. Kahl (1928b, Abb. 44a) found a second population which resembles the present species in body shape, but deviates distinctly in some other important features so that conspecificity with T. pediculiformis can be excluded (Fig. 103a); it is preliminarily classified as incertae sedis in Trachelostyla. Kahl (1932) again redescribed T. pediculiformis from life and recognised that it belongs neither to Stichochaeta (a synonym of Stichotricha) nor to Gonostomum and therefore established a new genus, which, however, is invalid (see genus section). In contrast, Wang & Nie (1932) accepted Maupas’ generic assignment, but they obviously misinterpreted the dorsal bristles as marginal cirri (which are therefore “very fine”) and considered the marginal rows as ventral rows. Trachelostyla spec. sensu Kahl is preliminarily classified in Cossothigma (Fig. 76e). Subsequently, Trachelostyla pediculiformis was redescribed in more or less detail several times (e.g., Kiesselbach 1936, Biernacka 1963, Kattar 1970, Jones 1974). Borror (1963) provided a rather detailed description of an American population. Very likely this is the first paper based on silver-impregnated material to confirm the pattern already recognised by Maupas (1883). Interestingly, Borror (1963) observed
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only three dorsal kineties, whereas six such rows are present in the neotype population. Borror (1972), who established Trachelostyla (see nomenclature), provided a detailed illustration of the ventral cirral pattern, which corresponds to that of Gonostomum, inter alia, in that the postoral ventral cirri (cirri IV/2, V/3, and V/4; see Berger 1999, p. 16) are displaced anteriorly. This pattern was confirmed recently by Gong et al. (2006), who discussed some differences between the many descriptions of T. pediculiformis, namely, the nuclear apparatus (see above), the caudal cirri, the number of frontoventral cirri, the dorsal kineties, and the cortical granules. Summarising the date, Trachelostyla pediculiformis seems to have “invariably” 18 frontalventral-transverse cirri, which can be easily homologised with the cirri of the “ordinary” 18-cirri hypotrichs (e.g., Oxytricha granulifera; details see Berger 1999; Lamtostyla granulifera, Fig. 40l, n) even without exact knowledge of cell division. The variability observed by some workers can be due to the following, as already discussed by Gong et al. (2006): (i) the cirral pattern is difficult to recognise in life in this species because it is opaque due to many cell inclusions; (ii) the cirral pattern shows, as in the other 18-cirri hypotrichs, a natural variability, that is, a cirrus is sometimes either absent, or supernumerary. Cohn (1866) is obviously the sole author who mentioned cortical granules (“gekörnte Linien”) in the present species. He found such granulated lines (rows) along the marginal rows and along the first, the middle, and last dorsal kinety (Fig. 97a, b). It is generally known that cortical granules often occur only along the cirral rows and the dorsal kineties. Thus, one cannot exclude that Cohn’s observation is correct. Stein (1859) also observed the cortical granules in several species very detailed and correct. By contrast, Gong et al. (2006) wrote that these organelles are only well recognisable with modern microscope techniques and therefore doubted that Cohn could truly tell the cortical granules apart from endoplasmic granules. Thus, they recommended that this difference should not be overinterpreted. A further difference between various descriptions is in the number of dorsal kineties which ranges from two (Kattar 1970) to six (Xu & Song 1999, Gong et al. 2006; Table 30). However, the exact dorsal kinety pattern (number and arrangement of kineties) is extremely difficult to recognise without protargol impregnation so that caution must be exercised. Kattar (1970), for example, very likely observed only the two laterally arranged dorsal kineties which are easily recognisable due to the long bristles. The same is true for the caudal cirri, which are clearly recognisable only when they are prominent (e.g., Stylonychia mytilus) or in protargol preparations. Anyhow, due to the neotypification of T. pediculiformis by Gong et al. (2006), the Chinese population is now relevant (Fig. 96a–j, Table 30). Stichochaeta corsica is the sole synonym of T. pediculiformis. The identity of these two species was first supposed by Hamburger & Buddenbrock (1929) and later accepted, inter alia, by Kahl (1932), Borror (1972), Maeda & Carey (1984), and Gong et al. (2006). Interestingly, the synonymy was never discussed in detail by most of these authors although there are some differences, for example, in the nu-
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Fig. 96a–e Trachelostyla pediculiformis (neotype population from Gong et al. 2006. a–c, from life; d, e, protargol impregnation). a: Ventral view of representative specimen, 112 µm. b: Left lateral view. c: Dorsal cilia and endoplasmic granules (length of cilia 7–8 µm). d, e: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 96 µm. Anteriorly displaced postoral ventral cirri circled, frontoventral cirri in rectangle, arrow in (d) marks pretransverse ventral cirri. For detailed labelling of cirri see Fig. 96f. Note that only three of the six dorsal kineties form a caudal cirrus; possibly, Trachelostyla pediculiformis has an unusual mode of dorsal kinety formation (see remarks at genus section). CC = caudal cirri, MA = macronuclear nodule, MI = micronucleus, PT = pretransverse ventral cirri, TC = transverse cirri, 1–6 = dorsal kineties. Page 478.
clear apparatus (one macronuclear nodule in S. corsica vs. around 10 in neotype of T. pediculiformis) and the contractile vacuole (in the rear body portion vs. likely lacking). However, both observations in S. corsica are obviously misobservations1 because neither a single, small ellipsoidal macronucleus, nor a contractile vacuole in the rear body end have never been reliably described in an interphasic hypotrich. Trachelostyla rostrata has two macronuclear nodules so that one cannot exclude that Gourret & Roeser (1888) overlooked the second (anterior) nodule. However, one also cannot exclude that Gourret & Roeser (1888) misinterpreted another organelle as macronucleus and that the contractile vacuole was displaced posteriorly due to the 1
The single macronuclear nodule was also interpreted as misobservation by Gong et al. (2006).
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Fig. 96f–i Trachelostyla pediculiformis (neotype population from Gong et al. 2006. f–i, protargol impregnation). f: Infraciliature of anterior body portion. Note the zigzag-shaped fibril along the adoral zone. g, h: Infraciliature of ventral side and nuclear apparatus of an early divider, size not indicated. Arrows in (g) mark oral primordia of proter and opisthe. Parental postoral ventral cirri circled. Long arrows in (h) mark micronuclei, short arrows mark replication band. i: Infraciliature of ventral side of middle divider, size not indicated. Asterisks mark primordia in dorsal kineties. AZM = distal end of adoral zone of membranelles (note that the distalmost 3–4 membranelles are enlarged), BC = buccal cirrus (cirrus II/2), E = endoral, FC = frontal cirri (cirri I/1, II/3, III/3), FVC = frontoventral cirri (cirri III/2, IV/3, VI/3, VI/4), LMR = left marginal row, P = paroral, PVC = anteriorly displaced postoral ventral cirri (cirri IV/2, V/3, V/4), RMR = right marginal row, 1, 6 = dorsal kineties. Page 478.
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preparation. Since no type material of S. corsica is available, we will never know what the French workers observed. Thus, I follow Hamburger & Buddenbrock (1929) and classify S. corsica as junior synonym of T. pediculiformis. Xu & Song (1999) described a Trachelostyla sp., likely from a Chinese marine habitat (Fig. 99a–e). A comparison with the neotype material described by Gong et al. (2006) shows no significant difference, except for the position of the rearmost postoral ventral cirrus, namely at the proximal end of the adoral zone (Fig. 99d) against close to the anterior two postoral ventral cirri (Fig. 96d). I do not know whether this difference is constant; if yes, then this population should be possibly separated at (sub)species level. Morphology: The description of the neotype population provided by Gong et al. (2006) is given first (Fig. 96a–j, Table 30). Subsequently, some relevant observations from the other populations are provided. The last paragraph deals with the Trachelostyla sp. described by Xu & Song (1999). Body size of neotype population 80–150 × 20–30 µm in life, on average about 120 × 25 µm. Body outline rather constant and characteristic because bipartite, that is, anterior fifth of body distinctly narrowed and set off from body proper, which has more or less parallel margins; posterior body end bluntly rounded. Body dorsoventrally flattened about 2:1, ventral side flat, dorsal side often vaulted in mid-body (Fig. 96a, b); flexible, but non-contractile. 9–17 macronuclear nodules arranged near cell trunk margin, that is, form elongate ring-shaped structure; individual nodules ovoid or ellipsoidal, about 5 × 4 µm in protargol preparations, containing small chromatin bodies. Two micronuclei inside ring of macronuclear nodules, one each in anterior and posterior region (Fig. 96e); interestingly, the same micronuclear pattern is described by Xu & Song (1999) for Trachelostyla sp. (Fig. 99e). No contractile vacuole observed. Pellicle thin. No cortical granules recognisable. Cytoplasm colourless and opaque, usually packed with granules 2–3 µm across (Fig. 96a, c) and 5–10 µm-sized food vacuoles. Movement rapid, showing great flexibility, sometimes jerky on microscope slide and between sediment particles. Adoral zone occupies about 40% of body length in life, and about 49% on average in protargol preparations (Fig. 96a, d, Table 30), composed of 40 membranelles on average; apical 3–5 membranelles strong, 15–20 µm long, and radially arranged; remaining membranelles distinctly shorter and arranged along left margin of narrowed anterior body portion; bases of largest membranelles about 3 µm wide. A zigzag-structure, likely composed of fibres, along adoral zone (Fig. 96d, f). Undulating membranes inconspicuous because short as compared to adoral zone; both membranes slightly curved, paroral about 13 µm long, endoral only about 6 µm. Cirral pattern and number of cirri of usual variability (Fig. 96d, f, Table 30). Cirral pattern basically as in other 18-cirri hypotrichs, except for the anteriorly displaced postoral ventral cirri. Three frontal cirri not distinctly larger than remaining cirri, distance between left and middle cirrus smaller than that between middle and right cirrus which is distinctly behind distal end of adoral zone (Fig. 96f). Buccal cirrus about at 50% of length of adoral zone, that is, distinctly ahead of anterior end
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of rather short undulating membranes. Frontoventral cirri form rectangular pattern with rear cirri about at level of buccal cirrus. Postoral ventral cirri, as in Gonostomum (for review, see Berger 1999, p. 367) and some other hypotrichs (e.g., Lamtostyla granulifera, Fig. 40n), right of proximal portion of adoral zone. Two pretransverse ventral cirri right of transverse cirri II/1 and III/1. Invariable five distinctly enlarged and 12–15 µm long transverse cirri arranged in hook- or Jshaped pattern. Right marginal row commences distinctly behind anterior body end, anteriormost cirri more widely spaced than remaining cirri, terminates at level of right transverse cirrus. Left marginal row begins left of proximal end of adoral zone, terminates distinctly ahead of rear cell end. Ventral and marginal cirri about 5–7 µm long. Dorsal bristles conspicuous because 7–8 µm long and stiff, arranged in six kineties of body length. Three inconspicuous caudal cirri about 5–7 µm long; middle and right cirrus often close to each other and arranged right of midline (Fig. 96d, e). Fig. 96j Trachelostyla pediculiObservations from other populations1 (Fig. 97a–o, formis (from Gong et al. 2006. 98a–k): Body length 100–200 µm (Maupas 1883), From life). Ventral view of a 150–200 µm (Kahl 1928b), 100–250 µm (Kahl 1932, neotype specimen. Long arrow 1933), about 150 µm (Biernacka 1963, Kattar 1970), marks enlarged, distalmost ado110 µm (Jones 1974), 136–196 µm (Maeda & Carey ral membranelles; short arrow marks transverse cirri. Page 1984), up to 200 µm (Carey 1992); body size 100 × 478. 20 µm (Cohn 1866), 129–255 × 24–32 µm (Bullington 1925, p. 271; no illustration), 114 × 22 µm (Wang & Nie 1932), 180 × 30 µm (Kiesselbach 1936), 135 × 28 µm (Borror 1963), 94.5–180 × 17.5–24.5 µm, anterior body portion 21–35 × 10.5 µm (Aladro Lubel 1985, Aladro Lubel et al. 1990). Ratio of length and width of head to body proper rather variable, depending on nutritional condition (Kahl 1932); in the Bay of Kiel (Baltic Sea) Kahl (1932) found gigantic forms where the heavily granulated body proper was distinctly enlarged against the relatively small head. Number of macronuclear nodules: 14–20 and two micronuclei, specimen shown in Fig. 97c with about 15 nodules (Maupas 1883); numerous nodules and usually two micronuclei (Kahl 1932), difficult to recognise even after staining (Kahl 1928b); 1 None of the populations, except that of Borror (1963, 1972), is described after protargol impregnation. Thus, the data about the cirral pattern (exact number and arrangement of cirri and dorsal kineties) must not be over-interpreted. Consequently, these data are omitted if they roughly fit the pattern described for the neotype population by Gong et al. (2006). Only when the pattern deviates very distinctly from that of the neotype material, then it is mentioned in the present review. For a brief characterisation of the synonym Stichochaeta corsica see remarks.
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Fig. 97a–o Trachelostyla pediculiformis (a, b, from Cohn 1866; c–h, from Maupas 1883; i, from Gourret & Roeser 1888; j, after Gourret & Roeser 1888 from Kahl 1932; k, from Kahl 1928b; l, from Kahl 1932; m, after Maupas 1883 from Kahl 1932; n, from Wang & Nie 1933; o, from Kahl 1933. From life). a–d, k–o: Dorsal (a, d), lateral (b), and ventral views, a, b = 100 µm, c, d, m = 200 µm, i, j = size not indicated, k = 150–200 µm, l = 180 µm, n = 114 µm, o = 100–250 µm. (i, j) show the synonym Stichochaeta corsica, which has, according to the original description, only one macronucleus (small arrow in j). e, g, h: Malformed specimens. f: Nuclear apparatus of dividing specimen. Page 478.
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Fig. 98a–k Trachelostyla pediculiformis (a, from Kiesselbach 1936; b, from Biernacka 1963; c, from Borror 1963; d, from Kattar 1970; e, from Borror 1972; f, from Jones 1974; g, from Maeda & Carey 1984; h, from Burkovsky 1984; i, from Aladro Lubel 1985; j, from Aladro Lubel et al. 1990; k, after Maeda & Carey
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nine plus two micronuclei (Kiesselbach 1936); 16–64(!) (Borror 1963); 18 (Kattar 1970); 32–64 and many micronuclei (Kool 1970); 13–18 (Hartwig 1973, p. 453); 26–30, about 2–3 µm across (Jones 1974; at least two micronuclei); 20–35, about 4 µm across (Aladro 1985, Aladro Lubel et al. 1990). Nuclear apparatus and contractile vacuole not recognised by Wang & Nie (1932). Contractile vacuole lacking according to Kiesselbach (1936), Biernacka (1963), Jones (1974), and Aladro Lubel et al. (1990), according to Kahl (1932) usually lacking. Cytoplasm usually packed with greasily shining granules, making cells bright-grey and opaque (Cohn 1866, Kahl 1932). Cortex (ectoplasm) fine, fragile (Kahl 1932). Lively movement (Kahl 1932); creeps between filaments of decaying algae, winds through tiny spaces feeling its way with the narrowed head, sometimes it moves hastily backwards in a straight or curved line, and the anteriormost adoral membranelles and the cirri at the rear cell end are likely used for jumping movements (Cohn 1866). Rotates to the left during movement (Bullington 1925). Highly thigmotactic (Bock 1953, Borror 1963). Adoral zone occupies almost 50% of body length (Carey 1992); composed of about 41 membranelles (Borror 1963). 11 cirri (14–17 µm long) right of adoral zone, frontal and transverse cirri 17–21 µm long (Aladro Lubel et al. 1990); 11–12 cirri right of adoral zone (frontal cirri enlarged), 23 left marginal cirri (12–14 µm long), 32–35 right marginal cirri, five transverse cirri about 28 µm long (Borror 1963). According to Kahl (1932), the elongated three bristles at the rear cell end are dorsal cilia and not caudal cirri (Fig. 97l). However, the description by Gong et al. (2006) shows that caudal cirri are present (Fig. 96d). Trachelostyla sp. sensu Xu & Song (1999, Fig. 99a–e, Table 30), described in Chinese, is possibly a distinct (sub)species. Further populations have to be studied to show whether the difference (rearmost postoral ventral cirrus distinctly set off) is constant. Body size 175–200 × 20–25 µm in life, that is, body very slender and head not very distinctly set off from body proper (Fig. 99a, b). Dorsal bristles 7–8 µm long. Further details (cirral pattern, dorsal kinety pattern, nuclear apparatus), see Fig. 99a–e and Table 30. Cell division: Kool (1970) provided a brief description of the cell division in T. pediculiformis (see also Foissner 1996, p. 109). Cortically it commences with the formation of fields in which the adoral zone originates as a longitudinal series of transverse streaks of cilia while the 4–6 frontal-ventral-transverse cirri anlagen are longitudinally arranged. Interestingly, both the proter and the opisthe form a new adoral zone. The anlagen formation resembles that of Stylonychia as described by Wallengren (for review, see Berger 1999); furthermore, the same number (18) of frontal-ventral-transverse cirri is formed (however, note that the 18-cirri pattern, including its formation is an apomorphy of the Hypotricha!). Both the right and left marginal row primordia originate right to the parental rows. The new adoral zone of the proter elongates and replaces the parental zone. The population studied by Kool
b
1984 from Carey 1992. a, b, d, f–k, from life; c, e, silver impregnation). Ventral views, a = 180 µm, b = 150 µm, c = 135 µm, d = 145 µm, e = 92 µm, f = 110 µm, g = 114 µm, h = 150 µm, i = 95 µm, j = 93 µm, k = size not indicated. Page 478.
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Fig. 99a–e Trachelostyla pediculiformis (from Xu & Song 1999. a–c, from life; d, e, protargol impregnation). a: Ventral view of a representative specimen, 187 µm. b: Shape variant. c: Specimen attached to substrate with transverse cirri and moving body. d, e: Infraciliature of ventral and dorsal side and nuclear apparatus, individual size not indicated. This population differs from the neotype material, besides the body size (see Table 30), in the location of the rearmost postoral ventral cirrus (rearmost arrow in d) which is distinctly set off from the other two cirri (anteriormost arrows); thus, conspecificity with neotype material is not certain. Frontal cirri connected by dotted line; right frontal cirrus and cirrus III/2 as well as frontoterminal cirri (cirri VI/3 and VI/4) connected by broken line. Frontoventral cirri circled. BC = buccal cirrus, CC = caudal cirri, MI = micronucleus, 1, 6 = dorsal kineties. Page 478.
(1970) has a rather high number of macronuclear nodules (32–64 against, e.g., 9–17 in neotype population). As is usual, they fuse to a single mass (plus three micronuclei). Gong et al. (2006) found an early and a middle divider and described the following features: (i) the frontal-ventral-transverse cirri primordia originate de novo on
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Fig. 100a Trachelostyla pediculiformis (from Hartwig 1973). Population dynamics in a habitat on the island of Heligoland. Page 478.
the cell surface in both proter and opisthe (Fig. 96g); (ii) replication bands occur in early stages of cell division (Fig. 96h); (3) dorsal kinety anlagen originate from the parental kineties (Fig. 96i). Recently, Shao et al. (2007) described the cell division in detail (see Addenda). Molecular data: The complete 18S rRNA gene sequence of T. pediculiformis is 1769 nucleotides long (Gong et al. 2004). The present species is included in phylogenetic analyses by Gong et al. (2006, p. 71) and Schmidt et al. (2007); however, the results are rather different (see remarks in genus section). Occurrence and ecology: Common in marine habitats (Kahl 1933, Patterson et al. 1989, Fenchel 1968), but also in inland salt waters (Kahl 1928b), and the brackish part of estuaries (Mazei & Burkovsky 2006, p. 259); According to Kahl (1932) mainly in mesosaprobic and sapropelic detritus. Fenchel & Jansson (1966) found it at all depths of the sediment with a maximum usually some centimetres below the surface. Hartwig (1973a, p. 129) recorded it mainly in weakly and moderately lotic areas, but not in strongly lotic regions of the sandy littoral area. From April to October most specimens occur in the upper (0–5 cm) sediment layer, during the cold season most specimens can be found in deeper sediment areas; during November and December 1969, Hartwig (1973a) even found specimens at a sediment depth of 35 cm where reductive conditions occurred. Due to the neotypification the type locality is the coast (39°10'N 117°10'E) of the Bohai Sea (China) near the city of Tianjin. Gong et al. (2006) found it there in the sandy littoral sediments at a water temperature of 18°C in August 2003. They set up a raw culture at 20°C room temperature using boiled seawater (salinity 30‰). Cohn (1866) found his specimens in an aquarium containing marine material from the is-
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land of Heligoland (North Sea) and from Dorsetshire (UK), Atlantic Ocean. Type locality of the synonym Stichochaeta corsica is the harbour of the city of Bastia (Corsica), Mediterranean Sea (Gourret & Roeser 1888). Further records substantiated by morphological data: regularly, but never abundant in the Brennermoor, a saline (25‰), silt peat bog near the north German city of Bad Oldesloe (Kahl 1928a, b); Baltic Sea, inter alia, in the Bay of Kiel (Kahl 1932); at about 7‰ salinity in the Bay of Danzig (Poland), Baltic Sea (Biernacka 1962, p. 78; 1963); up to 3800 ind. l-1 with a strong preference of the summer season in various sites at the sandy beaches of the islands of Sylt (Germany) and Jordsand (Denmark), North Sea (Hartwig 1973, Fig. 100a; see also Hartwig 1974, p. 17); Chichester Harbour (United Kingdom), North Sea (Maeda & Carey 1984; see also Carey & Maeda 1985, p. 565); harbour of Algiers (Algeria), Mediterranean Sea (Maupas 1883); Val di Lone, Rovinj (Croatia), Northern Adriatic Sea (Kiesselbach 1936); common in sand and other material (shell fragments, exoskeletons of isopods, dead oysters, and detritus) at Aligator Harbor (Florida), Gulf of Mexico (Borror 1963; see also Borror 1962, p. 343); in July in sand samples at Dauphin Island, Alabama Bay, Gulf of Mexico (Jones 1974); various marine sites in Mexico (Aladro Lubel 1985; Aladro Lubel et al. 1990, Aladro-Lubel et al. 1986, p. 240; 1988, p. 438; Mayén Estrada & Aladro Lubel 1987, p. 76; Madrazo Garibay & López-Ochoterena 1982, p. 293); beach (Praia do Embaré, Santos-São Vicente-Guarujá) near São Paulo (Brazil), Atlantic Ocean (Kattar 1970); Bay of Amoy (now Xiamen; China), Chinese Sea (Wang & Nie 1932; see also Wang & Nie 1934, p. 4210). Records not substantiated by morphological data: sand and mud flats near Plymouth (United Kingdom), Atlantic Ocean (Lackey & Lackey 1963, p. 803); Loch Eil and the Lynn of Lorne, west coast of Scotland, Atlantic Ocean (Wyatt & Pearson 1982, p. 301); sublittoral areas of the Sea of Cantabria, Bay of Biscay (Spain), Atlantic Ocean (Fernandez-Leborans 2000, p. 417; Fernandez-Leborans & Novillo 1993, p. 216; Fernandez-Leborans et al. 1999, p. 742); tidal sand flat off Crildumersiel (salinity 32‰), Jade Bay, North Sea (Hartwig 1984, p. 127); Königshafen, a harbour on the German North Sea island of Sylt (Küsters 1974, p. 175); Bran Sand, a sheltered beach in the Tees estuary (Great Britain) and other sandy beaches along the coast of North Yorkshire, North Sea (Hartwig & Parker 1977, p. 751; Parker 1981, p. 338); North Sea at Norwegian coast (Fjeld 1955; cited from Hartwig & Parker 1977, p. 751); Schlei, a large brackish water in North Germany (Bock 1960, p. 64; Jaeckel 1962, p. 13); fine sand (median grain size 95 µm) at a water depth of 22 m near Ålsgårde (Denmark), Kattegat (Fenchel 1969, p. 104); Bay of Kiel (Germany), Baltic Sea (Bock 1952, p. 81, 83); brackish bay at the Bottsand, Kieler Außenförde (Germany), Baltic Sea (Ax & Ax 1960, p. 12, 15, tolerating 15–20‰ salinity); beach of the island of Askö (Sweden), Baltic Sea (Fenchel & Jansson 1966, p. 176); brackish water in the Gdansk Bay, Baltic Sea (Czapik & Jordan 1976, p. 442; Czapik & Fyda 1992, p. 110); Amphioxus sand near the city of Marseilles (France), Mediterranean Sea (Vacelet 1961a, p. 15; 1961b, p. 3); harbour of Genoa (Italy), Mediterranean Sea (Gruber 1884, p. 482); Gulf of Naples (Italy), Mediterra-
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nean Sea (Nobili 1957, Tabella I; further records from Italy: Dini et al. 1995, p. 71); 201 ind. cm-2 in the 0–1 cm sediment layer of the lagoon at Sacca di Goro (Italy), North Adriatic Sea (Madoni 2006, p. 169); mesopsammon of Adriatic Sea near the city of Kuçova, Albania (Czapik 1952, p. 65); White Sea estuary (Mazei et al. 2002, p. 389); sites of White Sea distinctly influenced by freshwater (Burkovsky 1976, p. 288); Kandalaksha Gulf, White Sea (Azovsky et al. 1996, p. 30, as T. pediculiforme; Burkovsky 1968, p. 1409; 1970, p. 11; 1970b, p. 190; Mazei & Burkovsky 2005, p. 116; further records from White Sea: Burkovsky 1970a, p. 56; 1971a, p. 1570; 1971b, p. 1774; 1971c, p. 1289); Black Sea and saline Lake Tekirghiol, Romania (Tucolesco 1962, p. 814; 1965, p. 160); coarse ground of Caucasian coast of Black Sea (Azovskii & Mazei 2003, p. 904); at 21–26°C and 17–18‰ salinity at Bulgarian coast of Black Sea (Detcheva 1980, p. 35; 1982, p. 249; 1983, p. 72; 1992, p. 104; Kovaleva & Golemansky 1979, p. 275; further records from Black Sea: Azovsky & Mazei 2003, p. 84); saline lake near the Bay of Kruglaja, Crimea (Dagajeva 1930, p. 35); rare in a saline lake (Chadjebej-Liman; 50–70‰ salinity) near the city of Odessa, Ukraine (Butschinsky 1897, p. 195); Western Caspian Sea (Agamaliev 1967, p. 369; Agamaliyev 1974, p. 21); western coast of Novaya Zemlya (Russia), Barents Sea (Azovsky 1996, p. 6); Pechora Sea, a part of the Barents Sea (Azovsky & Mazei 2007, p. 62); Peter the Great Bay, Sea of Japan (Myskova 1976, p. 85); tidal marshes at Odiorne Point and/or Adams Point (New Hampshire, USA), Atlantic Ocean (Borror 1972a, p. 67; Martinez 1980, p. 370); sand and (saline) ponds near the Narragansett Laboratory, Rhode Island (USA), Atlantic Ocean (Lackey 1961, p. 277); littoral zone at St. John’s Bayou (New Orleans; USA), Gulf of Mexico (Smith 1904, p. 45); during June and September on Halodule beaudettei (sea grass) in the Tamiahua Lagoon (Mexico), Gulf of Mexico (Martínez-Murillo & Aladro-Lubel 1994, p. 14); north shore of Bermuda Island, Atlantic Ocean (Hartwig 1980, p. 426); Bay of Rio de Janeiro, Brazil (Faria et al. 1922, p. 113; Pinto 1925, p. 257; Petz 1999); in July in Sepetiba Bay in the south region of Rio de Janeiro State, Brazil (Wanick & Silva-Neto 2004, p. 5); Brazil (Cunha 1916, p. 67; as Stichochaeta pediculiforme). Trachelostyla pediculiformis feeds on bacteria (Borror 1963, Fenchel 1968, p. 117; 1969, p. 25; Hartwig 1973b, p. 329), according to Borror (1963) also on “other colourless microflora”. Well-growing cultures were obtained when specimens were isolated in a dish of filtered sea water containing a few rice grains (Borror 1963, Kool 1970). Martinez (1980) found following relevant temperatures: optimum for growth 31°C; upper limit for significant growth 34°C; upper limit for survival 35°; LD50 36.1°C. The lower limit for survival is, as in Epiclintes auricularis (for review see Berger 2006, p. 1119), -3°C (Hartwig 1973a, p. 148). Tolerates up to 10 mg l-1 H2S (Fenchel 1969, p. 139; according to Hartwig 1973a, p. 154, it tolerates 50 and more mg l-1 H2S) and according to Hartwig (1973a, p. 151, 155) oxidative as well as reductive areas.
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Trachelostyla caudata (Kahl, 1932) Borror, 1972 (Fig. 101a–d) 1932 Trachelostyla caudata spec. n. – Kahl, Tierwelt Dtl., 25: 597, Fig. 1141 (Fig. 101a; original description; no type material available and no formal diagnosis provided). 1933 Trachelostyla caudata Kahl 1932 – Kahl, Tierwelt N.- u. Ostsee, 23: 112, Fig. 17 24 (Fig. 101d; guide to marine ciliates). 1972 Trachelostyla caudata Kahl, 1932 – Borror, J. Protozool., 19: 15 (combination with Trachelostyla Borror, 1972; generic revision of hypotrichs and euplotids; see nomenclature). 1977 Trachelostyla caudata Kahl 1932 – Buitkamp, Decheniana, 130: 125 (brief review of Trachelostyla). 1982 Trachelostyla caudata Kahl, 1932 – Hemberger, Dissertation, p. 261 (revision of hypotrichs). 1984 Trachelostyla caudata Kahl, 1932 – Maeda & Carey, Bull. Br. Mus. nat. Hist. (Zool.), 47: 7, Fig. 2 (Fig. 101b; revision). 1992 Trachelostyla caudata (Kahl, 1930–5) Maeda and Carey, 1984 – Carey, Marine interstitial ciliates, p. 186, Fig. 736 (Fig. 101c; guide; redrawing of Fig. 101b; incorrect combining authors). 1999 Trachelostyla caudata (Kahl, 1932) Maeda and Carey, 1984 – AL-Rasheid, Agricultural Sciences, 4: 58, Fig. 8 (micrograph of protargol-impregnated specimen; incorrect combining authors). 2001 Trachelostyla caudata Kahl, 1932 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 89 (nomenclator containing all basionyms, combinations, and higher taxa of euplotids and hypotrichs). 2002 Trachelostyla caudata – Hu & Song, Hydrobiologia, 481: 177, Fig. 2g–i (Fig. 101a–c; brief discussion and key to Trachelostyla species). 2006 Trachelostyla caudata Kahl, 1932 – Gong, Song, Li, Shao & Chen, Europ. J. Protistol., 42: 72, Fig. 30N (Fig. 101a; brief review).
Nomenclature: No derivation of the name is given in the original description. The species-group name caudat·us, -a, -um (Latin adjective [m, f, n]; tailed, having a tail) refers to the tail-like posterior body end. Trachelostyla Kahl, 1932 is not available because no type species was fixed (see same chapter at genus section). Thus, Trachelostyla caudata Kahl, 1932 had to be transferred to the valid genus Trachelostyla Borror, 1972. Although Borror (1972) did not realise that Trachelostyla Kahl, 1932 was not available, I consider him as combining author because he (i) is also the author of the valid genus Trachelostyla; and (ii) considered T. caudata as valid member of Trachelostyla. Remarks: Trachelostyla caudata differs from the type species in the body shape, that is, by the tail-like posterior body portion (vs. not narrowed and broadly rounded) and the length of the dorsal bristles (12–15 µm [Kahl 1932] vs. 7–8 µm in neotype population of T. pediculiformis). Maeda & Carey (1984) redescribed T. caudata and largely confirmed Kahl’s observations. The micrograph provided by ALRasheid (1999a) shows a more or less tailed specimen with twenty or more macronuclear nodules. The cirral pattern is not recognisable so that the identification cannot be checked; according to the description, four frontoventral cirri, two rows of marginal cirri, and an oblique row of long transverse cirri are present. Interestingly, in all descriptions the number of cirri on the frontal area is rather low (4–5), that is, some cirri (e.g., postoral ventral cirri) are obviously lacking in this species (Fig. 101a–d); this would be a further important difference to the type species. However,
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a detailed investigation of the ventral and dorsal ciliature is needed for an in-depth discussion. Trachelostyla caudata sensu Petran (1967) and sensu AL-Rasheid (2001) are classified as insufficient redescriptions. Morphology: The following paragraphs are based on the original description (Kahl 1932) unless otherwise indicated. The description is supplemented by additional or deviating data by Maeda & Carey (1984). Since both descriptions are based only on live observations, some details (e.g., number of dorsal kineties) must not be over-interpreted. Kahl (1932) wrote that T. caudata looks like the type species, except for the data presented in the original description. Body length 150–220 µm in life (120–240 µm according to Kahl 1933); length:width ratio of specimen illustrated about 6:1 (Fig. 101a); length according to AL-Rasheid (1999a) 150 µm (from life?). Maeda & Carey (1984) mention a body length of 156 µm; however, according to Fig. 101b this size includes the anteriorly directed adoral membranelles and the transverse cirri; the length of the body alone is about 120 µm. Body outline spindle-shaped, that is, anterior (head) and posterior (tail) fourth distinctly narrowed. Body obviously fragile because it does not withstand cover glass pressure, thus difficult to observe in life; according to Maeda & Carey (1984), body fragile and elastic, but not contractile. Nuclear apparatus as in T. pediculiformis, that is, several (around 10 in Fig. 101a) macronuclear nodules arranged in central body portion; specimen illustrated by Maeda & Carey (1984) with 12 nodules (Fig. 101b); specimen shown by AL-Rasheid (1999a) with more than 20 nodules 3–5 µm across. Contractile vacuole not described by Kahl (1932), near proximal end of adoral zone according to Maeda & Carey (1984). Presence/absence of cortical granules not mentioned. Movement as in T. pediculiformis, but head region not lifted from substrate during forward movement (Maeda & Carey 1984). Oral apparatus basically as in the type species (Kahl 1932, Maeda & Carey 19841). Distalmost four (five according to Maeda & Carey 1984) adoral membranelles conspicuously splayed (Fig. 101a, b). Remaining membranelles rather short and brush-like (Maeda & Carey 1984). Cirral pattern not known in detail (Fig. 101a, b). Both according to Kahl (1932) and Maeda & Carey (1984) only four cirri on area right of adoral zone, that is, several of the 11 cirri present in the type species are lacking. No cirri on postoral area. Pretransverse ventral cirri not described by Kahl (1932) and Maeda & Carey (1984) whereas in the type species these cirri are present; possibly they have been overlooked in T. caudata because they are usually rather small and thus inconspicuous. Five strong transverse cirri, more prominent than in type species; right cirri extend by 2/3 of their length beyond body margin. Right marginal row commences near anterior cell end, ends about at level of transverse cirri (Fig. 101a); according to 1
Maeda & Carey (1984) described a “characteristically long membrane” along the lateral portion of the adoral zone of membranelles in T. pediculiformis (Fig. 98g). According to the redescription of the type species by Gong et al. (2006) this cannot be an undulating membrane because both paroral and endoral are very short in T. pediculiformis. Possibly it is the buccal lip. Anyhow, in T. caudata Maeda & Carey (1984) did not see this “characteristically long membrane”.
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Fig. 101a–d Trachelostyla caudata (a, from Kahl 1932; b, from Maeda & Carey 1984; c, from Carey 1992 [possibly a redrawing of Fig. 101b]; d, from Kahl 1933. a–d, from life). Ventral views, a = 150–220 µm, b = 120 µm, c = 150 µm (possibly inclusive adoral membranelles and transverse cirri, see morphology), d = 120–240 µm. Longitudinal arrow in (b) marks contractile vacuole, transverse arrow denotes anterior end of right marginal row (note difference to type population where this row commences near anterior cell end). AZM = adoral zone of membranelles, DB = dorsal bristles, TC = right transverse cirrus. Page 494.
Maeda & Carey (1984) it starts at base of narrowed head. Left marginal row extends from proximal end of adoral zone to near left transverse cirrus. Dorsal bristles 12–15 µm long, arranged in 8–10 rows (Fig. 101a). According to Maeda & Carey (1984) the dorsal bristles are not as long as those of T. pediculiformis; however, this contradicts the observations by Kahl (1932) and Gong et al. (2006; see remarks). Caudal cirri and/or elongated dorsal bristles at rear cell end lacking (Fig. 101a, b). Note, however, that details about the dorsal ciliature (number and arrangement of kineties, presence/absence of caudal cirri) must not be over-interpreted.
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Occurrence and ecology: Predominantly in (possibly confined to) marine habitats; benthic, mainly in fine sediments up to a depth of 20 cm (Hartwig 1973a, p. 128; Patterson et al. 1989, p. 211; Agamaliev 1970, p. 1279); however, the upper 5 cm, that is, the aerobic areas are preferred (Hartwig 1973a). The type locality of T. caudata is the Bay of Kiel (Germany), Baltic Sea, where Kahl (1932) discovered it on rather clean, sandy sediment at a water depth of 5–6 m; here and there abundant; rarely at the island of Heligoland (Kahl 1933). Maeda & Carey (1984; see also Carey & Maeda 1985, p. 565) found T. caudata in the fine sediments of East-Head, Chichester Harbour (England), mainly in sands containing a lower content of debris than the biotopes harbouring T. pediculiformis. They collected the samples in October 1982 and July 1983 at a water temperature of 14°C and 24°C, respectively. ALRasheid (1999a; 1999b, p. 353) found T. caudata in the upper sediment layer (0–3 cm) of the Jubail Marine Wildlife Sanctuary (Saudi Arabia), Arabian Gulf. Records from marine habitats not substantiated by morphological data: French coast of Atlantic Ocean (Fauré-Fremiet 1950, p. 67); sublittoral of Stoller Grund, a part of the Kiel Bay, Baltic Sea (Bock 1952, p. 83); up to 200 ind. l-1 in littoral sediments of the Island of Sylt (Germany), North Sea, with a preference of the warm season, that is, May to October (Hartwig 1973, p. 452; Küsters 1974, p. 175; see also Hartwig 1974, p. 17); Gulf of Naples (Italy), Mediterranean Sea (Nobili 1957, Tabella I; see also Dini et al. 1995, p. 71); Cantabrian Sea (Spain), Bay of Biscay, Atlantic Ocean (Fernandez-Leborans 2000, p. 417; Fernandez-Leborans & Novillo 1993, p. 216); Black Sea and saline Lake Tekirghiol, Romania (Tucolesco 1962, p. 813; Tuculescu 1965, p. 160); sandy beaches of Black Sea (Băcescu et al. 1967, p. 7; Kovaleva 1966, p. 1603; Kovaleva & Golemansky 1979, p. 270, 275; see also Detcheva 1992, p. 103); coarse ground of Caucasian coast of Black Sea (Azovskii & Mazei 2003, p. 904; Azovsky & Mazei 2005, p. 89); up to a depth of 7 cm in sandy sediment of the Odessa Bay, Black Sea (Dzhurtubayev 1978, p. 65; further record from Black Sea: Azovsky & Mazei 2003, p. 84); Apsheron coast of Caspian Sea (Agamaliev 1990, p. 61); Krasnovodsky Bay and Turkmen Bay at eastern coast of Caspian Sea (Agamaliev 1973, p. 1600; 1974a, p. 20); Bol’shoy Kyzylachag Bay and Kizlyar Bay in western Caspian Sea (Agamaliyev 1974, p. 21); Divichinsky lagoon, a low salinity site of Caspian Sea (Agamaliev 1986, p. 207; Agamaliyev & Aliyev 1983, p. 21; further records from Caspian Sea: Agamaliev 1967, p. 369; 1970, p. 1279; 1972, p. 7; 1974, p. 57; 1983, p. 36; Agamaliyev 1976, p. 91); Norwegian coast at Drøbak (Fjeld 1955; cited from Hartwig 1973, p. 453); sites of White Sea distinctly influenced by freshwater (Burkovsky 1976, p. 288); Chernaya River estuary, White Sea (Mazei & Burkovsky 2003, p. 111); Kandalaksha Bay, including polluted areas, and other sites of White Sea (Burkovsky 1970, p. 11; 1970a, p. 56; 1970b, p. 190; Azovsky et al. 1996, p. 30; Raikov 1962, p. 331; Mazei & Burkovsky 2005, p. 116); western coast of Novaya Zemlya, Barents Sea (Azovsky 1996, p. 6); sandy littoral of Barents Sea (Kovaljeva 1967, p. 83); Bay of Dalnie Zelentzy (Murmanskaya), Barents Sea, Russia (Raikov 1960; cited from Hartwig 1973, p.
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453); sand of Ussuri Gulf and Posjet Gulf, Japan Sea (Raikov 1963, p. 1757; Raikov & Kovaleva 1968, p. 331). Record from a freshwater habitat: Agrakhan Bay, a freshwater region (pH 7.1, average oxygen content 11.4 mg l-1) of Caspian Sea (Agamaliev 1986, p. 207). Food not mentioned by Kahl (1932) and Maeda & Carey (1984). For notes on the ecology, including nutrition and production, see Fauré-Fremiet (1950, p. 67), Agamaliev (1967a, p. 1426), Burkovsky et al. (1980, p. 331), Burkovskii & Azovskii (1985a, p. 616; 1985b, p. 815), and Burkovsky (1978, p. 326; 1984, e.g., p. 89; 1987, p. 648, 650).
Trachelostyla rostrata (Lepsi, 1962) comb. nov. (Fig. 102a, b) 1962 Trachelostyla rostrata n. sp. – Lepsi, Zool. Anz., 168: 465, Abb. 30 (Fig. 102a; original description; no type material available and no formal diagnosis provided). 1972 Trachelostyla rostrata Lepsi, 1962 – Borror, J. Protozool., 19: 19 (generic revision of hypotrichs and euplotids; see nomenclature). 1985 Trachelostyla pediculiformis – Small & Lynn, Phylum Ciliophora, p. 461, Fig. 44B (Fig. 102b; misidentification; illustration of a protargol-impregnated specimen). 1992 Trachelostyla rostrata Lepsi, 1964 – Carey, Marine interstitial ciliates, p. 186 (brief note; incorrect year). 2001 Trachelostyla rostrata Lepsi, 1962 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 90 (nomenclator containing all basionyms, combinations, and higher taxa of euplotids and hypotrichs). 2002 Trachelostyla pediculiformis – Lynn & Small, Phylum Ciliophora, p. 458, Fig. 56B (Fig. 102b; misidentification; illustration of a protargol-impregnated specimen). 2006 Trachelostyla rostrata Lepsi, 1962 – Gong, Song, Li, Shao & Chen, Europ. J. Protistol., 42: 69, 70, 72, Fig. 30M, R (Fig. 102a, b; brief review).
Nomenclature: No derivation of the name is given in the original description. The species-group name rostrat·us, -a, -um (Latin adjective [m, f, n]; beaked, having a beak, snout, trunk, or proboscis) refers to the narrowed anterior body end. Trachelostyla Kahl, 1932 is not available because no type species was fixed (see same chapter at genus section). Thus, Trachelostyla rostrata Lepsi, 1962 was established in an invalid genus. Borror (1972), who is the valid author of Trachelostyla (see genus section), did not classify the present species in Trachelostyla, but listed it as species of questionable systematic position. Thus, he cannot be considered as combining author. Since nobody else transferred T. rostrata to Trachelostyla Borror, 1972, this act is done in the present paper. Remarks: Trachelostyla rostrata is rather poorly described. However, the general appearance indicates that the classification in Trachelostyla is correct. The two macronuclear nodules are a significant difference to the other two species presently assigned to Trachelostyla, which have each about 10 or more nodules. Lepsi (1962) used nuclear staining so that it is unlikely that he misinterpreted other organelles as macronuclear nodules or overlooked small nodules. In addition, Small & Lynn (1985) found a Trachelostyla with only two macronuclear nodules, strongly indicat-
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ing that the American population is identical with the population studied by Lepsi (1962). By contrast, the differences in the cirral pattern are distinct (marginal rows lacking in type population vs. present in American population), but must not be over-interpreted because Lepsi (1962) may have overlooked these cirri which are distinctly smaller than the transverse cirri. Thus, I agree with Gong et al. (2006) that these two populations are conspecific. However, since Small & Lynn (1985) did not provide further details (e.g., live data, morphometry) I do not fix the American population as neotype. Maeda & Carey (1984) listed Lepsi (1962) in their reference section; however, I could not find a note about the present species in their paper. According to Carey (1992) they were unable to give it a firm taxonomic position. Hu & Song (2002) did not consider T. rostrata in their brief review, whereas Gong et al. (2006) classified it as valid species. Detailed investigation needed for a more in-depth discussion. Fig. 102a, b Trachelostyla rostrata Morphology: The present paragraph is solely (a, from Lepsi 1962; b, from Small & based on the somewhat superficial original de- Lynn 1985. a, from life; b, protargol scription and the illustration provided (Fig. 102a). impregnation?). a: Ventral view (120 to 170 µm) showing body outline, Body length 120–170 µm in life. Body short club- macronuclear nodules, and contractile shaped. Two widely separated macronuclear nod- vacuole. Cirral pattern obviously not ules. Contractile vacuole at proximal end of adoral completely recognised. b: Infracilazone. According to Lepsi (1962) neither marginal ture of ventral side, 85 µm. Pretranscirri, oblique ventral cirri, nor dorsal bristles pre- verse ventral cirri circled. FC = frontal cirri, MA = macronuclear nodule sent; however, this must not be over-interpreted. (micronucleus attached), UM = unduThe “lack” of the dorsal bristles indicates that they lating membranes. Page 498. are short, that is, around 3 µm. The illustration shows strong anterior adoral membranelles, two strong cirri (adoral membranelles?) near (at?) the proximal end of the adoral zone, and five prominent transverse cirri (Fig. 102a). Specimen illustrated by Small & Lynn (1985) about 85 × 13 µm after protargol impregnation. Anterior body third (head) distinctly narrowed, rear cell end rounded, margins of body proper parallel. Two macronuclear nodules slightly right of midline, each with a micronucleus. Adoral zone as in congeners, undulating membranes short, very close to proximal portion of adoral zone (Fig. 102b). Three frontal cirri, one(?) buccal cirrus, and eight further cirri (frontoventral cirri; anteriorly displaced postoral ventral cirri) right of adoral zone. No cirri on postoral area. Two pretrans-
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verse ventral cirri and five transverse cirri. In total thus 19 frontal-ventral-transverse cirri, that is, one more than in the type species, which has the ordinary number of 18 cirri. Right marginal row commences slightly ahead of level of buccal vertex, terminates somewhat behind level of transverse cirri; left row begins left of proximal end of adoral zone, terminates slightly ahead of level of transverse cirri. Dorsal ciliature (length of bristles; number and arrangement of dorsal kineties; presence/absence of caudal cirri) not known. Occurrence and ecology: Marine. Type locality of Trachelostyla rostrata is the Black Sea, where Lepsi (1962) discovered it in littoral-sediment at the Zoological Station Agigea-Dobrudscha, Romania. It was abundant in a sand culture. Small & Lynn (1985) discovered their population in sediments from the Chesapeake Bay, East Coast of the USA. No further records published.
Incertae sedis in Trachelostyla Stichochaeta pediculiformis Cohn (?) – Kahl, 1928, Arch. Hydrobiol., 19: 210, Abb. 44a (Fig. 103a). Remarks: Kahl found this population for some time in saline inland waters. Body shape as shown in Fig. 103a, that is, basically as in T. pediculiformis. Size not mentioned, but likely also similar as in T. pediculiformis. Body coarsely and darkly granulated. Since Kahl (1928b) was not certain about the details he announced a more detailed comparison between T. pediculiformis (Fig. 97k) and the present population; but in his 1932 monograph the latter population was no longer considered. Three frontal cirri and about six cirri right of adoral zone (frontalventral cirri and buccal cirrus/cirri). Five transverse cirri. According to Kahl, the present population has – besides the marginal rows (number not mentioned) – two well developed ventral rows. However, Fig. 103a shows five cirral rows (from right to left): (i) a right marginal row; (ii) a row commencing about at same level as right marginal row; (iii) a row (perhaps only a furrow?) commencing behind the buccal vertex; (iv) a row beginning left of the proximal portion of the adoral zone (possibly the inner left marginal row?); and (v) a left marginal row. Dorsal bristles roughly of ordinary length. Very likely only two macronuclear nodules. No further details described. Found in saline inland water (Salzwiesen von Altfresenburg; salinity 20‰; details see Kahl 1928a, p. 52) near the village of Bad Oldesloe in northern Germany. The higher level classification of this form is uncertain. Fig. 103a Stichochaeta pediculiformis (from Kahl 1928b. From life). Ventral view, size not indicated. DB = dorsal bristles, TC = transverse cirri. 1–5 = cirral rows (details see text). Page 500.
Trachelostyla
501
Table 30 Morphometric data on Trachelostyla pediculiformis (pe1, neotype population from Gong et al. 2006; pe2, Trachelostyla sp. from Xu & Song 1999) Characteristics a
Population mean
Body, length Body, width Adoral zone of membranelles, length Macronuclear nodule, diameter Macronuclear nodules, number Micronuclei, number Adoral membranelles, number Cirri on frontal area, number b Pretransverse ventral cirri, number Transverse cirri, number Right marginal cirri, number Left marginal cirri, number Caudal cirri, number Dorsal kineties, number
pe1 pe2 pe1 pe2 pe1 pe2 pe1 pe1 pe2 pe1 pe2 pe1 pe2 pe1 pe1 pe1 pe2 pe1 pe2 pe1 pe2 pe1 pe2 pe1 pe2
99.7 184.0 30.6 56.1 49.3 91.4 4.9 14.4 29.2 2.0 2.0 41.8 58.5 11.0 2.0 5.0 5.0 25.6 28.2 19.4 20.3 3.0 3.0 6.0 6.0
M
SD
SE
CV
Min
Max
n
– – – – – – – – – – – – – – – – – – – – – – – – –
13.0 10.5 7.5 4.6 6.7 7.9 1.4 1.9 1.3 0.0 0.0 3.3 2.0 0.0 0.0 0.0 0.0 2.5 1.7 2.3 1.4 0.0 0.0 0.0 0.0
– 4.0 – 1.8 – 3.0 – – 0.5 – 0.0 – 0.7 – – – 0.0 – 0.7 – 0.5 – 0.0 – 0.0
13.0 80.0 128.0 5.7 171.0 198.0 24.5 20.0 48.0 8.3 50.0 62.0 13.6 40.0 64.0 8.6 78.0 100.0 28.6 4.0 8.0 13.2 9.0 17.0 4.6 27.0 31.0 0.0 2.0 2.0 0.0 2.0 2.0 7.9 36.0 49.0 3.4 56.0 62.0 0.0 11.0 11.0 0.0 2.0 2.0 0.0 5.0 5.0 0.0 5.0 5.0 9.8 21.0 31.0 6.1 25.0 30.0 11.9 16.0 24.0 6.7 19.0 22.0 0.0 3.0 3.0 0.0 3.0 3.0 0.0 6.0 6.0 0.0 6.0 6.0
26 7 26 7 26 7 26 27 7 26 7 25 7 26 26 21 7 21 7 21 7 19 7 26 7
a
All measurements in µm. CV = coefficient of variation in %, M = median, Max = maximum value, mean = arithmetic mean, Min = minimum value, n = number of individuals investigated, SD = standard deviation, SE = standard error of arithmetic mean. Data based on protargol-impregnated specimens. b
In the neotype population, all cirri (except for marginal cirri) ahead of level of buccal vertex are included, that is, three frontal cirri, one buccal cirrus, four frontoventral cirri, and three postoral ventral cirri. The population described by Xu & Song (1999) has the same number, but they obviously counted the rearmost postoral ventral cirrus as pretransverse ventral cirrus.
Insufficient redescriptions Trachelostyla caudata Kahl – Petran, 1967, Ecol. Mar., 2: 188, Planşa V, Fig. D (Fig. 107b). Remarks: The cirral pattern (e.g., postoral row present) and the single (misobservation?) macronuclear nodule (vs. about 10 in original description) indicate a misidentification. According to the general appearance one cannot exclude that Petran (1967) observed a Stichotricha species. Psammobiotic at beach of Vama Veche, Bulgaria, Black Sea (see also Petran 1971, p. 154).
502
SYSTEMATIC SECTION
Trachelostyla caudata Kahl 1932 – AL-Rasheid, 2001, Tropical Zoology, 14: 140, Fig. 10, 44 (Fig. 107a). Remarks: The description and the illustration and micrograph lack the characteristic narrowed posterior body portion (tail) strongly indicating that the identification is incorrect. It can be excluded that the specimen is squeezed because the narrowed anterior portion (head) is well preserved. In addition, it has many more macronuclear nodules (about 45) than T. caudata (around 11) or T. pediculiformis (9–17 in neotype population). The illustration is somewhat confusing because the cirral pattern is shown as seen from the dorsal side. ALRasheid characterised the population as follows: “Hypotrich; thigmotactic, elongate rectangular, 80–120 × 20–35 µm. Anterior 1/3 attenuated, posteriorly broadly rounded, flexible, dorsoventrally flattened. AZM extending along left margin over 1/2 to 1/3 of body length, composed of about 40 thick membranelles. Paroral membrane short, on right of oral cavity. Frontoventral cirri four, two rows of marginals, oblique row of long five transverse cirri. Dorsal cirri fine, long. Macronuclei many, 3–5 µm in diameter, micronuclei two.” Found in sediment of sandy beaches (salinity 35‰) of the Farasan Islands, Red Sea.
Spirotrachelostyla Gong, Song, Li, Shao & Chen, 2006 2006 Spirotrachelostyla nov. gen.1 – Gong, Song, Li, Shao & Chen, Europ. J. Protistol., 42: 72 (original description). Type species (by original designation): Trachelostyla spiralis Dragesco & DragescoKernéis, 1986.
Nomenclature: Spirotrachelostyla is a composite of he speira (Greek noun; spiral, helix) and the genus-group name Trachelostyla (see there for derivation) and refers to the spirally twisted body. Has, like Trachelostyla, feminine gender (Gong et al. 2006). Characterisation (A = supposed apomorphy): Trachelostylidae with spindleshaped and twisted body (A). Additional characters: Only one of three species is described in detail; thus the list of additional characters is rather short and uncertain: body likely flexible; not distinctly flattened dorsoventrally; contractile vacuole lacking?; cortical granules likely lacking; dorsal bristles likely long (>5 µm). Remarks: The diagnosis of Spirotrachelostyla (see corresponding footnote) largely follows the description of S. tani. For example, Gong et al. (2006) mention “about 13 cirri scattered on the anterior peristomial region” although Hu & Song (2002) used the feature 13 vs. 5 cirri to separate S. tani from the very similar S. simplex. Further, Gong et al. (2006) characterised Spirotrachelostyla species as “generally sessile forms in a lorica”, although no lorica is described any of the three spe1
Gong et al. (2006) provided the following diagnosis: Spirally twisted trachelostylids with spindle-shaped body; about 13 cirri scattered on anterior peristomal region; two pretransverse ventral cirri present or absent; usually five transverse cirri; one left and one right row of marginal cirri not confluent posteriorly; caudal cirri present. Generally sessile forms in a lorica.
Spirotrachelostyla
503
cies included in this (Kahl 1932, Dragesco & Dragesco-Kernéis 1986, Hu & Song 2002). The general appearance of Spirotrachelostyla and especially the lack of cirri in the postoral area (except two small cirri in type species!) and the trachelostylid oral apparatus indicate a close relationship to Trachelostyla. According to Gong et al. (2006), Trachelostyla is related to Gonostomum Sterki, 1878; for a more detailed discussion of this topic, see remarks at Trachelostylidae. The main apomorphy of Spirotrachelostyla is obviously the twisted body, a feature which very likely evolved several times independently within the Hypotricha, for example in Spiroamphisiella (Fig. 27a, b, 28e, 29a), Strongylidium Sterki, 1878 (Kahl 1932, p. 551; Paiva & Silva-Neto 2007), and Stichotricha Perty, 1849 (see Foissner et al. 1991, p. 203). Due to the slender, twisted body the cirral pattern looks irregular and is therefore difficult to interpret. However, the general appearance indicates a close relationship to Trachelostyla, which has the same oral apparatus and lives also in the sea. Species included in Spirotrachelostyla (alphabetically arranged basionyms are given): (1) Stichotricha simplex Kahl, 1932; (2) Trachelostyla spiralis Dragesco & Dragesco-Kernéis, 1986; (3) Trachelostyla tani Hu & Song, 2002.
Key to Spirotrachelostyla species Main features for the identification are the number of macronuclear nodules, the body size, and the number of cirri in the frontal area. Thus, protargol impregnation is recommended. 1 16–20 macronuclear nodules (Fig. 104a, b). . Spirotrachelostyla spiralis (p. 503) - Two macronuclear nodules (Fig. 105a, 106a). . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2 Body length 85–120 µm; around eight cirri right of adoral zone (marginal cirri not included; Fig. 106a). . . . . . . . . . . . . . . . . Spirotrachelostyla simplex (p. 509) - Body length 135–210 µm; 13 cirri right of adoral zone (Fig. 105k). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Spirotrachelostyla tani (p. 506)
Spirotrachelostyla spiralis (Dragesco & Dragesco-Kernéis, 1986) Gong, Song, Li, Shao & Chen, 2006 (Fig. 104a–d) 1986 Trachelostyla spiralis n. sp. – Dragesco & Dragesco-Kernéis, Faune Tropicale, 26: 451, Planche 132, Fig. D–F (Fig. 104a–d; original description; slides likely deposited in private collection of J. Dragesco). 2001 Trachelostyla spiralis Dragesco and Dragesco-Kernéis, 1986 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 90 (nomenclator containing all basionyms, combinations, and higher taxa of euplotids and hypotrichs).
504
SYSTEMATIC SECTION
Fig. 104a–d Spirotrachelostyla spiralis (from Dragesco & Dragesco-Kernéis 1986. Protargol impregnation). a: Ventral view, 72 µm. Possibly the body outline is from a live specimen. b–d: Infraciliature of ventral side, b = 49 µm, c = 62 µm, d = 62 µm. Frontal cirri connected by anterior dotted line; frontoventral cirri circled; anteriorly displaced postoral ventral cirri connected by dotted line, and two further postoral cirri marked by arrows (note that these homologisations have to be confirmed by ontogenetic data). AZM = adoral zone of membranelles, BC = buccal cirrus (according to Dragesco & Dragesco-Kernéis 1986), CC = caudal cirri, FC = right frontal cirrus, LMR = left marginal row, MA = macronuclear nodule, P = paroral, PTVC = pretransverse ventral cirri, RMR = right marginal row, TC = right transverse cirrus, III/2 = cirrus behind right frontal cirrus. Page 503.
2002 Trachelostyla spiralis Dragesco & Dragesco-Kernéis, 1986 – Hu & Song, Hydrobiologia, 481: 178, Table 2, Fig. 2j (Fig. 104a; comparison with S. tani and key to Trachelostyla species). 2006 Spirotrachelostyla spiralis (Dragesco & Dragesco-Kernéis, 1986) nov. comb. – Gong, Song, Li, Shao & Chen, Europ. J. Protistol., 42: 72, Fig. 30P, Q (Fig. 104a, b; combination with Spirotrachelostyla).
Spirotrachelostyla
505
Nomenclature: No derivation of the name is given in the original description. The species-group name spiral·is, -is, -e (Greek adjective [m, f, n]; spiral, helical) likely refers to the spiral body. Type species of Spirotrachelostyla. Remarks: The present species differs from the congeners, inter alia, by the higher number of macronuclear nodules (16–20 vs. each two nodules) and the presence of two, admittedly inconspicuous, cirri in the postoral area. Ontogenetic data are needed to know the correct designation of these two “surplus” cirri. Spirotrachelostyla spiralis is described mainly (exclusively?) from protargol-impregnated specimens; thus, a redescription is needed. Morphology: Body size of contracted (likely protargol-impregnated) specimens 50–73 × 13–21 µm, on average 65 × 17 µm (n = 14). Body outline spindle-shaped, that is, anterior and posterior portion narrowed and middle cell portion widest. 16–20 macronuclear nodules in cell centre; individual nodules only about 3 µm across. Micronuclei not observed. Contractile vacuole not observed. Presence/absence of cortical granules and movement not known. Oral apparatus trachelostylid and obviously somewhat spirally. Adoral zone occupies roughly 50% of body length (24–43 µm, on average 36 µm; n = 12), composed of 24–42, on average 40 (n = 10) membranelles. Paroral rather short (about 10 µm), slightly curved and arranged very close to proximal portion of adoral zone, terminates at buccal vertex (Fig. 104b). Cirral pattern as shown in Fig. 104b–d. 10–12 cirri right of adoral zone, exact arrangement difficult to recognise because area rather narrow. Specimen shown in Fig. 104b with 11 cirri in this area; very likely these are the ordinary (slightly enlarged) frontal cirri (3 in number), the buccal cirrus (1), the frontoventral cirri (4), and the postoral ventral cirri (3), which are – as in Gonostomum (see Berger 1999) and Trachelostyla pediculiformis – ahead of the level of the buccal vertex (note, however, that ontogenetic data are needed for a correct interpretation of the pattern). Two small cirri close behind buccal vertex and two small pretransverse ventral cirri ahead of the two rightmost transverse cirri. Five strong transverse cirri arranged in Jshaped pseudorow; project distinctly beyond rear cell end. Marginal rows spirally arranged (Fig. 104a), right row commences about at 50% of length of adoral zone, terminates ahead of right transverse cirrus. Left row begins left of proximal portion of adoral zone, ends subterminally, that is, marginal rows do not overlap posteriorly. Length of dorsal bristles not observed; dorsal kinety pattern not known because it did not impregnate. Two caudal cirri (Fig. 104b). Occurrence and ecology: Saltwater (Patterson et al. 1989, p. 211). Type locality of S. spiralis are saltwater ponds near Lake Nokoué at the city of Cotonou, Benin (Dragesco & Dragesco-Kernéis 1986). Record not substantiated by morphological data: Lake Qarun, a salt lake in the Fayum Oasis, Egypt (Wilbert 1995, p. 282). Food not known.
506
SYSTEMATIC SECTION
Spirotrachelostyla tani (Hu & Song, 2002) Gong, Song, Li, Shao & Chen, 2006 (Fig. 105a–l, Table 31) 2002 Trachelostyla tani nov. spec.1 – Hu & Song, Hydrobiologia, 481: 174, Fig. 1a–l, Table 1 (Fig. 105a–l; original description; the holotype slide [accession number HD-00022201] and a paratype slide [HD-00022202] are deposited in the Laboratory of Protozoology, Ocean University of Qingdao, China). 2003 Trachelostyla tani Hu & Song, 2002 – Hu, Gong & Song, Pathogenic protozoa, p. 173, Fig. 5-9I–L (Fig. 105a, c, j, l; review about pathogenic ciliates). 2006 Spirotrachelostyla tani (Hu & Song, 2002) nov. comb. – Gong, Song, Li, Shao & Chen, Europ. J. Protistol., 42: 72, Fig. 30S–U (Fig. 105a, k, l; combination with Spirotrachelostyla).
Nomenclature: Hu & Song (2002) dedicated this species to Zhiyuan Tan, Institute of Oceanology, Chinese Academy of Sciences. Remarks: The present species differs from S. simplex by its body size (135–210 µm vs. 85–120 µm), the number of cirri on frontal area (13 vs. about 8 in specimen illustrated [value must not be over-interpreted because difficult to estimate in life]), and the prominence of the transverse cirri (rather inconspicuous vs. distinct). Perhaps a redescription of S. simplex will provide further, more convincing differences. Possibly, a subspecies classification would be more appropriate. If you are unable to decide between S. simplex and S. tani (e.g., because only few specimens available), then write S. simplex-group. Cossothigma dubium (Fig. 76a–d) and Spiroamphisiella hembergeri (Fig. 27a–i, 28a–k, 29a–n) have a very similar habitus, including the nuclear apparatus. However, these species have distinct frontoventral rows so that a confusion with S. tani is very unlikely. Morphology: Body size 135–210 × 25–35 µm in life. Body outline elongate spindle-shaped, that is, anterior and posterior body portion distinctly narrowed, middle portion widest; strongly twisted. Body conspicuously flexible, but not contractile; dorsoventrally not distinctly flattened, elliptical in cross section (Fig. 105a–c). Invariably two macronuclear nodules in widest body portion; individual nodules ovoid to long ellipsoidal, about 20 × 8 µm in protargol preparations. 2–5 micronuclei attached to macronuclear nodules, difficult to recognise in life (Fig. 105h, i, Table 31). Pellicle thin, no cortical granules recognisable. Contractile vacuole neither mentioned nor illustrated, indicating that this organelle is lacking. Cytoplasm dark greyish and opaque, contains numerous granules less than 2 µm across, crystals of ordinary shape, and food vacuoles (Fig. 105a, d). Movement moderately fast, swims forward by rotation about main body axis or crawls on substrate; during crawling most marginal cirri do not move. 1
Hu & Song (2002) provided the following diagnosis: Spirally twisted marine Trachelostyla with long and narrowed anterior portion, in vivo 135–210 × 25–35 µm; about 80 adoral membranelles, 13 frontoventral and 5 transverse cirri; marginal cirral row highly spiral, posteriorly influent; two dorsal kineties and 2 caudal cirri, constantly 2 macronuclear nodules.
Spirotrachelostyla
507
Table 31 Morphometric data on Spirotrachelostyla tani (from Hu & Song 2002) Characteristics a
mean
M
SD
SE
CV
Max
n
Body, length Body, width Adoral zone of membranelles, length Adoral membranelles, number Cirri on frontal area, number b Transverse cirri, number Left marginal cirri, number Right marginal cirri, number Macronuclear nodules, number Macronuclear nodules, length Macronuclear nodules, width Micronuclei, number Dorsal kineties, number
184.8 27.9 85.6 80.9 13.0 5.0 57.5 73.5 2.0 20.4 8.1 2.9 2.0
– – – – – – – – – – – – –
21.1 3.7 9.4 9.5 0.0 0.0 6.7 7.5 0.0 3.7 2.2 0.9 0.0
6.1 1.1 2.7 2.7 0.0 0.0 2.0 2.3 0.0 1.1 0.6 0.3 0.0
11.4 152.0 216.0 13.4 23.0 35.0 11.0 72.0 99.0 11.7 65.0 94.0 0.0 13.0 13.0 0.0 5.0 5.0 11.6 42.0 65.0 10.2 60.0 87.0 0.0 2.0 2.0 17.9 16.0 28.0 30.9 4.0 13.0 30.9 2.0 5.0 0.0 2.0 2.0
12 12 12 12 12 12 11 11 12 12 12 12 12
Min
a All measurements in µm. CV = coefficient of variation in %, M = median, Max = maximum value, mean = arithmetic mean, Min = minimum value, n = number of individuals investigated, SD = standard deviation, SE = standard error of arithmetic mean. Data based on protargol-impregnated specimens. b
Marginal cirri not included.
Oral apparatus more or less trachelostylid; adoral zone occupies about 45% of body length in protargol preparations and in life (Table 31, Fig. 105a), composed of 81 membranelles on average. Distalmost four membranelles somewhat enlarged (cilia about 15 µm long) and separated from remaining membranelles by gap, look like frontal cirri in life; major part of zone extends along left margin of narrowed anterior body portion, proximal portion distinctly curved and in buccal cavity (Fig. 105j, k). Cirral pattern rather constant, however, difficult to interpret because of the narrowed anterior body portion and the twisting of the cell (Fig. 105e, f, g–l, Table 31). Invariably 13 cirri on frontal area; cirri of rather different size at base, but all about 10 µm long; arranged as shown in Fig. 105k. Three frontal cirri with right one somewhat separated from the other two; two small cirri behind right frontal cirrus (Fig. 105j). Most cirri cannot be homologised with other cirral groups (frontoventral, buccal, postoral) of 18-cirri hypotrichs without ontogenetic study. No cirri on postoral area and ahead of the five transverse cirri1. Transverse cirri arranged in J-shaped pattern, about 16 µm long and thus project by about half of their length beyond rear cell end. Right marginal row commences near anterior body end, spirals to right transverse cirrus; left row commences slightly ahead of level of buccal vertex, spi1
The 18-cirri hypotrichs also have 13 frontal-ventral cirri, namely, three frontal cirri, four frontoventral cirri, one buccal cirrus, three postoral ventral cirri, and two pretransverse ventral cirri (details for designation see Fig. 6a in Berger 1999). Possibly this agreement in the number is purely by chance, but I strongly suggest that the 13, respectively, 18 cirri of S. tani are homologous to that of the 18-cirri oxytrichids, although it is difficult to imagine that the two pretransverse ventral cirri have moved so far anteriorly.
508
SYSTEMATIC SECTION
Spirotrachelostyla
509
rals to near rear cell end. Both rows arranged in a furrow clearly recognisable only in life; bases of marginal cirri rather long. Dorsal bristles 7–8 µm long, arranged in two kineties of body length. Each kinety with one caudal cirrus; however, caudal cirri very difficult to recognise in life (Fig. 105e, f). Occurrence and ecology: Marine. Type locality of S. tani is the coast of Qingdao (Tsingtao [36°08'N, 120°43'E], China), Yellow Sea, where Hu & Song (2002) discovered it in the mantle cavity of the scallop Chlamys farrei. Thus, Hu et al. (2003) mentioned it in their review on pathogenic ciliates. No further records published. Spirotrachelostyla tani feeds on bacteria, diatoms, and other algae. Hu & Song (2002) provided some ecological data: water temperature 3.0–5.7 °C, pH 8.3–8.6, salinity 34–36‰. Infection frequency is almost 100% when the shell of the host is 2–4 cm large; infection density is about 50–100 specimens per host (Hu & Song 2002).
Spirotrachelostyla simplex (Kahl, 1932) Gong, Song, Li, Shao & Chen, 2006 (Fig. 106a, b) 1932 Stichotricha simplex spec. n. – Kahl, Tierwelt Dtl., 25: 559, Fig. 9724 (Fig. 106a; original description; no formal diagnosis provided and no type material available). 1949 Stichotricha simplex – Froud, Jl. microsc. Soc., 90: 149, Fig. 216 (tiny redrawing of Fig. 106a; comparison of Stichotricha species). 1972 Trachelostyla simplex (Kahl, 1932) n. comb. – Borror, J. Protozool., 19: 15 (combination with Trachelostyla). 1982 Trachelostyla simplex (Kahl, 1932) Borror, 1972 – Hemberger, Dissertation, p. 262 (revision of hypotrichs). 1992 Stichotricha simplex Kahl, 1930–5 – Carey, Marine interstitial ciliates, p. 175, Fig. 687 (Fig. 106b; guide). 2001 Trachelostyla simplex (Kahl, 1932) Borror, 1972 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 81 (nomenclator containing all basionyms, combinations, and higher taxa of euplotids and hypotrichs). 2002 Trachelostyla simplex (Kahl, 1932) – Hu & Song, Hydrobiologia, 481: 176, 178, Table 2, Fig. 2k (redrawing of Fig. 106a; comparison with S. tani and key to Trachelostyla species).
b
Fig. 105a–l Spirotrachelostyla tani (from Hu & Song 2002. a–d, from life; e–l, protargol impregnation). a: Ventral view of a representative specimen, individual size not indicated (range 135–210 µm). b: Right lateral view. c: Slender specimen showing twisting. d: Cytoplasmic crystals. e, f: Infraciliature of rear body portion of same specimen, 69 µm. g: Infraciliature of ventral side, 173 µm. h, i: Nuclear apparatus, h = 142 µm, i = 160 µm. j–l: Infraciliature of oral region and ventral and dorsal side, 193 µm. Arrow in (j) marks inconspicuous gap in adoral zone. Note that the cirral pattern is very difficult to interpret because of the narrow, twisted anterior body portion. AZM = adoral zone of membranelles, CC = caudal cirri, FC = right frontal cirrus, LMR = left marginal row, MA = macronuclear nodule, MI = micronucleus, RMR = right marginal row, TC = transverse cirri, 1, 2 = dorsal kineties. Page 506.
510
SYSTEMATIC SECTION
2006 Spirotrachelostyla simplex (Kahl, 1932) nov. comb. – Gong, Song, Li, Shao & Chen, Europ. J. Protistol., 42: 72, Fig. 30O (Fig. 106a; combination with Spirotrachelostyla).
Fig. 106a, b Spirotrachelostyla simplex (a, from Kahl 1932; b, after Kahl 1932 from Carey 1992. From life). Ventral view, 85–120 µm. Oblique arrows mark anterior and posterior end of row of widely spaced cirri, transverse arrows mark long cirri at rear cell end (correct designation [transverse cirri? caudal cirri? marginal cirri?] only possibly after detailed redescription). AZM = adoral zone of membranelles, DB = dorsal bristles (erroneously illustrated as cirri in Carey’s redrawing), LMR = left marginal row, MA = macronuclear nodule, RMR = right marginal row. Page 509. 1
Nomenclature: No derivation of the name is given in the original description. The species-group name simplex (Latin; simple) likely refers to the reduced number of cirral rows because the two ventral rows present in Stichotricha are lacking in the present species. Incorrect subsequent spellings: Strichotricha simplex Kahl (Detcheva 1982, p. 249); Stychotricha simplex Kahl (Detcheva 1980, p. 34). Remarks: The present species is described only after live observation (Kahl 1932). Thus, a detailed redescription is needed. However, the discovery of S. tani shows that the cirral pattern described by Kahl (1932) is not a misobservation. Kahl (1932) classified S. simplex in Stichotricha Perty, 1849 because of the habitus. Borror (1972) transferred it to Trachelostyla, likely because of the lack of ventral rows. Recently, Gong et al. (2006) assigned it to Spirotrachelostyla because of the spiralling of the cell and the marginal rows. For a comparison of S. simplex and S. tani, see S. tani. Cossothigma (Fig. 76a–d) and Spiroamphisiella (Fig. 27a–i) have a similar habitus, but posses one distinct frontoventral row; Stichotricha species usually have two such rows (e. g., Foissner et al. 1991). Morphology: The description below is exclusively based on the text and the illustration of the original description. However, note that details of the ventral and dorsal infraciliature must not be over-interpreted because Kahl did not have the advantage of silver preparations. Body length 85–120 µm, body length:width ratio of specimen illustrated about 5.8:1 (Fig. 106a). Body outline slender spindle-shaped; anterior portion distinctly set off only in well-fed specimens. Body likely supple and slightly contractile (Kahl 1932, p. 557). Two macronuclear nodules (Fig. 106a). Contractile vacuole1 and cortical granules neither mentioned nor illustrated. No case observed.
Kahl (1932, p. 557) mentions in the genus section of Stichotricha that the contractile vacuole is in the ordinary position, that is, left behind the buccal vertex. However, I am uncertain whether or not this statement also refers to the present species.
Spirotrachelostyla
511
Oral apparatus roughly trachelostylid. Adoral zone occupies about 33% of body length, frontal-most three membranelles whirling (Fig. 106a). Cirral pattern not known in detail, but more or less as in S. tani. Description given by Kahl (1932) does not correspond very well with his illustration; according to Fig. 106a, likely three frontal cirri, two buccal cirri, and some (three in Fig. 106a) cirri behind right frontal cirrus. Cirri of marginal rows long and thin, especially in posterior portion. Right marginal row very likely begins distinctly ahead of level of buccal vertex, extends spirally onto left dorsolateral side posteriorly. Left row commences near buccal vertex, extends spirally onto dorsal surface and likely terminates on right side of cell. Some relatively long cirri at posterior end of cell; correct designation (marginal? transverse? caudal?) needs redescription and ontogenetic data (Fig. 106a). Dorsal bristles about 7 µm long; number and arrangement of kineties not mentioned in the original description (Kahl Fig. 107a, b Insufficient redescriptions. a: Trachelostyla caudata (from AL-Rasheid 2001), cir1932, p. 559); however, in the genus sec- ral pattern as seen from dorsal side from life, tion on page 557 Kahl writes that all “Sti- 94 µm; p. 502. b: Trachelostyla caudata (from Petran 1967), ventral(?) view, 150–200 µm; p. chotricha” species have three kineties. Occurrence and ecology: Marine 501. ACR = amphisiellid median cirral row, DB = dorsal bristles. and brackish (e.g., Patterson et al. 1989, p. 210). Type locality of Spirotrachelostyla simplex is the North Sea, where it was common, but never abundant at the islands of Heligoland and Sylt (Kahl 1932). Records not substantiated by morphological data: brackish bay at the Bottsand, Kieler Außenförde (Germany), Baltic Sea (Ax & Ax 1960, p. 12); mesopsammon (23 to 24°C, 6–17‰ salinity) of Black Sea at the village of Kitène, Romania (Detcheva 1980, p. 34; 1982, p. 249); sediments of Loch Eil, west coast of Scotland (Wyatt & Pearson 1982, p. 301); west coast of Caspian Sea (Agamaliev 1971, p. 384). Food not known. In experiments Ax & Ax (1960) found a salinity tolerance from 15–40‰, that is, Spirotrachelostyla simplex is likely euryhaline tolerating a range from 6–40‰ (see records above).
512
SYSTEMATIC SECTION
Taxa not Considered in the Trachelostylidae The following taxa, included in the trachelostylids by some workers, lack the main features of this group and are therefore not treated in this taxon. Gonostomum Sterki, 1878, Z. wiss. Zool., 31: 57. Type species (by original designation): Oxytricha affinis Stein, 1859. Remarks: Classified in the Trachelostylidae by Lynn & Small (2002). Gonostomum includes species mainly occurring in terrestrial habitats and with a non-narrowed anterior body portion. However, the shape of the adoral zone and the undulating membranes as well as the lack of dorsomarginal rows and kinety fragmentation are indeed reminiscent of Trachelostyla so that one cannot exclude that Gonostomum and Trachelostyla are closely related. However, since Gonostomum lacks the head and does not occur in the sea, I do not include it in the Trachelostylidae. A revision of Gonostomum was provided by Berger (1999, p. 367), where I assumed that its dorsal infraciliature was formed via a simplification process from the complicated Oxytricha pattern (kinety fragmentation and dorsomarginal kineties present). Hemisincirra Hemberger, 1985, Arch. Protistenk., 130: 408. Type species (by original designation): Uroleptus kahli, Buitkamp, 1977. Remarks: Classified in the Trachelostylidae by Lynn & Small (2002). Hemisincirra is a heterogenous group containing only terrestrial species which lack the trachelostylid oral apparatus. Preliminarily assigned to the amphisiellids (p. 387). Lamtostyla Buitkamp, 1977, Acta Protozool., 16: 270. Type species (by original designation): Lamtostyla lamottei Buitkamp, 1977. Remarks: Classified in the Trachelostylidae by Lynn & Small (2002). Lamtostyla comprises terrestrial species with a (usually) distinct amphisiellid median cirral row (p. 161). A trachelostylid oral apparatus is lacking, strongly indicating that a classification in the Trachelostylidae is incorrect. Psammomitra Borror, 1972, J. Protozool., 19: 8, 15. Type species (by monotypy): Mitra radiosa Quennerstedt, 1867. Remarks: Classified in the Trachelostylidae by Small & Lynn (1985). Now assigned to the Urostyloidea because a midventral complex is present (Song & Warren 1996; for review, see Berger 2006, p. 221). Consequently, the narrowed anterior body portion in Psammomitra and the trachelostylids has to be interpreted as convergence. Terricirra Berger & Foissner, 1989, Bull. Br. Mus. nat. Hist. (Zool.), 55: 35. Type species (by original designation): Perisincirra viridis Foissner, 1982. Remarks: Classified in the Trachelostylidae by Lynn & Small (2002). Terricirra comprises terrestrial species which do not have the trachelostylid oral apparatus. It is preliminarily assigned to the amphisiellids (p. 447).
Taxa not Considered in the Trachelostylidae
513
Urosoma Kowalewskiego, 1882, Pam. fizyogr., 2: 406. Type species (by monotypy): Urosoma cienkowskii Kowalewskiego, 1882. Remarks: Classified in the Trachelostylidae by Small & Lynn (1985). Urosoma comprises mainly terrestrial species which have a dorsomarginal kinety, but lack a kinety fragmentation, indicating that Urosoma belongs to the non-oxytrichid Dorsomarginalia. I discussed Urosoma in the review on oxytrichids (Berger 1999, p. 396). Urosomoida Hemberger in Foissner, 1982, Arch. Protistenk., Arch. Protistenk., 126: 115. Type species (by original designation): Uroleptus agilis Engelmann, 1862. Remarks. Classified in the Trachelostylidae by Small & Lynn (1985). Incorrectly spelled Urosomoides by Small & Lynn (1985, p. 461). Urosomoida comprises mainly terrestrial species which have a dorsomarginal kinety, but lack a kinety fragmentation, indicating that it belongs to the non-oxytrichid Dorsomarginalia. For review, see Berger (1999, p. 345).
Taxa of Unknown Position in the Hypotricha The phylogenetic positions of Apourosomoida and Bistichella are uncertain. Both lack dorsomarginal kineties and oxytrichid kinety fragmentation, strongly indicating that they do not belong to the dorsomarginalian part of the Hypotricha (Fig. 9a). Thus, I treat them as taxa of unknown position in the Hypotricha. For more detailed discussions about this topic, see remarks at genus sections.
Apourosomoida Foissner, Agatha & Berger, 2002 2002 Apourosomoida nov. gen.1 – Foissner, Agatha & Berger, Denisia, 5: 759 (original description). Type species (by original designation): Apourosomoida halophila Foissner, Agatha & Berger, 2002.
Nomenclature: Apourosomoida is a composite of the Greek prefix apo- (derived from) and the genus-group name Urosomoida (Foissner et al. 2002). No derivation of the genus-group name Urosomoida is given in the original description and in Berger (1999); it is a composite of the genus-group name Urosoma (tailed body), the thematic vowel ·o-, and the suffix -ida (similar) referring to a similarity with Urosoma Kowalewskiego, 1882. Apourosomoida has, like Urosomoida, feminine gender (Foissner et al. 2002). Characterisation (A = supposed apomorphy): Adoral zone of membranelles with gap at left anterior corner of body. Undulating membranes roughly in Gonostomum pattern. Three frontal cirri. Buccal cirrus present. Usually four frontoventral cirri in L-shaped pattern. Postoral cirral row present, formed only by anlage IV (A). Pretransverse ventral cirri lost and number of transverse cirri strongly reduced. One left and one right marginal row. Two dorsal kineties. Caudal cirri present. Frontalventral-transverse cirri anlage V lost (A?). Anlagen IV and VI are primary primordia. Dorsal kinety formation in Apourosomoida pattern2 (A). Highly saline habitat. Additional characters: Body slender, flexible; two macronuclear nodules; no contractile vacuole recognisable; cortical granules likely lacking; cytoplasm colourless; distributed in southern hemisphere. Remarks: The characterisation above is more or less exclusively based on the type species because the second species included has not yet been described in sufficient detail. We assigned Apourosomoida to the Oxytrichidae because the ventral 1 Foissner et al. (2002) provided the following diagnosis: Oxytrichidae Ehrenberg, 1838 with adoral zone formed like a question mark and interrupted by a minute gap at left anterior corner of body. Undulating membranes in Gonostomum pattern. Frontoventral cirri in V-shaped pattern; originate from five anlagen, of which the rightmost are primary primordia. Number of postoral cirri increased to a conspicuous row, originating from anlage IV. Single transverse cirrus produced by rightmost anlage V. 1 right and 1 left row of marginal cirri. 2 dorsal kineties, each with a caudal cirrus; proter kineties originate from ordinary anlagen, while opisthe’s kinety 1 is generated by posterior fragmentation of kinety. 2 For detailed description of this pattern, see cell division of A. halophila (Note: a dorsomarginal kinety and an oxytrichid kinety fragmentation are lacking!).
514
Apourosomoida
515
ciliature originates basically as in many members of this group (Foissner et al. 2002). However, we also discussed that it might be a representative of an own group, as indicated by the specific formation of the postoral cirral row and dorsal kineties. The similarities in cell division of Apourosomoida and many oxytrichids are very likely plesiomorphies resulting from the 18-cirri pattern already present in the last common ancestor of the hypotrichs. Now I suppose that Apourosomoida is not an oxytrichid because it lacks an oxytrichid dorsal kinety fragmentation, the main morphological apomorphy of this group (Berger 2006, p. 33; note that the fragmentation occurring in A. halophila is [very likely] not homologous to the fragmentation characterising the Oxytrichidae [for details, see cell division at A. halophila]). It probably branched off rather basally within the hypotrichs because it even lacks – like, for example, the urostyloids – a dorsomarginal kinety, that is, it is very likely not even a member of the Dorsomarginalia. The lack of frontal-ventral-transverse cirri anlage V in A. halophila, type of Apourosomoida, is strongly reminiscent of Vermioxytricha (p. 596), Erimophrya (p. 577), and Hemiurosoma (p. 614). However, these three taxa have dorsomarginal kineties, indicating that Apourosomoida is not closely related to them. Consequently, we have to assume that anlage V has been lost twice or several times independently (Foissner et al. 2002). The lack of a dorsomarginal kinety separates Apourosomoida not only from the genera mentioned above, but also from other seemingly similar taxa, for example Urosomoida (e.g., six frontal-ventral-transverse cirri anlagen, adoral zone without gap, four dorsal kineties; for review, see Berger 1999, p. 345). Apourosomoida produces (at least) the two rightmost frontal-ventral-transverse cirri anlagen via primary primordia (Fig. 110d), a feature also present in Urosoma and Gonostomum (Berger & Foissner 1997, Berger 1999). However, Urosoma has, inter alia, a dorsomarginal kinety, and in Gonostomum the dorsal kinety formation proceeds likely in the plesiomorphic way, namely, two anlagen are formed within each kinety. In the original description of Apourosomoida we compared the type species, inter alia, with Uroleptus natronophilus, a small species only known from an African soda lake. Its cirral pattern is similar to that of A. halophila so that it is transferred from Uroleptus – where it is certainly misplaced – to the present genus (details see A. natronophila). Urosomoida minima Hemberger, 1985 might also belong to Apourosomoida, as indicated by the bipartite adoral zone and the dorsal kinety pattern, which, however, is not illustrated and described in detail (Foissner et al. 2002; for review see Berger 1999, p. 366). However, Urosomoida minima has only three postoral ventral cirri and the distance between the first and middle cirrus is smaller than that between the middle and last (rearmost) cirrus. Both features are reminiscent of ordinary 18-cirri hypotrichs and because of the lack of ontogenetic data – which would prove or disprove the relationship to Apourosomoida – and the non-saline, European type locality, its original generic assignment is retained. However, to simplify identification, it is included in the key below.
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SYSTEMATIC SECTION
For a discussion of Cladotricha variabilis Ruinen, 1938, see A. halophila. Cladotricha variabilis sensu Borror & Evans (1979), which also has a postoral cirral row, has the ordinary number of six (I–VI) frontal-ventral cirri anlagen (vs. anlage V lacking in present genus) and very likely does not form transverse cirri, showing that it is not an Apourosomoida species. Species included in Apourosomoida: (1) Apourosomoida halophila Foissner, Agatha & Berger, 2002; (2) Uroleptus natronophilus Dietz, 1965.
Key to Apourosomoida species and similar species Note that A. natronophila is described only from live specimens, that is, the cirral pattern and the dorsal infraciliature are not known in detail. The key also includes Urosomoida minima (see remarks above) and Erimophrya quadrinucleata which has, like Apourosomoida species, an increased number of postoral ventral cirri arranged in line. 1 Two macronuclear nodules present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 - Four macronuclear nodules (Fig. 124c, d, 125a–g). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Erimophrya quadrinucleata (p. 595) 2 Body length in life 60–130 µm (e.g., Fig. 108a–f). . . . . . . . . . . . . . . . . . . . . . . . 3 - Body length in life(?) 35–50 µm (Fig. 112a, b). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apourosomoida natronophila (p. 530) 3 Right marginal row commences about at level of frontal cirri; 3–6, usually 5 postoral ventral cirri (Fig. 108e, f). . . . . . . . . . Apourosomoida halophila (p. 516) - Right marginal row commences about at level of rearmost frontoventral cirrus; 3 postoral ventral cirri (Fig. 110r in Berger 1999, p. 351). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Urosomoida minima (see Berger 1999, p. 366)
Apourosomoida halophila Foissner, Agatha & Berger, 2002 (Fig. 108a–f, 109a–f, 110a–v, Table 32) 2002 Apourosomoida halophila nov. spec.1 – Foissner, Agatha & Berger, Denisia, 5: 761, Fig. 166a–f, 167a–v, 398a–j, Table 147 (Fig. 108a–f, 109a–f, 110a–v; original description; the holotype slide [accession number 2002/329] and two paratype slides [2002/330, 331] have been deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; see also Aescht 2003, p. 387).
1
Foissner et al. (2002) provided the following diagnosis: Size about 100 × 20 µm in vivo; elongate ellipsoidal with pointed posterior end. 2 elongate ellipsoidal macronuclear nodules and 1 micronucleus. Adoral zone usually composed of 17 membranelles occupying 20% of body length. On average about 19 cirri each in right and left marginal row, 3 frontal cirri, 4 frontoventral cirri, 1 buccal and transverse cirrus each, 5 cirri in postoral row, and 2 caudal cirri. Dorsal kineties 1 and 2 composed of an average of 2, respectively, 8 bristles.
Apourosomoida
517
Nomenclature: The species-group name halophil·us, -a, -um (Latin adjective [m; f; n]; thriving in saline habitats) is a composite of the Greek words halós (salt) and philos (preferring), referring to the saline environments the species prefers (Foissner et al. 2002). Type species of Apourosomoida. Remarks: Apourosomoida halophila is well defined via the row of postoral ventral cirri originating from anlage IV only. The 18-cirri hypotrichs invariably have three postoral ventral cirri, which are produced by anlagen IV (one cirrus) and V (two cirri). There are only very few species which have a similar row. Erimophrya quadrinucleata has three longitudinally arranged postoral ventral cirri, but, inter alia, four macronuclear nodules and a distinct cortical crystalline reticulum (Fig. 124c, d, 125a–g). Apourosomoida natronophila is distinctly smaller and in Urosomoida minima the right marginal row is distinctly shortened anteriorly (see key for details). Ruinen (1938) discovered Cladotricha variabilis in highly saline waters from Australia (Fig. 111a–g). Probably, Ruinen mixed two species and therefore we fixed the quadrinucleate specimen shown in Fig. 111a as C. variabilis (Foissner et al. 2002, p. 771). Another specimen illustrated by Ruinen (1938) is possibly identical with A. halophila, as indicated by the body size and shape, the cirral pattern, and the highly saline habitat (Fig. 111d). However, the dorsal kinety pattern of the Australian specimen is not known so we cannot be certain whether the African and Australian populations are identical. Anyhow, the data show that species living in highly saline habitats of Africa and Australia are very similar or even identical. Cladotricha Gaievskaia, 1925 is an insufficiently defined group of hypotrichs, which, at present, is very likely not monophyletic (for brief comment see chapter Taxa not considered). Morphology: Body size 60–130 × 15–25 µm in life, usually around 100 × 20 µm, considerably shrunken and stouter in protargol and scanning electron microscopy preparations, namely, 71 × 21 µm, as also indicated by the high coefficient of variation (25%; Table 32); body length:width ratio 3.4:1 on average in protargol preparations (Table 32). Body shape rather constant, elongate ellipsoidal to bluntly cylindrical with anterior end narrowly rounded and posterior rather abruptly pointed; rarely slipper-shaped or curved specimens occur. Slightly to distinctly twisted along main body axis and flattened up to 2:1 dorsoventrally in oral area (Fig. 108a–c, e, 109a, c–e). Macronuclear nodules in middle third of cell left of midline, usually close together and connected by a fine thread, ellipsoidal to cylindrical (7:1), on average elongate ellipsoidal (3:1); contain many small, globular chromatin bodies. Micronucleus lacking or not impregnated in about 30% of specimens, others have one or two micronuclei attached to the macronuclear nodules at variable positions. No contractile vacuole recognisable. Cortex very flexible, lacks specific granules. Cytoplasm colourless, contains up to 7 µm-sized food vacuoles with bacterial remnants, some 3–5 µm long, ordinary crystals, and pale lipid droplets up to 3 µm across mainly in posterior body half. Glides and swims slowly on microscope slide.
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SYSTEMATIC SECTION
Fig. 108a–f Apourosomoida halophila (from Foissner et al. 2002. a–c, from life; d–f, protargol impregnation). a: Ventral view of a representative specimen, 96 µm. The paroral cilia increase in length from 2 µm anteriorly to 5 µm posteriorly. b: Slipper-shaped specimen showing details of oral area. c: Slender specimen. d: Infraciliature of dorsal side. Arrow marks anterior end of dorsal kinety 2 which, in some specimens, seemingly forms a third row of dorsal bristles (see f). e, f: Infraciliature of ventral and dorsal side and nuclear apparatus of holotype specimen, 75 µm. Arrowhead in (e) marks gap in adoral zone, short arrow de-
Apourosomoida
519
Adoral zone occupies about 20% of body length in life, while 26% on average in protargol preparations due to increased shrinkage of postoral body portion; composed of an average of three minute frontal and 14 ordinary ventral membranelles, separated by a distinct gap at left anterior corner of cell. Buccal field very narrow and flat, covered by an angularly projecting, hyaline lip bearing paroral. Both undulating membranes staggered side by side, composed of a single row of cilia; paroral cilia gradually increase in length from about 2 µm anteriorly to 5 µm posteriorly; endoral cilia extend into pharynx which is supported by fibres of ordinary length and direction (Fig. 108a, b, e, 109a–d, f, 110a, Table 32; see also Fig. 398g, j in Foissner et al. 2002). Cirral pattern and number of cirri of usual variability, except for frontoventral cirri, which vary from 3–6 (Fig. 108a, e, f, 109d–f, 110a, n, Table 32). Cirri usually composed of 4–6 cilia, rarely of two (last cirrus of left marginal row) or nine (frontal cirri). Frontal cirri form oblique pseudorow with left cirrus near gap of adoral zone, and right cirrus, as is usual, behind distal end of adoral zone. Frontoventral cirri arranged in L-shaped pattern, right cirri in flat furrow near right body margin, left cirrus is cirrus III/2. 3–6, usually five, postoral ventral cirri form straight or slightly oblique row in body midline beginning somewhat behind level of buccal vertex. Invariably one transverse cirrus at rear body end, forms rather conspicuous tuft together with caudal cirri and last cirri of left marginal row. Marginal rows obliquely arranged due to body torsion, right row commences subapically on dorsal side and extends onto ventral margin in mid-body to end far subterminally; left row commences left of buccal vertex, extends dorsolaterally in rear body portion and terminates close to cell end; cirri decrease in size and become more widely spaced in posterior third, especially in left row. Dorsal bristles about 2–3 µm long, loosely arranged in two rows (Fig. 108d, f, 109e, 110n, Table 32). Kinety 1 usually composed of a single anterior bristle at level of proximal end of adoral zone and one, rarely two bristles close to corresponding caudal cirrus; rarely an additional bristle occurs in mid-body. Kinety 2 composed of an average of seven bristles forming sigmoidal row in body midline, one or two bristles close to corresponding (= right) caudal cirrus; 1–3 anterior bristles more or less distinctly dislocated, seemingly forming third kinety along anterior portion of right marginal row (see cell division). Right caudal cirrus composed of nine cilia, left of about six. Cell division (Fig. 110b–v; see also Fig. 398h, i in Foissner et al. 2002): Foissner et al. (2002) could study cell division, which is indispensable to interpret the cirral pattern and the dorsal infraciliature of this species correctly. The principles are as
b
notes cirrus III/2, and long arrow marks rear end of postoral ventral row. Frontal cirri connected by dotted line; broken lines connect cirri originating from same anlage. Arrow in (f) marks a “third” dorsal kinety formed by some dislocated bristles of kinety 2 (see d). AZM = adoral zone of membranelles, BC = buccal cirrus, BL = buccal lip, FU = furrow, LMR = rear end of left marginal row, P = paroral, RMR = rear end of right marginal row, TC = transverse cirrus, I, IV, VI = frontal-ventral-transverse cirri anlagen, 1, 2 = dorsal kineties. Page 516.
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SYSTEMATIC SECTION
Fig. 109a–f Apourosomoida halophila (from Foissner et al. 2002. a–c, from life; d–f, scanning electron micrographs). a–c: Ventral views of a slender specimen. Arrows mark food vacuoles. Arrowheads denote gap between ventral and frontal membranelles. d, e: Ventral and dorsal overview. Arrow marks postoral ventral cirri. Arrowhead denotes a dislocated bristle from dorsal kinety 2, seemingly forming a third row of bristles together with the anteriormost bristle of kinety 2. f: Ventral side of anterior body portion. Small arrow marks ridge left of frontoventral cirri. Large arrow denotes gap between frontal and ventral membranelles. Arrowhead marks ontogenetically active cirrus III/2 forming anlage III of proter. Explanation of original labelling: BC = buccal cavity, BL = buccal lip, C = cirri, CC = caudal cirri, CR = cytoplasmic crystals, FC3 = right frontal cirrus (cirrus III/3), FM = frontal membranelles, FVR = frontoventral cirri, LR = left marginal row (d, e; right marginal row in f), MI = micronucleus, P = pharynx, PM = paroral, PO = anteriormost postoral ventral cirrus, RMR = right marginal row, TC = transverse cirrus, VM = ventral membranelles, 1, 2 = dorsal kineties. Page 516.
Apourosomoida 521
Fig. 110a–d Apourosomoida halophila (from Foissner et al. 2002. Protargol impregnation). a: Non-divider (75 µm) which is slightly twisted and thus shows only parts of the marginal rows (dorsal side see 110n). Frontoventral cirri circled. b: Very early divider (69 µm) showing that the oral primordium includes the postoral ventral cirri. c: Early divider (74 µm) showing that cirrus III/2 (short arrow) has modified to anlage III of the proter. The anterior portion of the oral primordium modifies to frontal-ventral-transverse cirri anlagen (long arrow). d: Early divider, 73 µm. The parental buccal cirrus (arrow) has modified to anlage II of the proter. Anlagen IV and VI (and possibly III) are primary primordia. OP = oral primordium, P = paroral, RMR = right marginal row, III, IV, VI frontal-ventral-transverse cirri anlagen, 2 = dorsal kinety 2. Page 516.
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SYSTEMATIC SECTION
Fig. 110e–g Apourosomoida halophila (from Foissner et al. 2002. Protargol impregnation. Parental structures white, new black). e, f: An early-middle and middle divider showing that, as is usual, the parental undulating membranes disorganise and form cirral anlage I (arrow). Note that two anlagen originate within each parental marginal row to produce the new rows. g: Middle-late divider showing cirral segregation within the anlagen as well as beginning migration of the single transverse cirrus (formed by anlage VI) and all cirri (except anteriormost cirrus) of anlage IV, which form the postoral ventral cirri (circled). TC = transverse cirrus of proter, I–IV, VI = frontal-ventral-transverse cirri anlagen. Page 516.
in many 18-cirri hypotrichs so that we described it only briefly, emphasising specific (apomorphic?) features. All parental cirri and dorsal bristles not involved in anlagen formation are resorbed in late dividers and postdividers. The parental buccal field and the undulating membranes are completely reorganised, while the adoral zone is only partially renewed, that is, the short1 ciliary row of the membranelles disappears for a short period in late dividers (Fig. 110i). Apourosomoida halophila has only five frontal-ventral-transverse cirri anlagen (see below). In the original description we designated them as anlagen I–V (Foissner et al. 2002). The rearmost frontoventral cirrus is certainly homologous to cirrus IV/3 of the 18-cirri hypotrichs, so that we can be sure that anlage IV is present. In addi1
Incorrectly designated as “short (leftmost) ciliary row” by Foissner et al. (2002, p. 765); correct is “rightmost”.
Apourosomoida
523
Fig. 110h–j Apourosomoida halophila (from Foissner et al. 2002. Protargol impregnation. Parental structures white, new black). Infraciliature of ventral side of middle and late dividers, h = 57 µm, i = 68 µm, j = 66 µm. Broken lines in opisthe of (i) connect cirri originating from same anlage (note that in this species anlage V is lacking!). Frontal cirri connected by dotted line; frontoventral cirri circled; postoral ventral cirri in rectangle (note that the anteriormost cirrus formed by anlage IV is the rearmost frontoventral cirrus in non-dividers! In the original description we incorrectly assumed that all cirri of anlage IV become postoral ventral cirri). Arrows in (j) mark transverse cirri and broken line connects cirrus IV/3 with postoral ventral cirri which are also formed by anlage IV. The number of cirri within the individual cirral groups is somewhat variable. TC = transverse cirrus of opisthe. Page 516.
tion, Apourosomoida has the ordinary frontoterminal cirri migrating anteriorly and forming two frontoventral cirri in morphostatic specimens. Since the frontoterminal cirri are formed by anlage VI in the 18-cirri hypotrichs, we can conclude that anlage V is lacking in the present species. Therefore, I designate the five anlagen as I–IV and VI. Morphogenesis begins with the formation of an oral primordium along the postoral cirral row, which is soon incorporated into the growing, triangular primordium, producing adoral membranelles very early (Fig. 110b, c). Some scattered basal body pairs remain at the anterior margin of the growing membranellar ribbon and organise to long cirral streaks (primary primordia) extending into the parental frontal field.
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SYSTEMATIC SECTION
Fig. 110k–m Apourosomoida halophila (from Foissner et al. 2002. Protargol impregnation. Parental structures white, new black). k: Infraciliature of ventral side of a very late divider, 59 µm. Arrow marks postoral ventral row of proter. l, m: Infraciliature of ventral side of proter postdividers, l = 43 µm, m = 33 µm. The transverse cirrus is still migrating in the specimen shown in (m) and cell shaping has not yet commenced. TC = transverse cirrus. Page 516.
Simultaneously, the buccal cirrus and cirrus III/2 modify to proter’s anlagen II and III proliferating basal bodies posteriorly; eventually, the anlage formed by cirrus III/2 contacts anlage III of the opisthe. Thus, three long cirral anlagen, so-called primary primordia, are recognisable in early dividers (Fig. 110c, d). Next the primary primordia divide transversely, providing each proter and opisthe with three anlagen (Fig. 110e, f). The proter anlagen originate as follows: anlage I is formed by the reorganising parental undulating membranes; anlage II is the modified parental buccal cirrus (II/2); anlage III is the modified cirrus III/2 plus the anterior portion of opisa third dorsal kinety along the anterior portion of the right marginal row (arrow). o, p: Early dividers showing that the anterior bristle of row 1 forms an anlage (arrow in p), while two anlagen are formed in kinety 2 (asterisks), one for the proter and one for the opisthe. q: A middle divider showing that the new opisthe kinety 1 is generated by fragmentation and migration to the left of the anlage in kinety 2 (arrow). This unique mode is a main feature of Apourosomoida. The macronuclear nodules fuse and the micronucleus commences division. LMR = left marginal row, MA = macronucleus, MI = micronucleus, RMR = right marginal row. Page 516.
d
Apourosomoida Legend continued on p. 524
525
Fig. 110n–q Apourosomoida halophila (from Foissner et al. 2002. Protargol impregnation). Dorsal views of a morphostatic specimens (n) and three dividers (u–q); corresponding ventral views, see Fig. 110a, e, none, f. n: Dorsal kinety 1 consists of two bristles only, one in anterior body half and another close to the left caudal cirrus. Dorsal kinety 2 is composed of an average of seven bristles, of which 1–3 anterior bristles are more or less distinctly dislocated to the right; seemingly forming
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SYSTEMATIC SECTION
Apourosomoida
527
Table 32 Morphometric data on Apourosomoida halophila (from Foissner et al. 2002) Characteristics a
mean
Body, length Body, width Body length:width, ratio Adoral zone of membranelles, length Body length:length of adoral zone, ratio Anterior end to paroral, distance Paroral, length Anterior body end to endoral, distance Endoral, length Anterior body end to buccal cirrus, distance Anterior body end to rearmost frontoventral cirrus, distance Anterior body end to postoral ventral row, distance Anterior body end to end of postoral ventral row, distance Anterior body end to right marginal row, distance Posterior body end to right marginal row, distance Posterior body end to left marginal row, distance Posterior body end to transverse cirrus, distance Anterior body end to first macronuclear nodule, distance Nuclear figure, length Macronuclear nodules, distance in between Anterior macronuclear nodule, length Anterior macronuclear nodule, width Macronuclear nodules, number Micronuclei, length Micronuclei, width Micronuclei, number Frontal adoral membranelles, number Ventral adoral membranelles, number Adoral membranelles, total number Frontal cirri, number Buccal cirri, number Frontoventral cirri, number Postoral ventral cirri, number Transverse cirri, number Right marginal cirri, number Left marginal cirri, number
b
M
SD
SE
CV
Min
Max
n
70.9 20.7 3.4 18.3 3.8 6.2 4.3 8.6 6.7 6.2 10.3
65.0 20.0 3.3 18.0 3.6 6.0 4.0 8.5 7.0 6.0 10.0
17.8 2.9 0.7 1.4 0.8 1.2 0.7 1.4 0.8 1.1 1.3
3.9 0.6 0.1 0.3 0.2 0.3 0.2 0.3 0.2 0.3 0.3
25.1 13.8 20.1 7.4 21.4 19.7 17.3 16.7 11.7 18.4 12.4
52.0 118.0 15.0 25.0 2.4 5.1 15.0 20.0 2.9 5.9 3.0 8.0 3.0 6.0 5.0 10.5 5.0 8.0 3.0 8.0 7.0 13.0
21 21 21 21 21 21 21 21 21 21 21
19.1 29.1
19.0 29.0
1.7 3.6
0.4 0.8
8.9 12.4
16.0 22.0
22.0 36.0
21 21
4.1 7.4 0.2 0.9 17.6
4.0 8.0 0.0 1.0 17.0
1.0 2.4 – – 3.3
0.2 0.5 – – 0.7
25.3 32.3 – – 18.7
2.0 4.0 0.0 0.0 12.0
6.0 13.0 2.0 2.0 27.0
21 21 21 21 21
35.8 2.0 16.6 5.0 2.0 3.3 2.1 0.8 3.0 14.1 17.1 3.0 1.0 4.3 4.9 1.0 19.9 17.7
33.0 2.0 16.0 5.0 2.0 3.0 2.0 1.0 3.0 14.0 17.0 3.0 1.0 4.0 5.0 1.0 20.0 18.0
12.6 1.7 5.0 0.8 0.0 0.6 0.5 – 0.0 0.7 0.7 0.0 0.0 0.6 1.0 0.0 1.9 1.7
2.8 0.4 1.1 0.2 0.0 0.1 0.1 – 0.0 0.2 0.2 0.0 0.0 0.1 0.2 0.0 0.4 0.4
35.4 83.7 30.4 15.5 0.0 18.9 24.5 – 0.0 4.8 3.9 0.0 0.0 15.0 19.8 0.0 9.5 9.7
20.0 0.0 10.0 4.0 2.0 2.0 1.0 0.0 3.0 12.0 15.0 3.0 1.0 3.0 3.0 1.0 16.0 15.0
68.0 5.0 29.0 6.0 2.0 4.5 3.0 2.0 3.0 15.0 18.0 3.0 1.0 6.0 6.0 1.0 25.0 22.0
21 21 21 21 21 21 21 21 21 21 21 21 21 21 21 21 21 21
Fig. 110r–v Apourosomoida halophila (from Foissner et al. 2002. Protargol impregnation). Corresponding ventral views, see Fig. 110g, j–m. Parental structures white, new black. r–t: A middle, late, and very late divider showing patterning of dorsal kineties and formation of a caudal cirrus each at the posterior end. Short arrows mark parental dorsal bristles; long arrows in (t) mark more or less distinctly dislocated bristles at anterior end of kinety 2; asterisks mark anterior bristle of kinety 1. u, v: Proter postdividers showing the new rows of dorsal bristles and the dividing macronucleus. CC = caudal cirri, RMR = right marginal row, TC = transverse cirrus, 1, 2 = dorsal kineties. Page 516.
528
SYSTEMATIC SECTION
Table 32 Continued Characteristics a Dorsal kineties, number Dorsal kinety 1, number of bristles Dorsal kinety 2, number of bristles Dorsal kinety "3", number of bristles Dorsal kineties 2 and "3", number of bristles Dorsal kinety 1, number of bristles in proter dividers Dorsal kinety 2, number of bristles in proter dividers Caudal cirri, number
mean
M
SD
SE
2.0 2.1 6.5 1.6 8.1 4.3
2.0 2.0 7.0 1.0 8.0 4.0
0.0 – 0.9 0.7 1.3 0.7
0.0 – 0.2 0.2 0.3 0.1
10.3
10.0
0.8
2.0
2.0
0.0
CV
Min
Max
n
0.0 – 14.3 46.7 15.8 15.3
2.0 2.0 5.0 1.0 6.0 4.0
2.0 3.0 8.0 3.0 10.0 6.0
21 21 21 21 21 21
0.2
8.2
8.0
12.0
21
0.0
0.0
2.0
2.0
21
a All measurements in µm. CV = coefficient of variation in %, M = median, Max = maximum value, mean = arithmetic mean, Min = minimum value, n = number of individuals investigated, SD = standard deviation, SE = standard error of arithmetic mean. Data based on protargol-impregnated specimens.
the’s anlage III; anlagen IV and VI originate from primary primordia produced by the opisthe. All anlagen of the opisthe originate from the oral primordium, which incorporates the postoral cirral row, a product of the previous (parental) anlage IV. Cirral segregation occurs in middle dividers and usually produces some surplus cirri, which are later resorbed. Cirral migration commences in middle-late dividers, while the final positioning occurs in early and late postdividers. The anteriormost cirrus of anlage IV forms the rearmost frontoventral cirrus, while the remaining cirri of this anlage migrate to the postoral area to become the postoral ventral cirri (Fig. 110i, j).1 By contrast, this cirral group is formed from anlagen IV (1 cirrus) and V (2 cirri) in 18-cirri hypotrichs (see Berger 1999, p. 16)! The single transverse cirrus is the rearmost cirrus of anlage VI, while the anterior two cirri become the frontoterminal cirri and form a part of the frontoventral cirri (Fig. 110g–m). The formation of the dorsal kineties is rather complicated in A. halophila and differs distinctly from the process known from other species; thus, it is a further type of dorsal kinety formation. It commences in early dividers. The anterior bristle of kinety 1 produces an anlage in the proter. The second, third, and likely the forth bristle of kinety 2 generate an anlage each in proter and opisthe (Fig. 110n–p). While growing, the posterior portion of the opisthe anlage 2 splits off and migrates leftwards to become dorsal kinety 1 of the opisthe. Thus, two dorsal anlagen each in proter and opisthe are recognisable in middle dividers (Fig. 110q, r). In late and very late dividers, when a caudal cirrus each has been formed at the rear end of the anlagen and cell furrowing is in progress, one to three bristles at the anterior end of kinety 2 migrate more or less distinctly rightwards (or are possibly displaced by cell shaping), producing a third kinety in the right anterior quadrant of the cell. We carefully checked this feature and can exclude that this row is a dorsomarginal kinety (Foiss1
In the original description, we incorrectly assumed that all cirri formed by anlage IV migrate to the buccal vertex to form the postoral row (Foissner et al. 2002, p. 767).
Apourosomoida
529
Fig. 111a–g Cladotricha variabilis (from Ruinen 1938. From life). a: Ventral view of a representative specimen, body length of species 100–150 µm. This quadrinucleate specimen was fixed as type of C. variabilis by Foissner et al. (2002). b: Anterior body end in dorsal view. c: Degenerated specimen. d: Specimen possibly identical with Apourosomoida halophila. e: Nuclear apparatus of two specimens. Foissner et al. (2002) supposed that Ruinen mixed at least two species (see remarks at A. halophila). f: Rear body end showing three caudal cirri or two caudal cirri and one transverse cirrus as in A. halophila? g: Ventral view of anterior body portion. Note the single cirrus left of the anterior end of the postoral ventral row, a feature not observed in A. halophila. AZM = adoral zone of membranelles, RMR = right marginal row. Page 517.
ner et al. 2002); actually it is the anterior portion of kinety 2, as evident from bristle counts in interphase specimens and proter dividers (Fig. 110s–v). The “fragmentation” of dorsal kinety 2 of the opisthe is certainly not homologous with the fragmentation characterising the oxytrichids because (i) the posterior fragment migrates leftwards (vs. rightwards in the oxytrichids); (ii) the anterior fragment also forms a caudal cirrus (vs. does not form a caudal cirrus); and (iii) the fragmentation occurs only in the opisthe (vs. in both filial products). For phylogenetic implications of the ontogenetic data, see remarks at genus section. The formation of the new marginal rows proceeds in the ordinary manner, that is, two anlagen occur in each parental row. As mentioned above, no dorsomarginal kineties are produced (Fig. 110e–j). Nuclear division proceeds in the ordinary manner (Fig. 110a, c–e, q–v).
530
SYSTEMATIC SECTION
Occurrence and ecology: Apourosomoida halophila is likely confined to terrestrial and semi-terrestrial habitats (Foissner et al. 2002, p. 50). Type locality is the Unjab river bed in the area where it crosses the main road to the village of Terrace Bay, about 150 m north-east of the road (20°10'S, 13°10'E), northern Namib Desert (Namibia). The river bed is a swampy area grown with grasses, Cyperus, Phragmites, and shrubs. Up to 1 m high sand hills accumulate around the grass Odyssea paucinervis (brakweed). The sample was taken from such a sand hill, which contained so much organic debris that the sand was dark in colour. In the laboratory, no ciliates developed within three days because the sample had a salinity of 18% (Foissner et al. 2002). Thus, we washed it three times with tap water, decreasing the salinity to 15%. After a few days, Apourosomoida halophila appeared and reproduced rapidly. With time, it could be adapted to ordinary sea water (3.5%), but did not grow as well as between 10% and 15%, indicating that it prefers very high salinities. In addition, the present species occurred in Namibian site 61, that is, a highly saline soil from the margin of the Etosha Pan (Foissner et al. 2002). Feeds on bacteria (Foissner et al. 2002).
Apourosomoida natronophila (Dietz, 1965) comb. nov. (Fig. 112a, b) 1965 Uroleptus natronophilus n. sp. – Dietz, Arch. Protistenk., 108: 25, Abb. 1a, b (Fig. 112a, b; original description; no formal diagnosis provided and very likely no type material available). 2001 Uroleptus natronophilus Dietz, 1965 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 98 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Uroleptus natronophilus Dietz, 1965 – Foissner, Agatha & Berger, Denisia, 5: 771 (comparison with A. halophila).
Nomenclature: No detailed derivation of the species-group name is given in the original description. The name is a composite of the Greek noun nitron (the derived form is natron; soda), the thematic vowel ·o-, and the Greek adjective phil·us, -a, -um ([m; f; n]; loving, preferring), referring to the fact that the type locality is a soda lake. Uroleptus is masculine (Aescht 2001, p. 304) and Apourosomoida is feminine; thus, the ending of the species-group name has to be changed from -us to -a. Remarks: Dietz (1965) assigned this species to Uroleptus Ehrenberg, 1831, and supposed a relationship to Uroleptus agilis Engelmann, 1862, now the type species of Urosomoida Hemberger in Foissner, 1982 (for review, see Berger 1999, p. 345). Urosomoida basically comprises 18-cirri hypotrichs with a slightly decreased number of pretransverse ventral and transverse cirri and with a dorsomarginal kinety, but without kinety fragmentation. The original description of the present species is not very detailed, however, the data indicate that it is (i) not a Uroleptus because zigzagging ventral cirri are lacking (vs. present) and (ii) not a Urosomoida because it has more than three postoral ventral cirri (vs. 3) which are arranged in line, indicating that they originate from a single anlage during cell division (vs. not in line and
Apourosomoida
531
Fig. 112a, b Apourosomoida natronophila (from Dietz 1965. From life?). a: Ventral view showing cirral pattern (details must not be over-interpreted) and nuclear apparatus, 45 µm. Cirrus marked by short arrow is possibly cirrus III/2, long arrow denotes row of postoral ventral cirri; asterisks mark elongated and enlarged cirri at rear cell end; possibly the rearmost cirrus is a transverse cirrus (correct designation only possibly after redescription). b: Swimming specimen showing torsion. AZM = distal end of adoral zone of membranelles, FC = right frontal cirrus, LMR = left marginal row, MA = posterior macronuclear nodule, P = paroral, RMR = right marginal row (possibly the anteriormost cirri are frontoventral cirri). Page 530.
formed from two anlagen). Perhaps it is an Apourosomoida because (i) the five postoral ventral cirri are arranged in line; (ii) the number of transverse cirri is obviously low (perhaps this cirral group is even lacking); (iii) the distal three membranelles are separated from the proximal portion of the adoral zone (see below); and (iv) it was found in a mud/water sample from an African soda lake, that is, the species lives, like the type species, in an African, highly saline habitat. Dietz (1965) very likely misinterpreted the distal three adoral membranelles as frontal cirri, and the three cirri behind as anterior portion of the right marginal row. A buccal cirrus and frontoventral cirri are not illustrated; however, since the species is very small (only 35–50 µm long!) these cirri are – if present at all – very difficult to recognise and therefore details of the cirral pattern described and illustrated should not be overinterpreted. Anyhow, a classification in Apourosomoida seems more reliable than a classification in Uroleptus or Urosomoida. Thus, Uroleptus natronophilus is transferred to the present genus (see heading). Of course, a detailed reinvestigation (live observation; infraciliature of ventral and dorsal side; cell division to confirm the characteristic details of Apourosomoida) is a prerequisite for a final confirmation of the present classification. Likely it is confined to soda lakes or even to the locus classicus, that is, Lake Elmenteita in Kenya. Morphology: Body size 35–50 × 9–11 µm in life(?). Body outline cuneiform, that is, anterior portion broadly rounded and margins of posterior portion converging with rear cell end narrowly rounded or even pointed (Fig. 112a). Body cross-section oval with ventral side slightly flattened. Rather resistant to cover-glass pressure and
532
SYSTEMATIC SECTION
carmine acetic acid. Two very narrowly spaced macronuclear nodules right1 of buccal vertex and postoral ventral cirri (Fig. 112a); nodules truncated where they meet. Micronuclei not observed. Contractile vacuole likely lacking. Movement conspicuous because body twists by a quarter to two thirds when swimming (Fig. 112b). Adoral zone occupies 27% of body length in specimen illustrated, distal three membranelles (misinterpreted as frontal cirri in original description) widely spaced and set off from proximal portion. Buccal cavity obviously narrow. Paroral(?) distinct and sail-like (Fig. 112a). Cirral pattern basically as shown in Fig. 112a; however, note that details should not be over-interpreted. Three frontal cirri not enlarged, behind distal adoral membranelles. No buccal cirrus illustrated (lacking? overlooked?). One cirrus slightly right and behind of right frontal cirrus; perhaps this is cirrus III/2. No frontoventral cirri illustrated; possibly misinterpreted as anterior portion of right marginal row. Five longitudinally arranged postoral ventral cirri in body midline, row commencing slightly behind level of buccal vertex. Specimen illustrated with about 26 right and 17 left marginal cirri (values must not be over-interpreted); marginal cirri long, thin, and widely spaced. 4–5 cirri at rear cell end elongated and enlarged (Fig. 112a); correct interpretation and designation only possibly after protargol impregnation (perhaps only one transverse cirrus is present, as in the type species). Dorsal infraciliature (length of bristles, number and arrangement of kineties, presence/absence of caudal cirri) not known; thus, present classification in Apourosomoida not certain (see remarks). Occurrence and ecology: Apourosomoida natronophila is possibly confined to African soda lakes. Type locality is Lake Elmenteita in Kenya, where Heinz Löffler (Austrian Academy of Sciences) collected a mud and water sample. The sample was checked for ciliates for the first time two years after collecting; when the sample was dry it was re-wetted with distilled water. The sample had an acid binding capacity of 2000. In a mixture of 1.5 parts of sample water and one part sea water, the present species died immediately. In a solution containing 2% NaCl and 0.8% Na2CO3 (NaHCO3), specimens survived and reproduced for several weeks (Dietz 1965). No further records published. Food not known; Apourosomoida natronophila, however, was ingested by Spathidium elmenteitanum Dietz, 1965 (Dietz 1965).
Bistichella gen. nov. Nomenclature: The genus-group name Bistichella is a composite of the Latin numeral bi- (two), the Greek substantive stich- (row, line), and the diminutive suffix -ella, referring to the two long frontoventral rows (rows V and VI) present in all five species included in this genus. Feminine gender because ending with -ella (ICZN 1999, Article 30.1.3).
1
In Foissner et al. (2002) we incorrectly wrote “in the left anterior half”.
Bistichella
533
Characterisation (A = supposed apomorphy): Hypotricha with two short frontal and two long frontoventral rows (A). Adoral zone of membranelles continuous. Three enlarged frontal cirri. More than one buccal cirrus (A). Postperistomial cirrus lacking. Transverse cirri lacking (type species, B. humicola?) or present. Caudal cirri lacking (type species, B. namibiensis, B. humicola?) or present. One left and one right marginal row. Dorsal kinety pattern of Gonostomum-type, that is, dorsomarginal row and dorsal kinety fragmentation lacking. Mainly terrestrial. Additional characters: Body flexible; contractile vacuole at left cell marginal about at mid-body or slightly ahead of it; cortical granules lacking; dorsal bristles short, that is, less than 5 µm long. Type species: Paraurostyla buitkampi Foissner, 1982. Remarks: Four species included in Bistichella (see next chapter) were previously assigned to Pseudouroleptus because they have – like the type species P. caudatus Hemberger, 1985 – two long frontoventral rows. However, Pseudouroleptus caudatus has a postperistomial ventral cirrus which is homologous to the postoral ventral cirrus IV/2 of the 18-cirri hypotrichs (Fig. 136a, h; Fig. 6a in Berger 1999). Since this cirrus, which is even conspicuous in live specimens, is lacking in the other four species, I remove them from Pseudouroleptus and put them into the new genus Bistichella because they cannot be included in any other group. Consequently, Pseudouroleptus is now confined to the type species comprising two subspecies. Since it has a dorsal kinety fragmentation it was classified in the oxytrichids by Berger (1999, p. 888). A review of Pseudouroleptus, including the new data by Foissner et al. (2002), is added at the end of the book in the Supplement to the Oxytrichidae chapter. The fifth species classified in Bistichella is Amphisiella namibiensis (Fig. 114a–g). It lacks the classical amphisiellid median cirral row, and since its cirral pattern is very similar to that of the other Bistichella species I transfer it to the present genus. I selected B. buitkampi as type species because it is well defined and described. Bistichella terrestris and B. humicola are described mainly/only after stained specimens and for B. procera a synonymy with B. terrestris cannot be completely excluded. Bistichella namibiensis differs from B. buitkampi, inter alia, by the presence of true transverse cirri. Bistichella buitkampi and B. namibiensis obviously lack caudal cirri. By contrast, Bistichella terrestris and B. procera have cirri of unknown origin at the posterior body end so that the presence of caudal cirri cannot be excluded. For B. humicola the situation regarding this cirral group is not known; in addition, this species lacks a second short frontoventral row and has only one buccal cirrus. Bistichella buitkampi has a longitudinal row between the rear portion of the marginal rows, a cirral group which is certainly not identical with the true transverse cirri present in B. namibiensis. Thus, one cannot exclude that Bistichella is non-monophyletic, although the overall-appearance of the five species included is rather similar. Possibly, the differences evolved within Bistichella.
534
SYSTEMATIC SECTION
All species included in the present genus were classified in the amphisiellids previously. Although the cell division is not known for any of the species one can conclude that they lack the typical amphisiellid median cirral row composed of at least two parts. Thus, I do not classify Bistichella in the Amphisiellidae, but consider it as taxon of unknown position in the Hypotricha. Since they obviously have the plesiomorphic dorsal kinety pattern, that is, lack a dorsomarginal row and kinety fragmentation, they obviously branched off, like the amphisiellids, outside the Dorsomarginalia. From the cirral pattern of B. buitkampi, B. namibiensis, and B. procera one can conclude that they have the ordinary (plesiomorphic) number of six (I–VI) frontal-ventral-transverse cirri anlagen. Thus, the cirral rows are designated as frontal row III (short row behind right frontal cirrus), frontoventral row IV (short row right of frontal row III), frontoventral row V (left long row), and frontoventral VI (right long row). Possibly the right row, which is homologous to the frontoterminal cirri of other hypotrichs, is too long to form a continuous row with the row formed by anlage V, as is the case in the amphisiellids. Of course, I cannot exclude that Bistichella is an amphisiellid which does not yet or no longer forms a continuous amphisiellid median cirral row. Possibly, relevant molecular data will show us the correct phylogenetic position of this genus. Species included in Bistichella (alphabetically arranged basionyms are given): (1) Amphisiella namibiensis Foissner, Agatha & Berger, 2002; (2) Paraurostyla buitkampi Foissner, 1982; (3) Pseudouroleptus procerus Berger & Foissner, 1987; (4) Pseudouroleptus terrestris Hemberger, 1985. Incertae sedis: (5) Uroleptus humicola Gellért, 1956b.
Key to Bistichella species Main features for the identification are the number of macronuclear nodules, the cirral pattern, and the number of dorsal kineties; thus, protargol impregnation is recommended. 1 Many (32 in specimen illustrated) macronuclear nodules (Fig. 118a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bistichella humicola (p. 556) - Two or 4 macronuclear nodules. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2 Four macronuclear nodules (Fig. 113a, f). . . . . . . . Bistichella buitkampi (p. 535) - Two macronuclear nodules. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3 Frontoventral rows V and VI extend to prominent transverse cirri (Fig. 114a–h). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bistichella namibiensis (p. 538) - Frontoventral rows V and VI terminate far ahead of inconspicuous transverse cirri (e.g., Fig. 116c). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4 Three dorsal kineties (Fig. 116a–f). . . . . . . . . . . . . . . Bistichella procera (p. 547) - Four dorsal kineties (Fig. 117a, b). . . . . . . . . . . . . . Bistichella terrestris (p. 554)
Bistichella
535
Bistichella buitkampi (Foissner, 1982) comb. nov. (Fig. 113a–f, Table 33) 1982 Paraurostyla buitkampi nov. spec.1 – Foissner, Arch. Protistenk., 126: 40, Abb. 4a–f, Tabelle 7 (Fig. 113a–f; original description; the holotype slide [accession number 1981/82; Aescht 2003, p. 382] is deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria). 1987 Pseudouroleptus buitkampi (Foissner, 1982) nov. comb. – Berger & Foissner, Zool. Jb. Syst., 114: 197 (combination with Pseudouroleptus). 2001 Pseudouroleptus buitkampi (Foissner, 1982) Berger and Foissner, 1987 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 69 (nomenclator containing all basionyms, combinations, and higher taxa of euplotids and hypotrichs).
Nomenclature: Foissner (1982) dedicated this species to Ulrich Buitkamp (Germany), who made basic studies about the systematics and ecology of soil ciliates (e.g., Buitkamp 1977, 1977a, 1979, Buitkamp & Wilbert 1974). Remarks: Foissner (1982) assigned this species to Paraurostyla Borror, 1972 obviously because of the cirral rows, whose arrangement is reminiscent of that of P. weissei, type of the genus. However, Paraurostyla weissei (Stein, 1859) Borror, 1972 has dorsomarginal rows and a dorsal kinety fragmentation, which are certainly lacking in B. buitkampi because it has only the ordinary three bipolar kineties (Fig. 113f). Further, the present species lacks transverse cirri (the cirral row in the posterior portion is obviously a true row and not a pseudorow which is characteristic for transverse cirri) and caudal cirri, whereas these cirral groups are well developed in Paraurostyla (for review, see Berger 1999, p. 841). When we described Pseudouroleptus procerus (now Bistichella procera) we recognised that Paraurostyla buitkampi has the same type of infraciliature and therefore transferred it to Pseudouroleptus. For a foundation of the transfer of both species to Bistichella, see genus section. Morphology: Body size 135–180 × 50–70 µm in life, on average 110 × 31 µm after protargol impregnation, that is, length:width ratio about 3.5:1 (Table 33). Body outline slightly dumbbell-shaped to sigmoidal, anterior and posterior end broadly rounded. Body flattened about 2:1 dorsoventrally, ventral side concave, dorsal convex, sometimes slightly twisted about main body axis (Fig. 113a–c). Invariably (n = 10) four macronuclear nodules left of cell midline arranged in two distinct pairs (Fig. 113a, f); individual nodules about 19 × 12 µm in size, with large, regularly distributed chromatin bodies. Micronuclei about 4 × 3 µm in life, inconspicuous, attached to macronuclear nodules (Fig. 113a, f). Contractile vacuole about in midbody, during diastole with short, lacunar collecting canals; excretory pore, as is usual, on dorsal side. Cortical granules lacking. Cytoplasm colourless, packed with 1
Foissner (1982) provided the following diagnosis: In vivo etwa 135–180 × 50–70 µm große, meist leicht hantelförmige, sehr biegsame Paraurostyla mit je 2 dicht hintereinander liegenden MakronucleusTeilen in der vorderen und hinteren Körperhälfte. 2 mäßig und 2 stark verkürzte Ventralreihen, die auf der rechten Körperhälfte von rechts oben schräg nach links unten verlaufen. Hinten eine vertikal orientierte Reihe von durchschnittlich 8 Transversalcirren(?). 3 Dorsalkineten.
536
SYSTEMATIC SECTION
Fig. 113a–d Bistichella buitkampi (from Foissner 1982. From life). a: Ventral view of a representative specimen, 164 µm. Arrow marks the short, longitudinal cirral row in the posterior body portion (termed transverse cirri in original description). Note the highly characteristic four macronuclear nodules which form two distinct pairs. b: Dorsal view showing contractile vacuole. c: Left lateral view showing dorsoventral flattening, 152 µm. d: Specimen during ingestion of a food organism (arrow); body length = 150 µm. Note the characteristic body shape which is due to the very flexible anterior body portion. AZM = adoral zone of membranelles, CV = contractile vacuole with collecting canals. Page 535.
tiny, shiny granules and 2–5 µm-sized, slightly yellowish, shiny globules. Usually many food vacuoles 8–20 µm across. Movement slow, burrowing. Adoral zone occupies 26% of body length and composed of 37 membranelles on average (Fig. 113a, b, e, Table 33); bases of largest membranelles about 10 µm wide. Buccal field deep. Undulating membranes two-rowed, roughly in Cyrtohymena-pattern (Berger 1999, p. 62), that is, paroral strongly curved anteriorly and endoral obliquely extending across buccal field with its anterior end near distal end of paroral (Fig. 113e). Pharyngeal fibres extend obliquely backwards. Cirral pattern and number of cirri of usual variability (Fig. 113a, e, Table 33). Frontal cirri distinctly enlarged, arranged in slightly oblique pseudorow with right cirrus behind distal end of adoral zone. 2–4, usually four buccal cirri along right side of paroral. 2–3, usually three, paramalar cirri behind right frontal cirrus (= frontal row III), ends at 15% of body length on average (Table 33). Frontal row IV com-
Bistichella
537
Fig. 113e, f Bistichella buitkampi (from Foissner 1982). Infraciliature of ventral and dorsal side and nuclear apparatus, 81 µm. Short arrow in (e) marks buccal cirral row, long arrow denotes the longitudinal cirral row in the posterior body portion (designated as transverse cirri in the original description). The three frontal cirri, which are distinctly enlarged, are connected by a dotted line. AZM = adoral zone of membranelles, E = endoral, FC = right frontal cirrus, LMR = left marginal row, MA = macronuclear nodule, MI = micronucleus, P = paroral, PF = pharyngeal fibres, RMR = right marginal row, III, IV = frontal rows, V, VI = frontoventral rows, 1–3 = dorsal kineties. Page 535.
posed of about five cirri, commences roughly at same level as row III, terminates at 19% of body length. Frontoventral row V composed of about 32 cirri, commences at 18% of body length in specimen illustrated (Fig. 33e), ends at 75% of body length on average. Frontoventral row VI begins near right frontal cirrus and terminates at 59% of body length, composed of 25 cirri on average. A short longitudinal cirral row (termed transverse cirri in original description) in rear body portion. Right marginal row distinctly shortened anteriorly, slightly shortened posteriorly. Left row commences left of proximal portion of adoral zone, extends to cell end leaving a distinct gap between marginal rows (Fig. 113e). Dorsal bristles about 2 µm long in life, arranged in three kineties; kinety 1 distinctly shortened anteriorly, ends near cell end; kineties 2 and 3 only slightly shortened anteriorly, but distinctly shortened posteriorly. Distance between individual bristles about equal in kinety 2, whereas bristles posteriorly more widely spaced than anteriorly in kineties 1 and 3 (Fig. 113f). Caudal cirri lacking.
538
SYSTEMATIC SECTION
Observations from a population from the Fuscher Tal, a valley in the Central Alps in Salzburg, Austria (kindly supplied by W. Foissner): body size about 140 × 55 µm in life; four macronuclear nodules; cortical granules lacking; dorsal bristles about 3 µm long. Observations from a New Zealand population (kindly supplied by W. Foissner): body size about 160 × 50–60 µm; body very flexible, with anterior body portion rather motile; four macronuclear nodules, in life about 17 × 10 µm; micronuclei not recognisable in life; contractile vacuole slightly ahead of mid-body, without distinct canals; cortical granules lacking; no cytoplasmic crystals; cirri about 15 µm long. Occurrence and ecology: Likely confined to terrestrial habitats (Foissner 1987, 1998). Type locality of Bistichella buitkampi is the Schloßalm area near the village of Bad Hofgastein, Salzburg, Austria (1950 m above sea-level), where Foissner (1982) discovered it in the soil of an alpine pasture. For some autecological data see Foissner & Peer (1985, p. 42). Further records: agricultural soil near the villages Frauendorf and Bierbaum, Lower Austria (Foissner et al. 1985, p. 108); upper soil layer (0–5 cm; pH 7.5; 1240 m altitude) on rocks grown with lichens and calciphil plants in the Fuscher Valley near the Vögerl Alm, Salzburg, Austria (Foissner 1987b, p. 57); soil from a meadow near Mount Herbert, New Zealand (W. Foissner, pers. comm.). Not recorded during a detailed survey of the ciliate cenosis of Namibian soils (Foissner et al. 2002). Bistichella buitkampi feeds on bacteria and ciliates (Colpoda inflata, Gonostomum affine), but food vacuoles also contain soil particles (Foissner 1982). The prey is fixed in the anterior half of the oral apparatus and ingested in portions, whereby the specimens have a characteristic body outline (Fig. 113d). Biomass of 106 specimens about 72 mg (Foissner 1987a, p. 127; 1998, p. 208).
Bistichella namibiensis (Foissner, Agatha & Berger, 2002) comb. nov. (Fig. 114a–g, 115a–h, Table 34) 2002 Amphisiella namibiensis nov. spec.1 – Foissner, Agatha & Berger, Denisia, 5: 656, Fig. 147a, b, h–l, 386a–g, 390f, Table 128 (Fig. 114a–g, 115a–h; original description; one holotype slide [accession number 2002/108 and six paratype slides [accession numbers 2002/5, 8, 109, 110, 147, 148] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Austria; Foissner et al. 2002, p. 37 and Aescht 2003, p. 392).
Nomenclature: The species-group name namibiensis refers to the country (Namibia), where the species was discovered (Foissner et al. 2002). 1
Foissner et al. (2002) provided the following diagnosis: Size about 250 × 80 µm in vivo; elongate ellipsoidal. 2 macronuclear nodules and 2–6 (mean = 4) micronuclei. Usually 2 long ventral rows extending to transverse cirri: right composed of 43, left of 40 cirri on average. On average 50 right marginal, 51 left marginal, 4 buccal, and 5 transverse cirri; 1–6 (mean = 4) cirri between buccal row and anterior end of ventral rows. 3 dorsal kineties. Adoral zone about one third of body length, composed of 44 membranelles on average.
Bistichella
Fig. 114a, b Bistichella namibiensis (from Foissner et al. 2002. From life). Ventral and lateral view of a representative specimen, 235 µm. Page 538.
539
Remarks: For a foundation of the transfer from Amphisiella to Bistichella and a general comment on the somewhat curious cirral pattern, see genus section. Some features of B. namibiensis are reminiscent of Uroleptoides polycirratus, for example, the large oral apparatus, the rather prominent frontal and transverse cirri, and the increased number of buccal cirri (Fig. 55a–d). However, this species has, as is usual for an amphisiellid, only one median cirral row. Apoamphisiella Foissner, 1997a, Parentocirrus Voß, 1997, and Paraurostyla fossicola (Kahl, 1932) Borror, 1972 also have a similar cirral pattern. However, they are oxytrichids, that is, they have, inter alia, a fragmenting dorsal kinety, dorsomarginal kineties, and a postoral ventral cirrus (Berger 1999, Blatterer & Foissner 2003). For separation from congeners, see key. Morphology: Body size 140–300 × 40–100 µm in life, usually near 250 × 80 µm; length:width ratio about 3:1 in life, while 2.8–6.3:1, on average 4.3:1 in the protargol preparations, where most specimens are rather distorted (Table 34). Body very flexible, but acontractile. Overall body shape elongate ellipsoidal, slightly narrowed posteriorly, both ends broadly rounded, dorsoventrally flattened up to 2:1 (Fig. 114a, b). Macronuclear nodules in middle third of cell, near or slightly left of midline, ellipsoidal, with numerous chromatin bodies 1–2 µm across. Micronuclei usually near or attached to macronuclear nodules, globular. Contractile vacuole near mid-body at left cell margin, during diastole with two collecting canals extending anteriad and posteriad. No specific cortical granules. Cytoplasm colourless, without crystalline in-
540
SYSTEMATIC SECTION
Fig. 114c, d Bistichella namibiensis (from Foissner et al. 2002. Protargol impregnation). Infraciliature of ventral and dorsal side and nuclear apparatus of holotype specimen, 260 µm. Broken line connects cirri originating from cirral anlage III (corresponding transverse cirrus not included). LMR = anterior end of left marginal row, MA = anterior macronuclear nodule, MI = micronucleus, RMR = right marginal row, TC = transverse cirri, V, VI = frontoventral rows V, VI, 1–3 = dorsal kineties. Page 538.
Bistichella
541
Fig. 114e Bistichella namibiensis (from Foissner et al. 2002. Protargol impregnation). Frontal cirri connected by dotted line, cirri left of anterior portion of frontoventral row V circled. Right frontal cirrus and cirrus III/2 connected by broken line. AZM = adoral zone of membranelles, BC = anteriormost buccal cirrus, E = endoral, FC = left frontal cirrus, LMR = left marginal row, P = paroral, III/IV = frontal rows, V, VI = frontoventral rows. Page 538.
542
SYSTEMATIC SECTION
Bistichella
543
clusions, contains some colourless, greasily shining globules 1–5 µm across and about 10–40 µm-sized food vacuoles. Glides quickly to and fro on microscope slide. Oral apparatus conspicuous because of the large, deep buccal cavity and curved paroral resembling cyrtohymenids (Fig. 114a, c, e, f, 115a–d, f, Table 34; for review on cyrtohymenids, see Berger 1999). Adoral zone occupies about one third of body length, proximal third of zone rather distinctly broadened and covered by a hyaline cortical process (lip); composed of an average of 44 membranelles, bases of largest membranelles about 20 µm wide. Structure of membranelles depends on zone region (Fig. 114f): those along buccal cavity composed of two long rows, one slightly shortened row, and a very short anterior row, which consists of only three basal bodies; the next 5–8 membranelles have the short anterior row composed of about 10 basal bodies; membranelles in distal third of zone composed of two long rows and one slightly shortened row of basal bodies, to which, except for last membranelle, a fairly short row is attached in midportion. Fibrillar system of adoral membranelles also depends on zone region (Fig. 114f): membranelles of region (a) with short fibre bundle directed to frontal field and bent dorsally; membranelles of region (b) with short fibre bundle directed to frontal field and curved dorsally and to the left, likely touching fibre bundle from neighbouring membranelle; membranelles of region (c) with very long submembranellar fibre bundle originating from left half of membranelle and extending to proximal end of zone; fibrillar associates of region (d) like those of region (c), but with fine, straight fibre extending between individual membranelles into buccal cavity; membranelles of region (e) also have intermembranellar fibres, but all other associates are lacking or inconspicuous.
b
b
Fig. 114f, g Bistichella namibiensis (from Foissner et al. 2002. Protargol impregnation). Fibrillar associates of structures in oral region and around transverse cirri (cp. Fig. 115b–g). Cirri have two fibre bundles, except for the left frontal cirrus and the transverse cirri, which lack the posterior bundle, and the buccal cirri, which lack the anterior bundle. The fibrillar associates of the adoral membranelles depend on the zone region: membranelles of regions (a) and (b) each have a short bundle; membranelles of region (c) have a very long submembranellar bundle extending to proximal end of zone; in region (d), a fine, straight fibre occurs between the individual membranelles and extends into the buccal cavity; the membranelles of region (e) possibly lack submembraneller bundles. The paroral fibres line the right wall of the buccal cavity, while those of the endoral line the bottom. a–e = regions of adoral zone (margins of regions marked by asterisks), LMR = left marginal row, RMR = right marginal row, TC = transverse cirri, V, VI = frontoventral rows. Page 538.
544
SYSTEMATIC SECTION
Fig. 115a–c Bistichella namibiensis (from Foissner et al. 2002. Protargol impregnation). Somatic and oral infraciliature. a: Ventral view. Bistichella namibiensis is characterised by (i) an elongate body with both ends broadly rounded, (ii) two ellipsoidal macronuclear nodules in middle third of cell, (iii) a right and a left marginal row frequently slightly overlapping posteriorly (arrowhead), (iv) two long frontoventral rows of almost body length, (v) three frontal cirri arranged right (in the original description we incorrectly wrote left) of midline, and (vi) an adoral zone of membranelles occupying about one third of body length. Note some slightly enlarged cirri between the distinctly curved paroral and the anterior portion of the frontoventral row V (arrow). Asterisk marks granule patch in bottom of buccal cavity. b, c: Anterior ventral portion at higher magnification. The frontal cirri are arranged right of midline with the rightmost cirrus close to distal end of adoral zone. They comprise 8–13 kineties with 6–8 cilia each. The buccal cirri, which form a row along the mid-portion of the paroral and attenuate from anterior to posterior, comprise 4–7 kineties with 6–9 cilia each. Both frontoventral rows (= amphisiellid median cirral rows in original description) commence behind the distal end of the adoral zone of membranelles. The paroral and the endoral are composed of closely spaced dikinetids, from which conspicuous fibre bundles originate (arrowheads). Argyrophilic granules form a conspicuous patch in the buccal bottom. Note that the short, fourth ciliary row of the proximal membranelles becomes longer in the frontal area of the zone (asterisk). Explanation of original labelling: AG = argyrophilic granules, AZM = adoral zone of membranelles, BU = buccal cirri, EM = endoral, FC = frontal cirri, LMR = left marginal row, LVR = frontoventral row V, MA = macronuclear nodule, PM = paroral, RMR = right marginal row, RVR = frontoventral row VI. Scale bar 60 µm. Page 538.
Bistichella
545
Fig. 115d–g Bistichella namibiensis (from Foissner et al. 2002. Protargol impregnation). Infraciliature. d: Oral area. Numerous argyrophilic granules form a V-shaped patch in the buccal cavity: the left fork extends along the adoral zone, the right along the posterior portion of the endoral. The paroral optically crosses the endoral in mid-buccal area; its fibre bundles (arrow) line the right wall of the cavity. The endoral traverses the bottom of the cavity, which is lined by the endoral fibre bundles (arrowheads). e: Fibrillar associates of structures in posterior ventral portion. The conspicuously long, antero-laterally directed fibres associated with the individual transverse cirri unite to a thick bundle (arrow). Note the slightly overlapping marginal rows (arrowhead). f: Buccal area at higher magnification. The undulating membranes are associated with 10–20 µm long fibre bundles (arrowheads). Fine, straight fibres occur between the individual membranelles and extend into the buccal area (arrow; cp. Fig. 114f). g: Detail of frontoventral rows (= amphisiellid median cirral rows in original description). Each cirrus has an anteriad (arrow) and posteriad (arrowhead) directed fibre bundle. Explanation of original labelling: AG = argyrophilic granules, AZM = adoral zone of membranelles, BU = buccal cirri, EM = endoral, FC = frontal cirri, LMR = left marginal row, LVR = frontoventral row V, PM = paroral, RMR = right marginal row, RVR = frontoventral row VI, TC = transverse cirri. Page 538.
546
SYSTEMATIC SECTION
Buccal cavity deep and wide, with conspicuous accumulation of argyrophilic granules right of adoral zone and in posterior half of right wall (Fig. 115a–d, f). Paroral composed of closely spaced, about 15 µm long cilia; straight in posterior half, strongly bent leftwards anteriorly, where it follows curvature of buccal cavity; optically crosses endoral in mid-buccal area. Endoral posteriorly slightly longer than paroral, so strongly curved that ends almost touch adoral zone of membranelles, anterior half traverses bottom of buccal cavity. Both membranes associated with 10–20 µm long fibre bundles, those of paroral line right wall of buccal cavity, those of endoral line bottom. Pharyngeal fibres surprisingly inconspicuous, extend obliquely backwards. Cirral pattern and number of cirri of usual variability (Fig. 114a, c, e, 115a, e, g, Table 34), except for buccal cirri (2–5), and cirri in frontal rows III and IV. Frontal cirri about 25 µm long in life, right of midline; right cirrus, as is usual, close to distal end of adoral zone of membranelles, composed of 8–13 kineties with 6–8 cilia Fig. 115h Bistichella namibiensis (from Foissner each. Buccal cirri along mid-portion of et al. 2002. Protargol impregnation). Infraciliaparoral, anteriormost cirrus invariably ture of ventral side and nuclear apparatus. Explanear level of right (= posteriormost) fronnation of original labelling: AZM = adoral zone tal cirrus, become slightly thinner from of membranelles, EM = endoral, MA = macronuanterior to posterior; composed of 4–7 kiclear nodules, MI = micronucleus, PM = paroral, TC = transverse cirri, VR frontoventral row V (= neties with 6–9 cilia each. 1–6, usually left amphisiellid median cirral row in original defour cirri between buccal row and anterior scription). Page 538. portion of left frontoventral row, usually arranged as shown in Fig. 14c, e, f, 115d, h, but several variations occur, for instance, cirri may be side by side (“midventral pattern”), or the left anterior cirrus is absent; cirri slightly enlarged, comprising 5–6 kineties with 5–8 cilia each. Two frontoventral rows commencing behind distal end of adoral zone and extending slightly obliquely to transverse cirri; in two out of 45 specimens a third such row occurs, and two specimens had some cirri between the anterior portion of the rows. Very likely, the right row is homologous to the frontoterminal cirri (see remarks at genus section).
Bistichella
547
Cirri of these rows about 15 µm long in life and usually composed of two kineties with 6–7 cilia each, anteriormost cirri of both rows frequently slightly enlarged, that is, composed of 3–5 kineties with 5–7 cilia each. Postperistomial cirri lacking. Transverse cirri about 25 µm long in life, inconspicuously projecting beyond rear body margin, form slightly curved, oblique pseudorow at, or slightly left of, midline; usually composed of 6–8 kineties with 5–6 cilia each. Fibrillar associates of cirri as shown in Fig. 115g, e; note lack of posterior fibre bundle in transverse cirri and anteriormost frontal cirrus as well as absence of anterior fibre bundle in buccal cirri. Marginal rows more or less widely overlapping posteriorly, that is, left row extends to or above midline at posterior end of cell, while right row commences dorsolaterally at level of anteriormost buccal cirrus and extends terminally to, or slightly beyond, midline; cirri usually composed of two kineties with 6–7 cilia each, anteriormost cirri of both rows frequently slightly enlarged, that is, consisting of 3–5 kineties with 5–7 cilia each. Dorsal bristles about 3 µm long in life, arranged in three rows slightly shortened anteriorly and posteriorly (Fig. 114d). Dorsomarginal kinety and kinety fragmentation obviously lacking. No caudal cirri. Occurrence and ecology: Bistichella namibiensis is possibly confined to terrestrial habitats (Foissner et al. 2002, p. 50). However, the large, blunt body indicates that it may be a limnetic species active mainly during floods. Type locality is the Wolfsnes water-hole (19°S 15°50'E) near the margin of the Etosha Pan (Namibia), where it occurred in a highly saline (15‰) sample (litter and soil from around decaying grass shrubbery) about 500 m off pan margin. Also found in two other sites of the Etosha National Park (Foissner et al. 2002), and in a forest soil (Kolmberg, 47°58'N 16°41'E) from Austria (Foissner et al. 2005). Bistichella namibiensis feeds on ciliates, for example, Drepanomonas revoluta and Gonostomum sp. (Foissner et al. 2002).
Bistichella procera (Berger & Foissner, 1987) comb. nov. (Fig. 116a–f, Table 33) 1987 Pseudouroleptus procerus nov. spec.1 – Berger & Foissner, Zool. Jb. Syst., 114: 195, Fig. 1–6, Table 2 (Fig. 116a–f; original description; the holotype slide [accession number 1986/61] and a paratype slide [1986/62] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; Aescht 2003, p. 394). 2001 Pseudouroleptus procerus Berger and Foissner, 1987 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 77 (nomenclator containing all basionyms, combinations, and higher taxa of euplotids and hypotrichs). 2002 Pseudouroleptus procerus – Lynn & Small, Phylum Ciliophora, p. 452, Fig. 33A, B (Fig. 116c, d; guide to ciliate genera).
1
Berger & Foissner (1987) provided the following diagnosis: In vivo about 170–250 × 30–40 µm (n = 4), vermicular, 2 unshortened dorsal kineties and 1 extremely reduced (1 basal body pair) dorsal kinety. 33 adoral membranelles, 47 left and 48 right marginal cirri on the average.
548
SYSTEMATIC SECTION
Nomenclature: No derivation of the name is given in the original description. The species-group name procer·us, -a, -um (Latin adjective [m, f, n]; slender, tall) refers to the slender body shape. For the problem with the type locality, see remarks. Remarks: Berger & Foissner (1987) studied three Austrian populations (meadow near the city of Salzburg; pasture near the village of Seekirchen; arable soil near the city of Vienna) of this species and provided a combined description of all populations which, however, agree very well so that conspecificity is beyond reasonable doubt. Unfortunately, we fixed the meadow near the city of Salzburg as type locality, although the type slides are from the population from the arable soil near the city of Vienna1. Thus, according to Article 76A.2 of the ICZN (1999) the statement about the type locality has to be corrected (see occurrence and ecology section). For a foundation of the transfer from Pseudouroleptus to Bistichella, see genus section. Bistichella procera is very similar to B. terrestris (Fig. 117a). They differ distinctly only in the number of dorsal kineties (3 [kinety 1 composed of only 1 bristle!] vs. 4). The body length:width ratio is also different (4–6:1 vs. 3:1 [from Fig. 117a]). However, the body shape data by Hemberger (1985) must not be overinterpreted because he studied live specimens somewhat superficially. The number of dorsal kineties is usually a rather constant feature so that I accept both species. Another feature separating B. procera and B. terrestris is the arrangement of the two short frontal rows. In B. procera they are behind the right frontal cirrus and right of it, whereas in B. terrestris they are behind the middle (very unusual!) and the right frontal cirrus. Further populations should be studied to show whether or not these differences can be confirmed. If not, Bistichella procera should be put into the synonymy of B. terrestris. If you are uncertain about the dorsal kinety pattern you should write Bistichella terrestris-group. Bistichella buitkampi has four macronuclear nodules (vs. two in present species), three bipolar dorsal kineties, and a wider body. Bistichella humicola, which has not Fig. 116a–f Bistichella procera (from Berger & Foissner 1987. a, b, from life; c–f, protargol impregnation. a, population from a meadow near the city of Salzburg; b, population from a pasture near the village of Seekirchen; c–f, type population from the Marchfeld area near the city of Vienna). a: Ventral view of a representative specimen, 171 µm. b: Ventral view of a twisted specimen, 246 µm. Arrow marks buccal cirri, arrowhead denotes notch at right posterior body margin. c, d: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 150 µm. Arrow in (c) marks transverse cirri, short arrow in (d) denotes dorsal kinety 1, which is composed of a single bristle only, and long arrow in (d) marks short cirral row (caudal cirri? at rear cell end). Frontal cirri connected by dotted line. e, f: Infraciliature of posterior body portion in ventral and dorsal view. Arrow in (e) marks transverse cirri, arrow in (f) denotes caudal (?) cirri. AZM = adoral zone of membranelles, CV = contractile vacuole, E = endoral, LMR = left marginal row, MA = macronuclear nodule, MI = micronucleus, P = paroral, RMR = right marginal row, III, IV = frontal rows (designated as ventral rows 1 and 2 in original description), V, VI = frontoventral rows (designated as ventral rows 3 and 4 in original description), 1–3 = dorsal kineties. Page 547. 1
According to a personal communication by Erna Aescht from the Upper Austrian Museum in Linz (LI), the designation on the type slides is “Marchfeld 1985”. Marchfeld is an area east of Vienna. For detailed description of the correct type locality, see occurrence and ecology.
d
Bistichella
549
550
SYSTEMATIC SECTION
Table 33 Morphometric data on Bistichella buitkampi (bui, from Foissner 1982), Bistichella humicola (hum, from Gellért 1956b), Bistichella procera (pro, type population; from Berger & Foissner 1987), and Bistichella terrestris (ter, from Hemberger 1985) Characteristics a Body, length
Species
bui hum h pro ter f Body, width bui pro ter f Adoral zone of membranelles, length bui pro Posterior macronuclear nodule, length bui c pro Posterior macronuclear nodule, width bui c pro Macronuclear nodules, distance in between pro Posterior micronucleus, length bui d pro Posterior micronucleus, width bui d pro Anterior body end to rear end of frontal bui row III, distance pro Anterior body end to rear end of frontal bui row IV, distance pro Anterior body end to rear end of frontobui ventral row V, distance pro Anterior body end to rear end of frontobui ventral row VI, distance pro Macronuclear nodules, number bui hum pro ter Micronuclei, number bui pro ter Adoral membranelles, number bui hum pro ter Frontal cirri, number bui hum pro ter Buccal cirri, number bui hum pro ter Frontal and frontoventral rows, number bui Frontal row III, number of cirri bui hum
mean
M
109.6 111.0 120.0 – 139.0 140.0 190.0 – 31.2 23.5 32.7 32.5 60.0 – 28.6 28.5 36.3 36.0 13.3 14.0 16.3 15.0 7.1 7.0 7.6 7.5 15.5 15.0 2.4 2.5 3.0 3.0 2.2 2.2 2.3 2.3 14.4 13.0 15.5 15.0 21.2 20.0 24.3 24.5 82.7 80.0 67.6 67.5 64.7 66.0 90.2 88.0 4.0 4.0 32.0 – 2.0 2.0 2.0 – 2.0 2.0 2.0 2.0 2.0 – 36.6 36.5 31.0 – 32.7 33.0 – – 3.0 3.0 3.0 – 3.0 3.0 3.0 – 3.6 4.0 1.0 – 3.8 4.0 4.0 – 3.9 4.0 2.7 3.0 9.0 –
SD
SE
CV
Min
Max
n
18.2 – 12.2 – 3.3 3.7 – 1.5 1.8 1.2 2.1 0.6 0.9 4.0 0.2 0.3 0.3 0.3 3.0 1.7 4.7 2.3 14.5 8.3 10.2 12.0 0.0 – 0.0 – 0.0 0.0 – 1.7 – 1.7 – 0.0 – 0.0 – 0.7 – 00.4 – 0.3 0.5 –
5.7 – 3.5 – 1.0 1.1 – 0.5 0.5 0.4 0.6 0.2 0.3 1.2 0.1 0.1 0.1 0.1 0.9 0.5 1.6 0.7 4.6 2.4 3.2 3.5 0.0 – 0.0 – 0.0 0.0 – 0.5 – 0.5 – 0.0 – 0.0 – 0.2 – 0.1 – 0.1 0.1 –
16.6 80.0 146.0 – – – 8.8 120.0 165.0 – – – 10.6 23.0 35.0 11.4 28.0 39.0 – – – 5.2 26.0 31.0 4.9 34.0 39.0 8.8 11.0 14.5 12.6 14.0 20.0 8.3 6.0 8.0 11.6 6.0 8.5 25.7 9.0 24.0 10.2 2.0 2.6 8.8 2.5 3.5 11.8 2.0 2.6 13.1 2.0 2.8 20.9 12.0 20.0 11.2 14.0 20.0 22.3 15.0 31.0 9.5 21.0 27.0 17.5 64.0 112.0 12.2 56.0 84.0 15.7 48.0 80.0 13.3 72.0 115.0 0.0 4.0 4.0 – – – 0.0 2.0 2.0 – – – 0.0 2.0 2.0 0.0 2.0 2.0 – – – 4.6 34.0 40.0 – – – 5.2 30.0 35.0 – 30.0 33.0 0.0 3.0 3.0 – – – 0.0 3.0 3.0 – – – 18.4 2.0 4.0 – – – 10.2 3.0 4.0 – – – 7.7 3.0 4.0 17.0 2.0 3.0 – – –
10 ? 12 ? 10 12 ? 10 12 10 12 10 12 12 4 12 4 12 10 12 9 12 10 12 10 12 10 ? 12 ? 4 12 ? 10 ? 12 ? 10 ? 12 ? 10 ? 12 ? 10 10 ?
Bistichella
551
Table 33 Continued Characteristics a Frontal row III, number of cirri Frontal row IV, number of cirri Frontoventral row V, number of cirri
Frontoventral row VI, number of cirri
Transverse cirri, number Cirral row in posterior body portion, number of cirri Left marginal cirri, number
Right marginal cirri, number
Dorsal kineties, number
Caudal cirri, number
Species
mean
M
SD
SE
CV
Min
Max
n
pro ter bui pro bui hum i pro ter bui hum i pro ter pro ter bui e
3.2 3.0 4.8 5.5 31.8 24.0 22.8 – 25.2 44.0 33.3 26.0 2.1 – 8.2
3.0 – 4.0 5.5 32.5 – 24.0 – 25.5 – 34.0 – 2.0 – 7.5
0.6 – 1.3 0.8 3.7 – 2.5 – 3.5 – 4.4 – 0.4 – 2.7
0.2 – 0.4 0.2 1.2 – 0.7 – 1.1 – 1.3 – 0.1 – 0.9
18.2 2.0 – – 27.3 3.0 14.5 4.0 11.7 25.0 – – 11.0 17.0 – 16.0g 14.1 20.0 – – 13.3 21.0 – – 16.6 2.0 – 2.0 33.5 6.0
4.0 – 7.0 7.0 37.0 – 25.0 18.0 30.0 – 29.0 – 3.0 3.0 15.0
12 ? 9 12 10 1 12 ? 10 1 12 ? 8 ? 10
bui hum i pro ter bui hum i pro ter bui pro ter pro b ter b
44.5 35.0 46.1 – 44.2 41.0 47.7 – 3.0 2.9 4.0 4.9 –
46.0 – 47.0 – 44.5 – 48.0 – 3.0 3.0 – 5.0 –
4.7 – 6.3 – 2.6 – 4.3 – 0.0 0.3 – 1.1 –
1.5 – 1.8 – 0.8 – 1.3 – 0.0 0.1 – 0.3 –
10.6 – 13.7 – 5.9 – 9.1 – 0.0 9.9 – 22.1 –
52.0 – 55.0 37.0 49.0 – 55.0 35.0 3.0 3.0 – 7.0 5.0
10 ? 12 ? 10 ? 12 ? 10 12 ? 12 ?
35.0 – 31.0 32.0 39.0 – 40.0 31.0 3.0 2.0 – 4.0 4.0
a All measurements in µm. CV = coefficient of variation in %, M = median, Max = maximum value, mean = arithmetic mean, Min = minimum value, n = number of individuals investigated, SD = standard deviation, SE = standard error of arithmetic mean, ? = sample size not known (if only one value is known, it is listed in the mean column; if two values are available they are listed as Min and Max). Data based on protargol-impregnated (bui, pro, ter) and opalblue-stained (hum) specimens. b
Whether these are indeed caudal cirri (Fig. 116d, long arrow; 117a) has to be confirmed by ontogenetic data.
c
It is not indicated which macronuclear nodule was measured.
d
It is not indicated which micronucleus was measured.
e
Designated as transverse cirri in original description.
f
Whether these data are from life or from protargol preparations is unclear.
g
This value is from Fig. 117a.
h
Method (from life, after fixation) not indicated.
i
From Fig. 118a (values must not be over-interpreted).
552
SYSTEMATIC SECTION
Table 34 Morphometric data on Bistichella namibiensis (from Foissner et al. 2002) Characteristics a Body, length Body, width Body length:width, ratio Anterior body end to distal end of adoral zone, distance Adoral zone of membranelles, length Length of adoral zone:body length, ratio (%) Distance 1 c Distance 2 c Distance 3 c Distance 4 c Anterior macronuclear nodule, length Anterior macronuclear nodule, width Micronuclei, diameter Macronuclear nodules, number Micronuclei, number Adoral membranelles, number Frontal cirri, number Buccal cirri, number Cirri left of anterior portion of frontoventral row V, number Frontoventral row V, number of cirri Frontoventral row VI, number of cirri Transverse cirri, number Left marginal cirri, number Right marginal cirri, number Dorsal kineties, number Middle dorsal kinety, number of kinetids
mean
M
SD
SE
CV
238.9 234.0 190 189.8b 57.3 53.0 65.9b 67.0 4.3 4.2 2.9b 2.8 31.5 33.0 24.1b 25.0 72.5 69.0 62.8b 62.0 30.9 30.1 53.2 48.0 207.3 200.0 50.5 45.0 220.4 215.0 27.7 28.0 13.2 14.0 4.1 4.0 2.0 2.0 3.6 4.0 43.6 44.0 3.0 3.0 3.7 4.0 4.1 4.0
47.8 20.7 12.3 10.3 1.0 0.4 7.5 5.5 12.3 5.8 4.4 17.8 44.2 16.5 47.3 3.1 1.9 – 0.0 1.1 2.6 0.0 0.7 1.3
12.4 4.7 3.2 2.4 0.3 0.1 1.9 1.3 3.2 1.3 1.1 4.6 11.4 4.3 12.2 0.5 0.5 – 0.0 0.3 0.7 0.0 0.2 0.4
3.5 3.3 0.4 4.4 4.9 0.0 2.2
0.9 0.9 0.1 1.9 1.3 0.0 0.6
39.6 43.1 5.0 50.9 50.2 3.0 29.5
40.0 42.0 5.0 50.0 51.0 3.0 29.0
Min
Max
n
20.0 10.9 21.5 15.6 23.7 15.1 23.6 22.8 17.0 9.2 14.3 33.4 21.3 32.6 21.5 11.1 14.4 – 0.0 29.3 5.9 0.0 18.8 32.8
143.0 152.0 41.0 50.0 2.8 2.3 19.0 15.0 59.0 50.0 25.1 32.0 125.0 24.0 134.0 24.0 10.0 4.0 2.0 2.0 39.0 3.0 2.0 1.0
320.0d 220.0 84.0 90.0 6.3 4.3 44.0 35.0 103.0 70.0 42.0 93.0 294.0 85.0 326.0 35.0 16.0 5.0 2.0 6.0 48.0 3.0 5.0 6.0
15 19 15 19 15 19 15 19 15 19 15 15 15 15 15 15 15 15 15 15 15 15 15 15
8.9 7.7 7.6 8.6 9.7 0.0 7.6
33.0 38.0 4.0 44.0 42.0 3.0 27.0
46.0 50.0 6.0 58.0 61.0 3.0 34.0
15 15 15 15 15 15 15
a
All measurements in µm. Data based, if not otherwise stated, on protargol-impregnated (Wilbert’s method) specimens (specimens deformed by preparation excluded). CV = coefficient of variation in %, M = median, Max = maximum value, mean = arithmetic mean, Min = minimum value, n = number of individuals investigated, SD = standard deviation, SE = standard error of arithmetic mean. b
Cells impregnated with protargol procedure A (Foissner 1991), where they are less distorted.
c
Distance 1 = anterior body end to anterior end of frontoventral row V. Distance 2 = anterior body end to posterior end of frontoventral row V. Distance 3 = anterior body end to anterior end of frontoventral row VI. Distance 4 = anterior body end to posterior end of frontoventral row VI. d
Note that this is not the maximum length (see feature Distance 4).
been described with modern methods, has many (about 32; Fig. 118a) macronuclear nodules and thus cannot be synonymous with B. terrestris or B. procera, which have two nodules. For separation from B. namibiensis, which is also binucleate, see key.
Bistichella
553
Morphology: Note that the following description is based on the combination of the data from three Austrian populations. The morphometry (Table 33) and the infraciliature (Fig. 116c–d) refer to the type population. Body size 170–250 × 30–40 µm in life; average size of type population 139 × 33 µm in protargol preparations (Table 33). Body vermicular, that is, margins parallel or slightly converging posteriad, both ends rounded, sometimes a small notch on right margin immediately ahead of posterior end (Fig. 116b); very flexible (especially under cover glass), slightly to distinctly twisted about main body axis, and only inconspicuously flattened dorsoventrally. Macronuclear nodules 21–28 × 10–14 µm (n = 3) in life, as is usual, slightly left of median. Micronuclei attached to macronuclear nodules. Contractile vacuole near left body margin, distinctly ahead of middle of cell (about at 45% of body length in specimen illustrated; Fig. 116b); during diastole with distinct collecting canals. Cortical granules lacking. Pellicle along ventral and marginal cirral rows crenelated. Cytoplasm colourless, with some greasily shining 0.5–2.0 µm large inclusions, numerous 1–2 µm-sized, colourless globules, and many colourless cloddy particles about 1–8 µm across. Movement without peculiarities. Adoral zone occupies 26% of body length and composed of 33 membranelles on average (Fig. 116a–c, Table 33); bases of largest membranelles about 5–8 µm wide in life (n = 3). Buccal area deep. Undulating membranes distinctly curved in life. Pharyngeal fibres conspicuous both in life and in protargol preparations (Fig. 116a–c). Cirral pattern and number of cirri of usual variability (Table 33). Three distinctly enlarged frontal cirri, in life about 12 µm long; left cirrus usually slightly larger than the middle and right one. Usually four, sometimes three buccal cirri along anterior portion of paroral. 2–4, usually three cirri behind right frontal cirrus (= frontal row III1); this row terminates at 11% of body length on average (Table 33). Frontal row IV composed of 5.5 cirri on average, terminates at 17% of body length; frontoventral row V composed of 23 cirri on average, terminates at 49%; and frontoventral row VI composed of 33 cirri on average and terminates at 65% (Fig. 116c, Table 33). Between posterior end of marginal rows a small group of cirri (transverse cirri?). Right marginal row extends onto dorsolateral surface anteriorly, ends subterminally ahead of supposed transverse cirri (Fig. 116e). Left marginal row commences left of proximal end of adoral zone, terminates at rear cell end. Ventral and marginal cirri in life about 8–10 µm long. Dorsal bristles about 3 µm long, arranged in three kineties; however, kinety 1 usually composed of only a single bristle about at level of buccal vertex; kineties 2 and 3 roughly bipolar (Fig. 116f, d). A short cirral row at posterior dorsal surface (Fig. 116d, f); whether these are caudal cirri or another cirral group can be decided only after studying cell division. Occurrence and ecology: Likely confined to terrestrial habitats of Holarctis and Palaeotropis (Foissner 1987, 1998, Foissner et al. 2002). Unfortunately, the type lo1
In the original description this row was designated as ventral row 1. The remaining three rows right of row III were designated as ventral rows 2–4 in the original description.
554
SYSTEMATIC SECTION
cality of B. procera was incorrectly given in the original description (see remarks). The correct type locality of B. procera is the village of Siebenbrunn in the Marchfeld area east of the city of Vienna (Austria), where we discovered it in a sample from an arable soil (= sample site of population 3 in Berger & Foissner 1987, p. 195). We found it also in the upper (0–2 cm) soil layer of a pasture near the village of Seekirchen, Salzburg, Austria (site E in Foissner et al. 1987a), and in a meadow near the city of Salzburg, where we discovered it in the upper 5 cm soil layer (Berger & Foissner 1987). For a detailed description of this site, see Foissner et al. (1987a, site A). Further record: dry roadside ditch at main road between towns of Aus and Helmeringshausen at the escarpment of the Southern Namib Desert (26°20’S 16°25’E; Foissner et al. 2002, p. 62). Bistichella procera feeds on autotrophic (Euglena sp.) and heterotrophic flagellates, naked amoebas, and ciliates like Colpoda sp. (Berger & Foissner 1987). Biomass of 106 specimens 120 mg (Foissner 1987, p. 127; 1998, p. 208).
Bistichella terrestris (Hemberger, 1985) comb. nov. (Fig. 117a, b, Table 33) 1982 Pseudouroleptus terrestris n. spec.1 – Hemberger, Dissertation2, p. 42, Abb. 5 (Fig. 117a, b; see nomenclature). 1985 Pseudouroleptus terrestris n. spec. – Hemberger, Arch. Protistenk., 130: 399, Fig. Abb. 2 (Fig. 117a, b; original description, see also footnotes on previous entry; no formal diagnosis provided; type slides are deposited in the Institut für landwirtschaftliche Zoologie, University of Bonn, Germany). 2001 Pseudouroleptus terrestris Hemberger, 1985 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 77 (nomenclator containing all basionyms, combinations, and higher taxa of euplotids and hypotrichs).
Nomenclature: No derivation of the name is given in the original description. The species-group name terrestr·is, -is, -e (Latin adjective [m, f, n]; living on the land or in the soil) refers to the habitat where the species was discovered. Remarks: This is the second species assigned to Pseudouroleptus by Hemberger (1985). However, it differs from the type species P. caudatus by the lack of a postperistomial ventral cirrus, strongly indicating that these two species are not closely related. Thus, the present species is transferred, together with some other species, to Bistichella (further details, see genus section). The cirral pattern on the frontal area of B. terrestris shows a peculiarity which is difficult to understand without ontogenetic data. Behind the middle frontal cirrus is a short cirral row, indicating that this row originates from anlage II, like the middle frontal cirrus and the buccal cirri, which form, as in the other Bistichella species, also a short row. Bistichella buitkampi, B. namibiensis, and B. procera also have two short cirral rows (exclusive buccal cirral row) in the frontal area. However, in these 1 2
This name is disclaimed for nomenclatural purposes (ICZN 1999, Article 8.3). See next footnote. See same footnote at Uroleptoides binucleatus.
Bistichella species they are arranged behind the right frontal cirrus (frontal row III) and right of it (frontal row IV). Possibly, the figure provided by Hemberger (1982, 1985) is not quite correct in this respect because he did not use a camera lucida. However, if Fig. 117a is correct then B. terrestris likely produces its frontal-ventraltransverse cirri only from five anlagen, that is, anlage IV would be absent. In the description below the two long rows are designated, as in the other Bistichella-species, as frontoventral rows V and VI. Bistichella terrestris has, like B. procera, some cirri at the posterior cell end. To decide whether these are caudal cirri or another cirral group, cell division data are needed. The most significant difference between these two species is in the number of dorsal kineties (four vs. three [kinety 1 composed of only one bristle]). Hemberger (1985) studied the species mainly after protargol preparations, that is, life data (e.g., presence/absence of cortical granules, consistence [flexible, rigid] of cell) are lacking. Thus, a detailed redescription is needed. Morphology: Body size 190 × 60 µm in life(?), that is, body length:width ratio 3.2:1 and not 4:1 as indicated in the original description. Body outline as shown in Fig. 117a,
555
Fig. 117a, b Bistichella terrestris (from Hemberger 1982, 1985. Protargol impregnation). Infraciliature of ventral side and nuclear apparatus, 190 µm. Short arrow marks cirral row behind middle frontal cirrus; this position indicates that it originates, like the buccal cirri (long arrow), from anlage II. However, this has to be confirmed by ontogenetic data. Frontal cirri connected by dotted line. AZM = adoral zone of membranelles, CC = caudal cirri (?; ontogenetic data needed), CV = contractile vacuole, LMR = left marginal row, MA = posterior macronuclear nodule, MI = micronuclei, P = paroral, RMR = right marginal row, TC = transverse cirri, III = frontal row, V, VI = frontoventral rows. Page 554.
556
SYSTEMATIC SECTION
that is, posterior margins converging, rear end narrowly rounded, but not pointed. Two ellipsoidal macronuclear nodules, each with a micronucleus (Fig. 117b). Contractile vacuole near left cell margin at 30% of body length in specimen illustrated (Fig. 117a). Presence/absence of cortical granules, cell inclusions, and movement not described. Adoral zone occupies about 20–25% of body length, composed of 30–33 membranelles (Fig. 117a, Table 33). Undulating membranes curved, in parallel, one (endoral?) somewhat shorter than the other. Three enlarged frontal cirri arranged in slightly oblique pseudorow with right cirrus, as is usual, at distal end of adoral zone. Buccal cirri right of undulating membranes. Three cirri behind middle frontal cirrus (this position indicates that it originates from the same anlage as the buccal cirri; see remarks). Frontal row III ends at 13% of body length in specimen illustrated. Frontal row IV possibly lacking (see remarks). Frontoventral row V commences slightly behind level of distal end of adoral zone, terminates at 45% of body length in specimen illustrated, whereas row VI ends at 64% (Fig. 117a). Transverse cirri inconspicuous, arranged somewhat subterminally, 12–15 µm long. Right marginal row commences about at level of right frontal cirrus, distinctly shortened posteriorly. Left marginal row begins left of proximal end of adoral zone, almost continuous with cirral group (caudal cirri?) at posterior cell end. Marginal cirri about 10 µm long. Dorsal bristles about 4 µm long, arranged in four kineties; no further details (length of kineties, arrangement) given. 4–5 cirri at posterior cell end, possibly these are caudal cirri. Occurrence and ecology: Bistichella terrestris is very likely confined to terrestrial habitats (Foissner 1987, 1998). The type locality is not given by Hemberger (1985), who mentions only “suspension of mull rendzina soil”. However, this soil sample is not from Peru, the sole country mentioned in the materials and methods section, but from Germany (see Hemberger 1982, p. 2 and Foissner 2000a, p. 259). According to a personal communication by Norbert Wilbert (University of Bonn, Germany), the site is near the village of Mechernich (50°36'N 6°39'E). It is a beech forest on the “Kakushöhle Nordhang” and the outcrops are a middle Devonian dolomite and a Pleistocene tufaceous limestone. No further records published. Food not known. Biomass of 106 specimens 330 mg (Foissner 1987, p. 127; 1998, p. 209).
Incertae sedis in Bistichella Bistichella humicola (Gellért, 1956) comb. nov. (Fig. 118a, b, Table 33) 1956 Uroleptus humicola n. sp. – Gellért, Acta biol. hung., 6: 345, Abb. 10 (Fig. 118a, b; original description; no formal diagnosis provided and very likely no type slides available).
Bistichella
557
1982 Pseudouroleptus humicola1 – Hemberger, Dissertation2, p. 42 (revision of hypotrichs). 1985 Pseudouroleptus humicola – Hemberger, Arch. Protistenk., 130: 399 (combination with Pseudouroleptus; see nomenclature and footnote on previous entry). 2001 Pseudouroleptus humicola (Gellért, 1956) Hemberger, 1985 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 98 (nomenclator containing all basionyms, combinations, and higher taxa of euplotids and hypotrichs).
Nomenclature: No derivation of the species-group name humicola is given in the original description. It is a composite of the Latin noun hum·us (soil), the thematic vowel ·i-, and the Latin verb colere (to live in) and means living in soil. Usually, species-group names ending with -cola are considered as appositive substantives and are thus not changed when transferred to a genus of different gender (Werner 1972, p. 138). Hemberger (1985) did not formally transfer Uroleptus humicola to Pseudouroleptus because he mentioned P. humicola only in a brief remark on P. terrestris. In spite of this I consider Hemberger (1985) as combining author because he recognised the resemblance of these species. Remarks: Bistichella humicola is described mainly after fixed and opalbluestained specimens. Thus, some live data (e.g., presence/absence of cortical granules) are lacking. Further, one cannot be sure that Gellért (1956b) recognised the cirral pattern quite correctly. There are two major differences to the other Bistichella species described after protargol impregnation, namely, the present species has (i) only one buccal cirrus (vs. more than one), and (ii) only one short cirral row (vs. two). In addition, nothing is known about the dorsal kinety pattern. Thus, the assignment of U. humicola to Bistichella is uncertain although the general appearance is strongly reminiscent of B. terrestris (Fig. 117a, 118a). Consequently, I classify Gellért’s species as incertae sedis in Bistichella. For a more proper classification, a detailed redescription including cell division is needed. In spite of the differences to the other Bistichella species, the cirral rows of B. humicola are designated as is usual, that is, the short row, which is obviously behind the right frontal cirrus is designated as frontal row III, and the two long rows are designated as frontoventral rows V and VI. Morphology: Body length 120 µm (after sublimate fixation?); body length:width ratio about 4:1 (Fig. 118a). Body outline elongate elliptical with margins converging posteriorly. Consistency of body not described, likely flexible. Many macronuclear nodules scattered throughout the cell, except in the anterior and posterior portion. Contractile vacuole near left cell margin at 50% of body length; during diastole with collecting canals, pulsates every 13–17 s. Presence/absence of cortical granules, cytoplasmic inclusions, and movement not described. Adoral zone occupies 30% of body length in specimen illustrated, composed of about 31 membranelles (Fig. 118b, Table 33). Paroral (= adoral membrane in original description) long and strongly curved leftwards anteriorly; endoral short, extends from proximal end of adoral zone anteriorly. Between endoral and proximal portion 1 2
This combination is disclaimed for nomenclatural purposes (ICZN 1999, Article 8.3). See next footnote. See same footnote at Uroleptoides binucleatus.
558
SYSTEMATIC SECTION
Fig. 118a, b Bistichella humicola (from Gellért 1956b. Sublimate fixation and opalblue-staining). Infraciliature of ventral side and nuclear apparatus, 120 µm. (b) is a detail from (a). Long arrow in (b) marks the buccal cirrus, short arrow denotes the lateral membranellar cilia. Bistichella humicola has only one short frontal row (III) and one buccal cirrus (long arrow) whereas the other three Bistichella species have two frontal rows and a short buccal cirral row. AZM = distal end of adoral zone of membranelles, CV = contractile vacuole with collecting canals, E = endoral, FC = right frontal cirrus, LMR = anterior end of left marginal row, MA = macronuclear nodules, P = paroral, RMR = anterior end of right marginal row, III = frontal row, V, VI = frontoventral rows. Page 556.
of adoral zone a row of long cilia, likely, the lateral membranellar cilia (Fig. 118b). Protargol preparations are needed for a better description of the oral apparatus. Cirral pattern roughly as in congeners (Fig. 118a, b, Table 33). Three enlarged frontal cirri. One buccal cirrus right of anterior portion of paroral. Frontal row III extends to near level of buccal vertex; frontoventral row V terminates at 64% of body length in specimen illustrated, frontoventral row VI ends at rear cell end. Postperistomial cirrus and transverse cirri obviously lacking. Right marginal row dis-
Bistichella
559
tinctly shortened anteriorly, terminates at rear cell end. Left marginal row commences left of proximal end of adoral zone, slightly shortened posteriorly, that is, rows separated. Dorsal ciliature (length of bristles, number and arrangement of kineties) not known. According to Fig. 118a, caudal cirri likely lacking; however, this has to be checked in protargol preparations. Occurrence and ecology: Likely confined to terrestrial habitats. Type locality of Bistichella humicola is the south-western region of the hill Magoska north-east of the Hungarian village of Boldogkóváralja, where Gellért (1956) discovered it in humus under moss on hypersthenaugitandesit rocks. Record not substantiated by morphological data: chernozem soil from the centre of the Great Hungarian Plain about 15 km south of the main road no. 33 between the villages Hajdúszoboszló and Nagyhegyes, Hungary (Szabó 2000, p. 14). Feeds on debris (Gellért 1956b) and bacteria (Szabó 2000).
Taxa of Unknown Position in the Non-oxytrichid Dorsomarginalia In our monograph on soil ciliates from Namibia we established some oxytrichid genera (Erimophrya, Hemiurosoma, Vermioxytricha) which lack the frontal-ventraltransverse cirri anlage V1 and which have, inter alia for that reason, less than 18 frontal-ventral-transverse cirri (Foissner et al. 2002). However, all taxa lack dorsal kinety fragmentation, the main feature of the Oxytrichidae (Berger 1999), strongly indicating that these three genera are misplaced in the oxytrichids. Since they form a dorsomarginal kinety they belong to the Dorsomarginalia, a large group of hypotrichs comprising all taxa having this feature, which very likely evolved only once (Berger 2006, p. 33, 38). I do not know whether or not the three genera form a monophyletic group, because the reduction or the increase of the number of frontalventral-transverse cirri anlagen obviously occurred several times in the hypotrichs. Examples for a decrease of the number of anlagen are the genera mentioned above (anlage V lost) and Nudiamphisiella (anlage IV lost; see below), examples for an increase are the urostyloids (for review, see Berger 2006) and some oxytrichids, like Paraurostyla (for review, see Berger 1999) and Neokeronopsis (for review, see Berger 2006). Consequently, the lack of anlagen should not be used as apomorphy without support by further features, for example, peculiarities of the dorsal kinety pattern. In addition, the frontoventral cirri of Hemiurosoma are not in the ordinary V-shaped pattern (e.g., Fig. 130h), but arranged as in Urosoma, which has, however, 18 frontal-ventral-transverse cirri and therefore six cirral anlagen (for review, see Berger 1999). For a more detailed discussion of the evolution of major taxa within the Hypotricha, see chapter 2 of the general section. The present chapter also includes Nudiamphisiella, a genus previously classified in the amphisiellids. However, both species included form a dorsomarginal kinety, but lack an oxytrichid kinety fragmentation so that I preliminarily classify it as taxon of unknown position in the non-oxytrichid Dorsomarginalia.
Nudiamphisiella Foissner, Agatha & Berger, 2002 2002 Nudiamphisiella nov. gen.2 – Foissner, Agatha & Berger, Denisia, 5: 693 (original description). Type species (by original designation): Nudiamphisiella interrupta Foissner, Agatha & Berger, 2002.
Nomenclature: Nudiamphisiella is a composite of the Latin adjective nudus (naked) and the genus-group name Amphisiella (see there for derivation); it refers to the lack
1
Note that Foissner et al. (2002) designated the five anlagen I–V, instead of I–IV, VI. Foissner et al. (2002) provided the following diagnosis: Amphisiellidae with several cirri left of amphisiellid median cirral row, which originates from the two rightmost anlagen. The rightmost dorsal kinety develops dorsomarginally. Transverse cirri and postperistomial cirrus lacking.
2
560
Nudiamphisiella
561
of transverse cirri (Foissner et al. 2002). Feminine gender because ending with the Latin diminutive suffix -ella (ICZN 1999, Article 30.1.3). Characterisation (A = supposed apomorphy): Dorsomarginalia with continuous adoral zone of membranelles formed like a question mark. Three frontal cirri. Buccal cirrus present. Left of anterior portion of frontoventral row1 two or more cirri, which originate from anlage III only (A?). Anlage IV lacking (A?). Frontoventral row originates from anlagen V (posterior portion) and VI (anterior portion). Postperistomial cirrus and transverse cirri lacking. One left and one right marginal row. Dorsomarginal kinety present. Dorsal kinety fragmentation lacking. Caudal cirri present. Additional characters: The two species assigned have, in addition to the features mentioned in the previous paragraph, the following, plesiomorphic characteristics in common: body flexible; two macronuclear nodules; contractile vacuole about in mid-body at left cell margin; about 25 adoral membranelles on average; buccal cirrus near anterior end of paroral; frontoventral row terminates roughly in midbody; dorsal bristles 3–4 µm long. Remarks: The type species has a unique combination of features (see corresponding footnote), preventing the assignment to a known genus. Consequently, we established the monotypic genus Nudiamphisiella (Foissner et al. 2002). The frontoventral row, which is distinctly interrupted in N. interrupta, is formed from two anlagen; thus, we assigned it to the Amphisiellidae. Recently I recognised that Amphisiellides illuvialis is likely more closely related to N. interrupta than to A. atypicus, type of Amphisiellides, because both in A. illuvialis and N. interrupta the frontoventral row is formed, as in the amphisiellids, from the two rightmost anlagen against from the rightmost anlage only in A. atypicus. Further, both N. interrupta and A. illuvialis have a dorsomarginal kinety, whereas such a kinety is lacking in A. atypicus. By contrast, the presence of a dorsal kinety fragmentation in A. atypicus requires the transfer of Amphisiellides from the amphisiellids to the oxytrichids for which the dorsal kinety fragmentation is the main apomorphy (Berger 1999). Simultaneously, Amphisiellides illuvialis is (preliminary) assigned to Nudiamphisiella because of the agreements with N. interrupta discussed above. The classification of Nudiamphisiella in the amphisiellids is also uncertain because of the dorsomarginal kinety, which is lacking in all other amphisiellids. Unfortunately, at present we do not know whether the last common ancestor of the amphisiellids never had a dorsomarginal kinety or lost this specific feature. Anyhow, the presence of a dorsomarginal kinety assigns at least Nudiamphisiella to the Dorsomarginalia (Berger 2006, p. 33, 38). The ontogenetic data show that both in N. interrupta and Amphisiellides illuvialis the frontal-ventral cirri anlage IV is lacking because the frontoventral row is formed, as is usual, by the two rightmost anlagen: anlage V forms the posterior portion,
1
The specific term amphisiellid median cirral row is replaced by the neutral term frontoventral row because Nudiamphisiella is very likely not an amphisiellid.
562
SYSTEMATIC SECTION
whereas the anterior portion is formed by anlage VI because the front part of the row is homologous to the frontoterminal cirri of other hypotrichs (Berger 1999, 2006). Hemiamphisiella quadrinucleata also has a dorsomarginal kinety and a cirral pattern which is rather similar to that of N. interrupta (Fig. 62d, e). Thus, one cannot exclude that H. quadrinucleata is more closely related to N. interrupta than to H. terricola (type of Hemiamphisiella), which lacks a dorsomarginal kinety. However, further data on Hemiamphisiella quadrinucleata are needed for a more proper classification. Species included in Nudiamphisiella (alphabetically arranged according to basionyms): (1) Amphisiellides illuvialis Eigner & Foissner, 1994; (2) Nudiamphisiella interrupta Foissner, Agatha & Berger, 2002.
Key to Nudiamphisiella species and a similar species Hemiamphisiella quadrinucleata is very similar to N. interrupta. However, with the following key they can be easily distinguished. 1 Area between posterior portion of marginal rows without cirri (e.g., Fig. 119a, g); cortical granules present (Fig. 119b, j, 62b). . . . . . . . . . . . . . . . . . . . . . . . . . 2 - Area between posterior portion of marginal rows with 1–5, usually 2 longitudinally arranged cirri (Fig. 120b, h–j); cortical granules lacking. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nudiamphisiella illuvialis (p. 569) 2 Two macronuclear nodules (Fig. 119a). . . . . Nudiamphisiella interrupta (p. 562) - 4–6, usually 4 macronuclear nodules (Fig. 62a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemiamphisiella quadrinucleata (p. 318)
Nudiamphisiella interrupta Foissner, Agatha & Berger, 2002 (Fig. 119a–j, Tables 35, 36) 2002 Nudiamphisiella interrupta nov. spec.1 – Foissner, Agatha & Berger, Denisia, 5: 694, Fig. 154a–i, 398k, Table 138 (Fig. 119a–j; original description; the holotype slide [accession number 2002/370] and five paratype slides [2002/343, 351, 371–373] have been deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; see also Aescht 2003, p. 388).
Nomenclature: The species-group name interrupt·us, -a, -um (Latin adjective [m; f; n]; interrupted) refers to the interrupted “amphisiellid median cirral row” (Foissner et al. 2002). Type species of Nudiamphisiella.
1
Foissner et al. (2002) provided the following diagnosis: Size about 150 × 50 µm in vivo; elongate ellipsoidal. Cortical granules globular, colourless to yellowish, form widely spaced rows. On average 2 macronuclear nodules, 27 adoral membranelles, 30 cirri in right and 23 in left marginal row, 14 cirri in discontinuous amphisiellid median cirral row, 3 cirri left of amphisiellid median cirral row, l buccal cirrus, 3 caudal cirri, and 4 dorsal kineties.
Nudiamphisiella
563
Fig. 119a–f Nudiamphisiella interrupta (from Foissner et al. 2002. From life). a: Ventral view of a representative specimen, 150 µm. b: Colourless to yellowish granules 0.5–1.0 µm across form loose rows in cortex. These cortical granules are a major difference to Nudiamphisiella illuvialis, which lacks such granules. c: Cytoplasmic crystals are up to 6 µm long. d–f: Ventral views of shape variants of Benin population, about 100–120 µm long. Page 562.
Remarks: The most similar species, as concerns the ventral and dorsal ciliature, are N. illuvialis and Hemiamphisiella quadrinucleata. Nudiamphisiella interrupta and N. illuvialis – and very likely also H. quadrinucleata – have a dorsomarginal row (Fig. 62e, 119h, i, 120c, r, s), a feature lacking in the amphisiellids. This indicates that these three species do not belong to the Amphisiellidae. However, their position in the Dorsomarginalia Berger, 2006, that is, hypotrichs with a dorsomarginal kinety, is difficult to estimate and therefore meaningful molecular data should be
564
SYSTEMATIC SECTION
awaited before the assignments are adapted (further details, see remarks in genus section and under H. quadrinucleata). In spite of the above mentioned similarity, the three species differ in several features making separation relatively simple (see key and Table 35). Afroamphisiella species, which also lack transverse cirri, do not have a dorsomarginal kinety and caudal cirri (Fig. 74j). Paragastrostyla Hemberger, 1985 – with P. lanceolata Hemberger, 1985 as type species – is a urostyloid and lacks a buccal cirrus and a dorsomarginal kinety (for review, see Berger 2006, p. 613). Morphology: Body size 120–170 × 35–55 µm in life, length:width ratio about 3:1 in life and protargol preparations (Fig. 119a, d–f; Table 36). Body outline elongate elliptical with anterior end often somewhat obliquely truncated and posterior end broadly rounded; left margin usually more convex than right. Body very flexible, but acontractile. Two, rarely four, macronuclear nodules slightly left of midline; individual nodules 20–25 × 10–15 µm in life, that is, ellipsoidal on average, sometimes globular or elongate ellipsoidal; chromatin bodies inconspicuous. Usually one globular, minute micronucleus attached to each macronuclear nodule. Contractile vacuole near mid-body at left cell margin, during diastole without collecting canals. Cortical granules inconspicuous and easily confused with cytoplasmic inclusions, form widely spaced longitudinal rows, do not distinctly impregnate with the protargol method used; when methyl green-pyronin is added, they become red, but are not released (Fig. 119b, j); individual granules inconspicuous because only 0.5–1.0 µm across and almost colourless, most distinct in Benin cells (Fig. 119j), very indistinct in Venezuelan specimens; close beneath cortex ellipsoidal structures, likely mitochondria, as in most Urosoma species (for review, see Berger 1999, p. 396). Cytoplasm colourless and rather strongly granulated, contains ordinary crystals 1–6 µm in size, mainly in posterior body portion, some small lipid droplets, and 7–10 µm-sized food vacuoles. Movement without peculiarities, that is, swims and glides rapidly on microscope slide and debris, showing great flexibility. Adoral zone occupies 19–32%, on average 26% of body length, roughly in Gonostomum pattern (see Berger 1999), that is, extends straight along left body margin, performing right bend and slight clockwise rotation to plunge into buccal cavity; number of adoral membranelles unusually variable (Table 36), bases of largest membranelles about 7 µm wide in life. Buccal cavity flat and narrow. Buccal lip conspicuously projecting and extending from surface of cell at right angles, bears paroral and covers posterior portion of buccal cavity and right half of proximal end of adoral zone (Fig. 119f). Undulating membranes almost straight and both likely composed of a single row of basal bodies; paroral with about 5 µm long cilia, begins on average 3 µm ahead of endoral and is about 8 µm long; endoral usually alongside paroral, about 11 µm long, terminates near proximal end of adoral zone of membranelles. Pharyngeal fibres clearly recognisable in protargol preparations, of ordinary length and structure (Fig. 119a, g). Cirral pattern and number of cirri rather variable (Fig. 119a, g, h; Table 36). All cirri about 15 µm long in life, those of marginal rows and frontoventral row usually
Nudiamphisiella
565
Fig. 119g–i Nudiamphisiella interrupta (from Foissner et al. 2002. Protargol impregnation). g, h: Infraciliature of ventral and dorsal side and nuclear apparatus of holotype specimen, 150 µm. Arrows denote the lateral break separating the anterior and posterior portion of the frontoventral row. The broken line connects the cirri originating from anlage III. The three frontal cirri are connected by a dotted line. i: Infraciliature of ventral side of a late divider, 145 µm (parental structures white, new black). Arrow marks dorsomarginal kinety (= dorsal kinety 4) of opisthe. Note that cirral anlage IV is lacking, because the right frontal cirrus and the cirri behind are formed by anlage III, and the frontoventral row is formed, as is usual, by the anlagen V and VI. The ventral and dorsal morphogenesis proceeds as in N. illuvialis, except that in N. illuvialis some cirri of anlage VI migrate posteriorly. AZM = adoral zone of membranelles, E = endoral, FC = right frontal cirrus, FVR = frontoventral row, LMR = left marginal row, MA = macronuclear nodule, MI = micronucleus, P = paroral, RMR = right marginal row, I–III, V, VI = frontal-ventral cirral anlagen, 1–4 = dorsal kineties (4 = dorsomarginal kinety). Page 562.
composed of two ciliary rows with four cilia each. Frontal cirri distinctly enlarged, form transverse pseudorow; right cirrus, as is usual, behind distal end of adoral zone. Buccal cirrus enlarged (3 × 3–4 basal bodies), right of anterior end of paroral and ahead of endoral. Usually three slightly enlarged cirri behind right frontal cirrus, form short row terminating on average at 25% of body length; few specimens have two short rows (indicating that anlage IV is present in such specimens). Frontoven-
566
SYSTEMATIC SECTION
tral row composed of an anterior portion and a posterior portion, which is shifted by about 5 µm leftward, usually terminating at 53% of body length. Postperistomial cirrus and transverse cirri absent. Right marginal row commences on dorsolateral surface, terminates about 5 µm ahead of rear body end. Left row commences left of proximal end of adoral zone, ends terminally near body midline, that is, marginal rows separated posteriorly (Fig. 119g). Dorsal bristles 3–4 µm long in life, arranged in four kineties (Fig. 119h). Kinety l shortened anteriorly and less densely ciliated than kinety 2, which begins, like kinety 3, near anterior end; kinety 4, a dorsomargiFig. 119j Nudiamphisiella interrupta (from Foissner et nal kinety (Fig. 119i), composed of al. 2002. Interference contrast micrograph; Benin only 4–7 bristles and thus terminatpopulation). Dorsal side of posterior body half showing distinctly ahead of mid-body. ing loose rows of colourless to yellowish cortical granCaudal cirri inconspicuous in life beules 0.5–1.0 µm across (arrowheads). Page 562. cause hardly longer than marginal cirri, composed of 2 × 2 basal bodies; usually one cirrus each associated with bristle rows 1–3, rarely 2–3 cirri with row l and two cirri with row 3. Cell division: We found some dividers and illustrated a late divider (Foissner et al. 2002; Fig. 119i). As in N. illuvialis, five cirral anlagen are recognisable. In late dividers, all cirri of anlage VI (= anlage V in Foissner et al. 2002) migrate anteriad and produce the anterior portion of the frontoventral row, while all cirri of anlage V (= anlage IV in Foissner et al. 2002) form the posterior portion (Fig. 119i). However, the alignment of the two portions is rather incomplete, that is, they are laterally separated by 3–9 µm in interphase specimens (Fig. 119g). Dorsal ontogenesis proceeds also as in N. illuvialis, that is, the anlagen for the dorsal kineties 1–3 develop within the parental rows. By contrast, kinety 4 originates dorsomarginally, that is, from an anlage at/near to the right marginal row (Fig. 119i). This mode of formation corresponds type 2 of Foissner & Adam (1983), respectively, the Urosomoida-pattern of Berger & Foissner (1997) and Berger (1999). The presence of a dorsomarginal kinety indicates that N. interrupta, respectively, Nudiamphisiella is a member of the Dorsomarginalia. Occurrence and ecology: Nudiamphisiella interrupta is likely confined to terrestrial habitats (Foissner et al. 2002, p. 54). To date found only at three sites of the
Nudiamphisiella
567
Table 35 Differences between Nudiamphisiella interrupta, Nudiamphisiella illuvialis, Hemiamphisiella quadrinucleata, and Amphisiellides atypicus Characteristics
N. interrupta
N. illuvialis H. quadrinucleata
A. atypicus
Cirral row between posterior portion of marginal rows
absent
present (longitudinal)
absent
Macronuclear nodules, number Cortical granules
2 present
2 lacking
usually 4 present
distinctly interrupted 2
± continuous 2
± continuous likely 3
present (transverse; thus a pseudorow) about 23 presence/absence not known continuous very likely 1
present absent
present absent
present absent
absent present
Frontoventral row Number of anlagen forming the frontoventral row Dorsomarginal kinety Dorsal kinety fragmentation
Table 36 Morphometric data on Nudiamphisiella interrupta (from Foissner et al. 2002) Characteristics a
mean
M
Body, length 136.0 138.0 Body, width 45.4 45.0 Body length:width, ratio 3.0 3.1 Adoral zone of membranelles, length 35.1 36.0 Body length:length of adoral zone, ratio 3.9 3.8 Anterior cell end to buccal cirrus, distance 15.2 16.0 Anterior cell end to short frontal cirral row, distance f 10.2 10.0 Anterior cell end to end of short cirral row, distance f 23.1 23.0 Anterior cell end to frontoventral row, distance 6.0 6.0 Anterior cell end to end of anterior portion of 32.3 32.0 frontoventral row, distance Interruption in frontoventral row, lateral distance b 5.3 5.0 Anterior cell end to posterior portion of 34.8 34.0 frontoventral row, distance Anterior cell end to end of frontoventral row, 72.0 74.0 distance Body length:length of frontoventral row, ratio 1.9 1.8 Anterior cell end to right marginal row, distance 7.2 8.0 Posterior cell end to right marginal row, distance 5.0 5.0 Anterior cell end to paroral, distance 16.8 17.0 Paroral, length 8.0 8.0 Anterior cell end to endoral, distance 19.3 20.0 Endoral, length 11.0 11.0 Posterior cell end to middle caudal cirrus, distance 3.0 2.0 Anterior cell end to first macronuclear nodule, 28.6 30.0 distance Nuclear figure, length 68.7 70.0 Macronuclear nodules, distance in between 26.2 26.0 Anterior macronuclear nodule, length 20.9 22.0 Anterior macronuclear nodule, width 9.2 9.0 Anterior macronuclear nodule length:width, ratio 2.3 2.3 Posterior macronuclear nodule, length 21.6 22.0
SD
SE
CV
Min
Max
n
13.7 5.9 0.4 4.1 0.6 2.6 2.1 4.3 1.8 5.6
2.9 1.2 0.1 0.9 0.1 0.5 0.4 0.9 0.4 1.2
10.1 111.0 158.0 13.0 35.0 58.0 13.1 2.4 3.8 11.7 27.0 44.0 14.2 3.1 5.2 17.0 11.0 20.0 20.9 6.0 14.0 18.5 16.0 32.0 31.0 3.0 9.0 17.4 17.0 46.0
23 23 23 23 23 22 23 22 23 23
1.6 4.4
0.3 0.9
29.1 12.6
3.0 24.0
9.0 44.0
23 23
10.2
2.1
14.2
50.0
91.0
23
0.3 1.8 2.0 3.0 1.1 3.0 1.6 1.6 3.5
0.1 0.4 0.4 0.6 0.2 0.6 0.3 0.3 0.7
16.3 24.3 40.3 17.7 14.3 15.7 14.1 53.2 12.3
1.5 4.0 2.0 11.0 6.0 14.0 8.0 1.0 19.0
3.0 10.0 6.0 22.0 10.0 25.0 15.0 6.0 34.0
23 23 22 23 23 23 22 23 23
10.1 6.4 4.5 1.3 0.7 4.9
2.1 1.3 0.9 0.3 0.1 1.0
14.6 24.2 21.8 14.6 28.1 22.8
38.0 17.0 10.0 6.0 1.1 10.0
84.0 38.0 28.0 12.0 4.0 30.0
23 23 23 23 23 23
568
SYSTEMATIC SECTION
Table 36 Continued Characteristics a Posterior macronuclear nodule, width Macronuclear nodules, number c Anterior micronucleus, diameter Micronuclei attached to anterior macronuclear nodule, number Micronuclei attached to posterior macronuclear nodule, number Micronuclei, total number Adoral membranelles, number Frontal cirri, number Buccal cirri, number d Short frontal row, number of cirri f Anterior part of frontoventral row, number of cirri Posterior part of frontoventral row, number of cirri Frontoventral row, total number of cirri Right marginal cirri, number Left marginal cirri, number Caudal cirri, number Dorsal kineties, number Kinetids in kinety 1, number Kinetids in kinety 2, number Kinetids in kinety 3, number Kinetids in kinety 4, number
mean
M
SD
SE
CV
Min
Max
n
9.6 2.0 2.8 1.2
9.0 2.0 3.0 1.0
1.4 0.0 – –
0.3 0.0 – –
15.1 0.0 – –
7.0 2.0 2.5 1.0
12.0 2.0 3.0 2.0
23 22 23 23
1.0
1.0
–
–
–
0.0
2.0
23
2.2 26.8 3.0 1.0 2.7 6.9 7.0 13.9 30.0 23.7 3.3 4.0 11.9 17.4 14.4 5.5
2.0 27.0 3.0 1.0 3.0 7.0 7.0 14.0 30.0 24.0 3.0 4.0 12.0 17.0 14.0 5.5
0.7 2.8 0.0 0.0 0.6 1.2 1.4 2.0 4.2 2.5 – 0.0 1.0 1.1 1.6 0.9
0.1 0.6 0.0 0.0 0.1 0.3 0.3 0.4 0.9 0.5 – 0.0 0.4 0.4 0.6 0.3
30.3 10.5 0.0 0.0 24.4 17.9 19.7 14.2 14.1 10.6 – 0.0 8.3 6.5 11.1 16.8
1.0 21.0 3.0 1.0 1.0 4.0 4.0 9.0 20.0 19.0 3.0 4.0 11.0 16.0 12.0 4.0
4.0 32.0 3.0 1.0 4.0 9.0 10.0 19.0 37.0 27.0 6.0 4.0 14.0 19.0 17.0 7.0
23 23 23 22 23 23 23 23 23 23 23 23 8 9 8 8
a
All measurements in µm. Data based on mounted, protargol-impregnated (Foissner’s protocol), and randomly selected specimens from a non-flooded Petri dish culture. CV = coefficient of variation in %, M = median, Max = maximum, mean = arithmetic mean, Min = minimum, n = number of individuals investigated, SD = standard deviation, SE = standard error of arithmetic mean. b
See Fig. 119g (arrows).
c
Rarely specimens with four macronuclear nodules occur.
d
Rarely specimens without buccal cirrus occur.
e
Right frontal cirrus not included.
f
The short row is formed by anlage III (= paramalar cirral row).
southern hemisphere (Foissner et al. 2002), namely (i) at the type locality (site 49 in Foissner et al. 2002), that is, the Bambatsi Guest Farm (20°10’S 15°25’E; altitude 1150 m) between the towns of Khorixus and Outjo, Namibia, where we discovered it with high abundance in mud and soil from road puddles; (ii) in a soil sample from near the National University of Benin (collected by Jean Dragesco; Fig. 119d–f, j); and (iii) in a soil sample from Venezuela. Nudiamphisiella interrupta feeds on bacteria (Foissner et al. 2002).
Nudiamphisiella
569
Nudiamphisiella illuvialis (Eigner & Foissner, 1994) comb. nov. (Fig. 120a–t, Tables 35, 37) 1994 Amphisiellides illuvialis n. sp.1 – Eigner & Foissner, J. Euk. Microbiol., 41: 244, Fig. 1–24, Table 1 (Fig. 120a–t; original description; according to the original description, the holotype slide and a paratype slide [accession numbers 1994/10, 1994/11] have been deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; according to Aescht 2003, p. 388, four syntypes [1993/10, 11, 44, 103] have been deposited). 2001 Amphisiellides illuvialis Eigner and Foissner, 1994 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 8 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Amphisiellides illuvialis – Lynn & Small, Phylum Ciliophora, p. 453, Fig. 37A–C (Fig. 120b, c; guide to ciliate genera).
Nomenclature: The species-group name illuvialis (Latin adjective; living in dirty water) is a composite of the Latin noun illuvies (dirty water, morass) and the suffix -al·is (living in), and refers to the habitat (litter of disused pigpen) where the species was discovered (Eigner & Foissner 1994). Lynn & Small (2002) used the term midventral cirral file to designate the amphisiellid median cirral row. This is somewhat misleading because the term midventral (cirri/rows/complex) should be confined to the urostyloids (for review, see Berger 2006). Remarks: Eigner & Foissner (1994) assigned this species to Amphisiellides because it has (i) transverse and caudal cirri and (ii) more than one cirrus left of the anterior portion of the frontoventral row (note that these cirri are from anlage III only, and not from anlagen III and IV as in other species with more than one cirrus left of the anterior portion of the frontoventral row). As major difference to the type species A. atypicus they mentioned the nuclear apparatus (two macronuclear nodules vs. 23). Further differences exist in the arrangement of the transverse cirri (longitudinal [note that these cirri are not true transverse cirri] vs. transverse), the body size (100 µm vs. 200 µm), the number of cirri in the frontoventral row and the marginal rows (see Table 35), and the morphogenesis (Eigner & Foissner 1994). Interestingly, Eigner & Foissner (1994) did not discuss two important morphogenetic features of the type species of Amphisiellides, namely, (i) that the frontoventral row obviously originates from the rightmost anlage only against from the two rightmost anlagen in the present species, and (ii) that very likely dorsal kinety fragmentation occurs and a dorsomarginal kinety is lacking in A. atypicus, whereas fragmentation is lacking and a dorsomarginal kinety is present in A. illuvialis. These differences are rather significant and strongly indicate that these two species are not closely related. This assumption is supported by the different transverse cirri. The type species has true transverse cirri (enlarged and originating from different anlagen), whereas the longitudinal row of cirri in the present species can hardly be des1
Eigner & Foissner (1994) provided the following diagnosis: Lanceolate, in vivo 50–140 × 20–50 µm. 2 macronuclear segments, 25 adoral membranelles, 1 buccal cirrus at anterior end of paroral membrane, 2 cirri right and 3 cirri left of amphisiellid cirral row, 2 transverse cirri, and 3 caudal cirri on average.
570
SYSTEMATIC SECTION
Nudiamphisiella
571
ignated as transverse cirral row, except for the rearmost cirrus (for terminology, see chapter 1.7 in general section). All these differences lead me to transfer A. illuvialis to Nudiamphisiella. However, the classification in Nudiamphisiella is possibly also not definite because there are some differences to the type species, for example, in the shape of the undulating membranes (short and more or less straight in N. interrupta [Fig. 119g] vs. long and curved in N. illuvialis [Fig. 120b]). In life, the present species is difficult to separate from binucleate members of the amphisiellids. They differ in the presence/absence of certain cirral groups (e.g., transverse cirri, caudal cirri, number of cirri left of anterior portion of amphisiellid median cirral row, number and arrangement of dorsal kineties). Thus, protargol preparation is needed to recognise the ventral and dorsal infraciliature correctly. Morphology: Eigner & Foissner (1994) found two populations (see occurrence and ecology section). The German population was very similar to the type material and was not studied in detail. Body size in life highly variable, that is, 50–140 × 20–50 µm, on average around 120 × 30 µm; body length:width ratio 3.7:1 in protargol preparations (Table 37). Body outline usually lanceolate, that is, distinctly tapering anteriorly and posteriorly (Fig. 120a); rarely as shown in Fig. 120d–f. Body dorsoventrally flattened by about 2:1 (Fig. 120g); highly flexible. Macronuclear nodules left of midline, ellipsoidal, shape and size highly variable (Fig. 120a, c); chromatin bodies in life 1.5–2.5 µm across. Several micronuclei 2.0–3.5 µm across, adjacent to macronuclear nodules, only faintly stained by protargol. Contractile vacuole slightly ahead of mid-body near left cell margin (Fig. 120a). Cytoplasm colourless, about 4 µm long yellowish crystals mainly in posterior body portion. Conspicuous cortical granules lacking. Food vacuoles 10–20 µm across. Movement not described, likely without peculiarities. Adoral zone occupies about 30% of body length, composed of an average of 25 membranelles of ordinary fine structure. Bases of largest membranelles in life about 8 µm wide, cilia of membranelles up to 13 µm long. Buccal cavity inconspicuous, that is, flat and narrow, buccal lip slightly curved, parallels undulating membranes (Fig. 120a, b). Cirral pattern and number of cirri rather variable (Fig. 120a, b, h–j, l, Table 37). All cirri about 10 µm long. Three slightly enlarged frontal cirri almost transversely arranged; right cirrus, as is usual, near distal end of adoral zone. Usually one buccal cirrus right of anterior end of paroral. Frontoventral row commences near right fron-
b Fig. 120a–g Nudiamphisiella illuvialis (from Eigner & Foissner 1994. a, d–g, from life; b, c, protargol impregnation). a: Ventral view of a representative specimen, 105 µm. b, c: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 80 µm. Short arrows mark cirri right of frontoventral row, long arrow denotes cirral row (designated as transverse cirri in original description) between posterior portion of marginal rows. Frontal cirri connected by dotted line. d–f: Ventral view of rare shape variants. g: Right lateral view. AZM = adoral zone of membranelles, BC = buccal cirrus, CC = caudal cirri, CV = contractile vacuole, FC = right frontal cirrus, FVR = rear end of frontoventral row, LMR = left marginal row, MA = macronuclear nodule, MI = micronucleus, RMR = right marginal row, 1–4 = dorsal kineties (4 = dorsomarginal kinety). Page 569.
572
SYSTEMATIC SECTION
Nudiamphisiella
573
tal cirrus, terminates at 54% of body length on average (Table 37). Usually 2–3 cirri left of anterior portion of frontoventral row (originating from anlage III only!); in about 3% of specimens at distinct row left of frontoventral row (Fig. 120i). About 45% of specimens without ventral cirri right of frontoventral row (Fig. 120h), about 45% with 1–2 cirri right of frontoventral row (Fig. 120b), and about 10% with a conspicuous row right of the frontoventral row (Fig. 120j); rarely, specimens occur with few cirri right of this additional row (Fig. 120j). Usually a longitudinal row of two (rarely of up to five) cirri between rear portion of marginal rows (Fig. 120b, h–j); occasionally two such rows present (Fig. 120l). Eigner & Foissner (1994) designated all these cirri as transverse cirri, which is, however, misleading because a transverse cirrus is, per definition, only the posteriormost, very often slightly to distinctly enlarged and set off cirrus of an anlage. Thus, only the rearmost cirrus of this row can be designated as transverse cirri (see general section for details on terminology). Postperistomial cirrus and distinct transverse cirri lacking. Right marginal row commences on dorsolateral surface distinctly behind anterior body end, extends – like left row – almost to rear cell end leaving small gap; left row commences left of proximal portion of adoral zone (Fig. 120b). Dorsal cilia 3–4 µm long, arranged in four kineties. Kineties 1–3 distinctly shortened anteriorly, kinety 4 – a dorsomarginal row – only slightly shortened anteriorly, terminates, however, at 28% of body length in specimen illustrated (Fig. 120c). Occasionally a faintly stained second dorsomarginal row from previous generation. Caudal cirri usually at posterior end of kineties 1 and 2 (Fig. 120c, s). Cell division (Fig. 120k–t): This part of the life cycle is described in great detail by Eigner & Foissner (1994). Four micrographs of a non-divider and of three dividing stages document the most important features of the process (not shown in present revision). Note that Eigner & Foissner (1994) did not use the Wallengren system to designate the cirral anlagen. Their anlagen 1–5 correspond the anlagen I–III, V, VI in the present review, that is, anlage IV is, like in the type species, lacking in N. illuvialis because the frontoventral row is formed, as is usual, by the anlagen V and VI.
b Fig. 120h–n Nudiamphisiella illuvialis (from Eigner & Foissner 1994. Protargol impregnation). Variability of cirral pattern (h–j) and infraciliature of ventral side of dividing specimens (k–n). Note that in N. illuvialis anlage IV is lacking because the cirri left of the anterior portion are from anlage III and the frontoventral row is formed by the anlagen V and VI. h: Specimen with only one transverse cirrus. i: Specimen with a distinct cirral row behind the right frontal cirrus. The rearmost cirrus of anlage VI can be designated as transverse cirrus; the cirrus ahead of it has to be designated as pretransverse ventral cirrus. j: Specimen with distinct cirral row right of frontoventral row. Some cirri are produced by an additional anlage right of anlage VI. k: Very early divider (105 µm) showing that the formation of the oral primordium (arrow) commences left of the posterior portion of the frontoventral row. l: Early divider (105 µm) showing two streaks (arrowheads) extending from oral primordium. Arrow marks an anlage which originates from cirri of the frontoventral row or, less likely, de novo. m: Early to middle divider (125 µm) showing opisthe’s anlagen I–III, V, VI. The streak (arrow) right of the frontoventral row appears unchanged as compared to the previous stage. n: Middle divider (114 µm) showing five anlagen per filial product. Marginal primordia are recognisable. FVR = frontoventral row, I–III, V, VI = frontal-ventral-(transverse) cirri anlagen. Page 569.
574
SYSTEMATIC SECTION
Nudiamphisiella
575
The division commences with the proliferation of basal bodies near the left side of the posterior cirri of the frontoventral row, which appears unchanged (Fig. 120k). Later, a large, wedge-shaped primordium develops in the central body portion. Very likely, cirri of the posterior portion of the frontoventral row are disaggregated and involved in the primordium. Two streaks originate from the right anterior end of the oral primordium and extend left and right of the frontoventral row (Fig. 120l, arrowheads). A third streak develops right of the frontoventral row from a modified cirrus of the parental row or, possibly, de novo (Fig. 120l, m, arrow). In a later stage, five frontal-ventral cirral anlagen are recognisable (Fig. 120m). The anlage right of the frontoventral row is almost unchanged. The parental buccal cirrus has modified to a short streak. In the next stage, five anlagen are recognisable each in the proter and the opisthe (Fig. 120n–p). The anlagen for the proter develop slightly later than those of the opisthe, whose anlagen originate from the oral primordium only. By contrast, anlage I of the proter originates from the parental undulating membranes; anlage II from the buccal cirrus; anlage III from the cirri behind the right frontal cirrus (= cirri left of the anterior portion of the frontoventral row); anlage IV is obviously lacking; anlagen V and VI from the middle portion (rear segment) of the frontoventral row (the exact origin of anlage VI is not known; possibly it is formed de novo). Some cirri may develop between the posterior portions of the anlagen V and VI (Fig. 120p); likely they form the second row between the rear end of the marginal rows found in about 3% of specimens (Fig. 120l). In late dividers, the new cirri arrange in the species-specific manner. Some cirri of the (new) anlagen II, III, V, and VI are resorbed during late stages so that their number is smaller during interphase than in dividers. The anlagen produce the following parts of the infraciliature (note that the pattern is, as expected, very similar to that of other species): anlage I forms the endoral, the paroral, and the left frontal cirrus; anlage II forms the middle frontal cirrus and the buccal cirrus; anlage III forms the right frontal cirrus and the cirri behind of it (= cirri left of anterior portion of frontoventral row); anlage V (= anlage 4 in Eigner & Foissner 1994) forms the posterior part of the frontoventral row; anlage VI forms the anterior part (homologous
b Fig. 120o–t Nudiamphisiella illuvialis (from Eigner & Foissner 1994. Protargol impregnation). Infraciliature of ventral (o, p, r, t) and dorsal (q, s) side of dividing specimens. Parental structures white, new black. Broken lines connect cirri originating from same anlage (not shown everywhere). o: Middle to late divider (83 µm) with five anlagen per filial product. Arrows mark anlagen for dorsomarginal kinety (= dorsal kinety 4). p: Detail of cirral formation and migration. Note that the anterior portion of anlage VI (homologous to frontoterminal cirri) forms the anterior portion of the frontoventral row. White arrow marks two cirri which very likely form a second row of cirri between the posterior portion of the marginal rows (cp. Fig. 120l). q, s: Dorsal morphogenesis (q = 76 µm, s = 70 µm) is interesting in N. illuvialis because a dorsomarginal kinety (arrows) is formed (details see species description). Arrowhead in (q) denotes parental dorsomarginal kinety. r: Late divider (115 µm) showing formation of final cirral pattern. Arrows mark anlagen for dorsomarginal kinety; frontoterminal cirri, which form the anterior portion of the frontoterminal row, are circled t: Postdivider (76 µm) showing that alignment of anterior and posterior portion of frontoventral row is just finished after cytokinesis. MA = fused macronucleus, I–III, V, VI = frontal-ventral(transverse) cirri anlagen. Page 569.
576
SYSTEMATIC SECTION
Table 37 Morphometric data on Nudiamphisiella illuvialis (from Eigner & Foissner 1994) Characteristics a Body, length Body, width Adoral zone of membranelles, length Anterior body end to rear end of frontoventral row, distance Anterior macronuclear nodule, length Anterior macronuclear nodule, width Posterior macronuclear nodule, length Posterior macronuclear nodule, width Macronuclear nodules, number Adoral membranelles, number Buccal cirri, number Cirri left of anterior portion of frontoventral row, number b Frontoventral row, number of cirri Cirri right of frontoventral row, number Cirri between rear portion of marginal rows, number c Left marginal cirri, number Right marginal cirri, number Caudal cirri, number
mean
M
SD
SE
CV
Min
Max
n
98.0 103.0 26.7 26.5 28.7 29.0 53.6 51.0
14.3 6.8 2.7 12.3
– – – –
14.6 25.6 9.5 23.0
56.0 132.0 16.0 46.0 24.0 35.0 19.0 80.0
28 28 28 28
24.3 7.2 21.2 7.0 2.0 24.6 1.0 3.0
27.0 7.0 19.5 6.0 2.0 25.0 1.0 3.0
7.0 2.0 10.4 2.3 – 2.7 – 1.5
– – – – – – – –
28.8 27.6 49.3 33.5 – 10.8 – 50.0
9.0 4.0 9.0 4.0 1.0 21.0 1.0 1.0
32.0 12.0 40.0 12.0 4.0 35.0 2.0 6.0
27 27 12 12 28 28 28 14
13.0 2.1 2.3
12.0 0.0 2.0
3.6 3.8 0.8
– – –
27.6 181.0 34.8
5.0 0.0 1.0
27.0 13.0 5.0
28 28 25
19.4 25.6 3.5
20.5 26.0 3.0
4.3 4.3 1.1
– – –
22.0 16.7 31.4
8.0 15.0 2.0
27.0 36.0 7.0
28 28 28
a All measurements in µm. CV = coefficient of variation in %, M = median, Max = maximum value, mean = arithmetic mean, Min = minimum value, n = number of individuals investigated, SD = standard deviation, SE = standard error of arithmetic mean. Data based on protargol-impregnated specimens. b
Cirrus III/2 included.
c
Designated as transverse cirri by Eigner & Foissner (1994; see remarks and description).
to the frontoterminal cirri of other hypotrichs) of the frontoventral row and the row (= transverse cirri according to Eigner & Foissner 1994) between the posterior portion of the marginal rows; the cirri right of the frontoventral row present in about 55% of specimens are also produced by anlage VI. In very late dividers, the frontoventral row is more or less perfectly aligned, and the rear portion of anlage VI migrates posteriorly (Fig. 120r, t). The division of the marginal rows proceeds in the plesiomorphic mode, that is, within each parental row two primordia occur (Fig. 120n, o, r). However, from/near the anterior end of the right marginal row primordium the dorsomarginal kinety (= dorsal kinety 4) is formed (Fig. 120o, q–s). The presence of a dorsomarginal kinety indicates that Nudiamphisiella illuvialis does not belong to the amphisiellids (see genus section). Caudal cirri develop at the rear end of kineties 1 and 2, rarely also on kinety 3. As is usual, the dorsomarginal row does not form a caudal cirrus. The nuclear apparatus divides plesiomorphically, that is, the two macronuclear nodules fuse to a single mass and later divide twice (Fig. 120q, r).
Erimophrya
577
Occurrence and ecology: Terrestrial and limnetic (Foissner 1998, p. 199). Type locality is the village of Schrötten, Deutsch Goritz (46°47’N 15°49’E; 320 m altitude), Styria, Austria, where Eigner & Foissner (1994) discovered it in a litter sample from a disused pigpen. They found a second population near the village of Eching (Bavaria, Germany) in a tiny track puddle which was heavily polluted by sewage from an activated sludge plant. No further records published. Feeds on bacteria and ciliates (Eigner & Foissner 1994). Biomass of 106 specimens about 38 mg (Foissner 1998, p. 199).
Erimophrya Foissner, Agatha & Berger, 2002 2002 Erimophrya nov. gen.1 – Foissner, Agatha & Berger, Denisia, 5: 791 (original description). Type species (by original designation): Erimophrya glatzeli Foissner, Agatha & Berger, 2002.
Nomenclature: Erimophrya is a composite of the Greek nouns erimos (desert) and ophrya (eyebrow l cilium l ciliate), meaning a ciliate type occurring in deserts. Feminine gender (Foissner et al. 2002). Characterisation (A = supposed apomorphy): Dorsomarginalia with continuous (type) or bipartite adoral zone of membranelles. Undulating membranes roughly in Oxytricha pattern. Three frontal cirri. Buccal cirrus present. Frontoventral cirri arranged in V-shaped pattern. Frontal-ventral-transverse anlage V lost and anlagen II–IV, VI form less than the plesiomorphic number of 17 cirri so that the number of postoral ventral cirri, pretransverse ventral cirri, and transverse cirri is reduced (A?). One right and one left marginal row. Dorsomarginal kinety present. Dorsal kinety fragmentation lacking. Caudal cirri present. Terrestrial. Additional characters: The four species are rather similar and therefore have several further (plesiomorphic?) features in common: body length 90–110 µm on average in life. Body acontractile, flexible. Two or four (E. quadrinucleata) macronuclear nodules in middle body third left of midline. Buccal cavity narrow and flat. Undulating membranes short, slightly curved and indistinctly intersecting. All cirri of about same size. Buccal cirrus right of anterior end of paroral. Transverse cirri very close to posterior body end. Dorsal bristles 2–4 µm long, rather widely spaced. Two caudal cirri. Remarks: Erimophrya was established for two Urosomoida-like hypotrichs from dunes of the Namib Desert because they have only five frontal-ventral-transverse cirri anlagen; in addition, the number of postoral ventral cirri, pretransverse ventral cirri, and transverse cirri is more strongly reduced than in Urosomoida Hemberger in Foissner, 1982 (Foissner et al. 2002; for review of Urosomoida see Berger 1999, p. 345). A few years later, two further species clearly belonging to Erimophrya were 1
Foissner et al. (2002) provided the following diagnosis: Oxytrichidae with oral apparatus in Oxytricha pattern and V-like arranged frontoventral cirri. Number of postoral and transverse cirri distinctly reduced. 1 right and 1 left row of marginal cirri. 3 or 4 dorsal kineties. Caudal cirri present. Ontogenesis in Urosomoida pattern, but with only 5 fronto-ventral-transverse cirral anlagen streaks.
578
SYSTEMATIC SECTION
discovered in Austrian pine forest soils (Foissner et al. 2005). The four species are very similar, especially in life and at superficial observation, but they can be easily distinguished in protargol preparations, mainly by morphometric features, such as the number of macronuclear nodules, postoral ventral cirri, transverse cirri, and dorsal kineties (Table 38). Vermioxytricha lacks postoral ventral cirri, transverse cirri, and caudal cirri and has only two dorsal kineties (p. 596). For separation of Erimophrya from Hemiurosoma see there (p. 614). Hemisincirra (p. 387) lacks caudal cirri and, more important, very likely does not form a dorsomarginal kinety, a feature clearly assigning Erimophrya, Vermioxytricha, and Hemiurosoma to the Dorsomarginalia. Since the ontogenesis and the general cirral pattern of Erimophrya are similar to those of the 18-cirri hypotrichs, it was assigned to the Oxytrichidae by Foissner et al. (2002). For a foundation of the exclusion from the oxytrichids, see introduction to nonoxytrichid Dorsomarginalia (p. 560). The loss of a frontal-ventral-transverse cirri anlage is a striking feature. Since I do not know at which level it is an apomorphy (e.g., for Erimophrya and other taxa, like Vermioxytricha and Hemiurosoma, separately or once for all these taxa) I do not mark it as such in the characterisation above. 18-cirri hypotrichs have three postoral ventral cirri originating from two anlagen, namely, anlage IV (cirrus IV/2) and anlage V (V/3, V/4) (Fig. 3a). Three Erimophrya species have two or three postoral ventral cirri (Table 38). However, since anlage V is lacking in this genus they are formed exclusively from anlage IV. Thus, the three postoral ventral cirri of, for example, Erimophrya quadrinucleata (Fig. 125c, f) are not homologous to the three postorals of the ordinary 18-cirri hypotrichs. Which of the three cirri in E. quadrinucleata is homologous to IV/2 of the 18cirri hypotrichs cannot be said unequivocally; I suppose the anteriormost. Anyhow, it is a further example of convergence in the hypotrichs. However, a realised convergence does not cause problems in systematics. Urosomoida antarctica Foissner, 1996b also lacks one postoral ventral cirrus, but has the full set of two pretransverse ventral cirri and five transverse cirri, proving that it forms six (I–VI) anlagen during cell division (for review, see Berger 1999, p. 365). Thus, it is likely not closely related to Erimophrya. Urosomoida monostyla Foissner, Agatha & Berger, 2002 (p. 784) and U. deserticola Foissner, Agatha & Berger, 2002 (p. 787) also have only one or two postoral ventral cirri, indicating a relationship to the present genus. However, ontogenetic data should be awaited for a more proper classification. Species included in Erimophrya (alphabetically arranged basionyms are given): (1) Erimophrya arenicola Foissner, Agatha & Berger, 2002; (2) Erimophrya glatzeli Foissner, Agatha & Berger, 2002; (3) Erimophrya quadrinucleata Foissner, Berger, Xu & Zechmeister-Boltenstern, 2005; (4) Erimophrya sylvatica Foissner, Berger, Xu & Zechmeister-Boltenstern, 2005.
Erimophrya
579
Key to Erimophrya species Note that the differences between the four species described so far are not very conspicuous, that is, identification needs detailed live observation and protargol impregnation because the number of postoral ventral cirri, transverse cirri, and dorsal kineties are important. If you cannot identify your specimen/population with the key below, see also Hemisincirra (transverse cirri present; p. 387), Hemiurosoma (transverse and caudal cirri present, four dorsal kineties; see p. 614 or Foissner et al. 2002, p. 834), and Vermioxytricha (postoral ventral cirri, pretransverse ventral cirri, transverse cirri, and caudal cirri lacking; p. 596). 1 2 3 -
Four macronuclear nodules (Fig. 125a, b, g) Erimophrya quadrinucleata (p. 595) Two macronuclear nodules. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Four dorsal kineties (Fig. 121a–g). . . . . . . . . . . . . . . Erimophrya glatzeli (p. 579) Three dorsal kineties (Fig. 122f, 123c). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Two postoral ventral cirri; two transverse cirri; cortical crystal reticulum present (Fig. 123a–f, 124a, b). . . . . . . . . . . . . . . . . . . . . . . Erimophrya sylvatica (p. 590) One postoral ventral cirrus; one transverse cirrus; cortical crystal reticulum absent (Fig. 122a–f). . . . . . . . . . . . . . . . . . . . . . . . . . Erimophrya arenicola (p. 586)
Erimophrya glatzeli Foissner, Agatha & Berger, 2002 (Fig. 121a–g, Table 38) 2002 Erimophrya glatzeli nov. spec.1 – Foissner, Agatha & Berger, Denisia, 5: 791, Fig. 172a–g, Table 153 (Fig. 121a–g; original description; the holotype slide [accession number 2002/246] and two paratype slides [2002/247, 248] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; see Foissner et al. 2002, p. 39 and Aescht 2003, p. 387).
Nomenclature: Wilhelm Foissner dedicated this species to Gerhard Glatzel, a famous forest ecologist at the University of Vienna, Austria (Foissner et al. 2002). Type species of Erimophrya. Remarks: Erimophrya glatzeli differs from E. arenicola, which occurs in the same area, in the features given in the key plus an ontogenetic feature: Erimophrya arenicola lacks the anarchic field of basal bodies left of the transverse cirrus. Taken together, the differences allow a clear separation of these two species. However, in life E. glatzeli is easily confused with medium-sized hypotrichs, especially Urosomoida agiliformis (for review see Berger 1999, p. 356). Thus, check the number of postoral ventral cirri (two vs. three), which are easily recognisable with differential interference contrast. 1
Foissner et al. (2002) provided the following diagnosis: Size about 110 × 25 µm in vivo; elongate elliptical. On average 2 widely separated, elongated ellipsoidal macronuclear nodules, 21 adoral membranelles, 20 cirri each in right and left marginal row, 2 postoral cirri, 1 transverse cirrus, 2 caudal cirri, and 4 dorsal kineties.
580
SYSTEMATIC SECTION
Erimophrya
581
Morphology: Body size 80–150 × 20–30 µm, on average about 110 × 25 µm in life; body length:width ratio highly variable, that is, 3.3–6.6:1, on average 4.7:1 both in life and protargol preparations (Table 38). Body outline usually very elongate elliptical with a small concavity at buccal vertex; posterior portion more distinctly narrowed than anterior, ocassionally with bluntly pointed end or rather broad, highly resembling Urosomoida agiliformis (Fig. 121a, e, f). Body flexible, but acontractile, dorsoventrally flattened up to 2:1 with rather distinct furrow along dorsal kinety 4. Macronuclear nodules in middle body third left of midline, rear nodule usually closer to body margin than front one, invariably connected by a fine, granulated thread, broadly (1.5:1) to very elongate (5:1) ellipsoidal, on average 2.6:1 and widely separated; chromatin bodies highly variable, often some large and many small ones. A globular to ellipsoidal micronucleus attached to left side of each macronuclear nodule, about 2.5 µm in diameter and thus inconspicuous (Fig. 121a, c, e, f). Contractile vacuole slightly to distinctly ahead of mid-body at left cell margin (Fig. 121a). Cytopyge subterminal on dorsal side. No specific cortical granules. Cytoplasm colourless, contains some lipid droplets and 5–15 µm-sized food vacuoles. Swims and glides rather rapidly. Adoral zone occupies 18–30%, on average 22%, of body length; of usual shape and structure, consists of an average of 20 membranelles; bases of largest membranelles about 6 µm wide in life. Buccal cavity narrow and flat; buccal lip angularly projecting and thus prominent, covers posterior portion of adoral zone and bears paroral. Undulating membranes curved and almost side by side, optically intersecting slightly in anterior quarter; paroral commences a few micrometers ahead of endoral, which is longer posteriorly, likely consists of closely spaced dikinetids. Pharyngeal fibres distinct in life and in protargol preparations, extend to at least mid-body right of midline (Fig. 121a, b, Table 38). Cirral pattern very constant, number of cirri of usual variability (Fig. 121a, b, Table 38). Most cirri in life about 10 µm long. Frontal cirri slightly enlarged with right cirrus behind distal end of adoral zone. Buccal cirrus right of anterior end of paroral. Invariably four frontoventral cirri arranged in V-shaped pattern, three of which
b
Fig. 121a–g Erimophrya glatzeli (from Foissner et al. 2002. a, from life; b–g, protargol impregnation). a: Ventral view of a representative specimen with a distinct furrow along dorsal kinety 4, 106 µm. Arrows mark caudal cirri. b, c: Infraciliature of ventral and dorsal side and nuclear apparatus of holotype specimen, 96 µm. Dotted line connects frontal cirri, broken lines connect cirri originating from same anlage (transverse cirrus very likely originates, like the frontoterminal cirri, from anlage VI). Frontoventral cirri circled. Arrow marks cirrus III/2, asterisk denotes cirrus IV/3. d: Middle divider (85 µm) showing that only five frontal-ventral-transverse cirri anlagen are formed (anlage V lacking). Arrow marks dorsomarginal kinety of opisthe. e, f: One of the smallest (e, 69 µm) and largest (f, 115 µm) specimen found in the protargol slides, demonstrating the great variability in body size and shape. Note the threadlike structure connecting the macronuclear nodules. g: Arrangement of dorsal kineties in broad specimen, 78 µm. Note the blank area between kineties 2 and 3 (asterisk). AZM = adoral zone of membranelles, CC = caudal cirri, CV = contractile vacuole, FC = right frontal cirrus, FT = frontoterminal cirri, MA = anterior macronuclear nodule, MI = micronucleus, RMR = rear end of right marginal row, TC = transverse cirrus, I–IV, VI = frontal-ventral-transverse cirri anlagen, 1–4 = dorsal kineties. Page 579.
582
SYSTEMATIC SECTION
(IV/3, VI/3, VI/4) form a conspicuous bow with frontal cirri. Usually two, rarely one or three postoral ventral cirri distinctly behind buccal vertex; in the original description we incorrectly supposed that the anteriormost cirrus (usually cirrus IV/2; see Berger 1999, Fig. 6a) has been reduced (for detailed discussion, see cell division below). Usually one, rarely two transverse cirri between last cirrus of marginal rows, that is, in body midline near rear cell end, feigning confluent marginal rows. Right marginal row commences about at level of posterior frontoterminal cirrus, ends subterminally, while left row ends at level of transverse cirrus, that is, almost at rear cell end; left row commences, as is usual, left of proximal end of adoral zone; distances between individual cirri increase slightly from anterior to posterior in both marginal rows, while size of cirri hardly changes. Dorsal bristles 3–4 µm long in life, arranged in four kineties forming very constant pattern (Fig. 121c, g, Table 38). Kineties 1 and 2 slightly shortened anteriorly, each composed of about 10 bristles and associated with a caudal cirrus. Kinety 3 slightly shortened anteriorly, extends to rear body end, convex, leaving blank a fusiform area in body midline because kinety 2 is concave; lacks a caudal cirrus. Kinety 4 originates dorsomarginally and is composed of only 3–6 bristles extending along a rather distinct furrow in anterior quarter of cell. Caudal cirri about 15 µm long and obliquely spread (Fig. 121a). Cell division (Fig. 121d): Foissner et al. (2002) found some dividers showing that ontogenesis commences with the formation of an anarchic field of basal bodies each left of the posterior postoral ventral cirrus and the single transverse cirrus, quite similarly to Urosomoida agiliformis (for review, see Berger 1999). The frontal-ventral-transverse cirri anlagen of proter and opisthe originate independently and produce supernumerary cirri that are reduced during cirral patterning. Four middle dividers invariably showed that only five cirral anlagen develop in E. glatzeli (Fig. 121d). In 18-cirri hypotrichs the three postoral ventral cirri are formed from anlagen IV (cirrus IV/2; usually anteriormost cirrus) and V (V/3 and V/4) (for review, see Berger 1999, Fig. 6a). Since the ordinary number of four frontoventral cirri is present in E. glatzeli we can be sure that anlage IV is present; consequently we have to assume that the two postoral ventral cirri present (Fig. 121b) are formed from anlage IV, that is, anlage IV forms, unlike in the 18-cirri hypotrichs, two postoral ventral cirri. This is the sole possibility to explain the Erimophrya-cirral pattern when only five cirral anlagen are present. The increased number of postoral ventral cirri formed from anlage IV is reminiscent of Apourosomoida halophila (Fig. 110i–m). In this species, anlage IV produces the rearmost frontoventral cirrus (IV/3) plus 3–6, usually five, postoral ventral cirri. However, Apourosomoida has a rather different dorsal kinety pattern, that is, lacks a dorsomarginal row and has a curious kinety fragmentation, which is, however, very likely not homologous to the fragmentation of the oxytrichids. Occurrence and ecology: Very likely confined to terrestrial habitats (Foissner et al. 2002, p. 56). Type locality of Erimophrya glatzeli is the Sossus Vlei of the Na-
Erimophrya
583
Table 38 Morphometric data on Erimophrya arenicola (are, from Foissner et al. 2002), Erimophrya glatzeli (gla, from Foissner et al. 2002), Erimophrya quadrinucleata (qua, from Foissner et al. 2005), and Erimophrya sylvatica (syl, from Foissner et al. 2005) Characteristics a Body, length
Body, width
Body length:width, ratio
Adoral zone of membranelles, length
Body length:length of adoral zone, ratio
Anterior body end to paroral, distance
Paroral, length
Anterior body end to endoral, distance
Endoral, length
Anterior body end to anteriormost frontoventral cirrus, distance Anterior body end to rearmost frontoventral cirrus, distance
Anterior body end to buccal cirrus, distance
Anterior body end to right marginal row, distance
species are gla qua syl are gla qua syl are gla qua syl are gla qua syl are gla qua syl gla qua syl gla qua syl gla qua syl gla qua syl gla qua syl are gla qua syl are gla qua syl are gla qua syl
mean
M
85.0 85.0 103.2 102.0 92.7 96.0 83.0 83.0 12.4 12.0 21.8 21.0 13.0 13.0 11.7 12.0 6.9 6.7 4.8 4.7 7.4 7.1 7.1 7.4 18.1 18.0 22.4 22.0 20.5 21.0 17.1 17.0 4.7 4.8 4.6 4.6 4.6 4.4 4.9 4.8 6.1 6.0 7.5 8.0 5.6 5.0 8.3 9.0 5.5 5.0 5.6 6.0 8.2 8.0 9.6 10.0 8.4 8.0 9.5 9.0 6.6 7.0 6.2 6.0 6.0 6.0 5.2 5.0 4.8 5.0 10.3 10.0 13.9 14.0 10.7 11.0 9.8 10.0 6.5 6.0 6.1 6.0 8.0 8.0 6.0 6.0 12.3 12.5 11.0 11.0 9.7 10.0 9.7 10.0
SD
SE
CV
Min
Max
n
6.6 16.6 11.1 8.4 1.5 2.1 2.4 1.0 0.8 0.9 1.7 0.9 1.1 0.9 1.9 1.2 0.4 0.7 0.6 0.5 0.8 1.2 1.0 1.3 0.9 – 0.7 1.2 0.9 0.9 0.2 0.9 1.1 0.5 0.6 0.8 1.5 0.8 0.9 0.8 0.8 1.2 0.6 1.6 1.9 1.6 1.9
1.4 3.6 2.6 1.8 0.3 0.5 0.5 0.2 0.2 0.2 0.4 0.2 0.2 0.2 0.4 0.3 0.1 0.2 0.1 0.1 0.2 0.3 0.2 0.3 0.2 – 0.2 0.3 0.2 0.2 0.2 0.2 0.3 0.1 0.1 0.2 0.3 0.2 0.2 0.2 0.2 0.3 0.1 0.3 0.4 0.4 0.4
7.8 16.0 12.0 10.1 11.9 9.9 18.3 8.6 11.7 18.6 22.4 12.0 6.2 3.9 9.3 7.0 8.9 15.0 13.7 9.4 12.9 16.2 17.3 15.3 16.4 – 8.5 12.6 11.2 9.2 13.7 15.4 19.0 9.7 12.5 7.5 10.7 7.0 9.1 12.2 12.6 14.4 10.5 13.1 17.5 16.7 19.0
74.0 76.0 68.0 72.0 10.0 20.0 9.0 10.0 6.0 3.3 4.7 5.2 16.0 21.0 16.0 15.0 3.9 3.3 3.4 4.0 5.0 5.0 4.0 7.0 4.0 5.0 7.0 7.0 7.0 8.0 5.0 5.0 4.0 5.0 4.0 9.0 12.0 10.0 8.0 5.0 5.0 6.0 5.0 10.0 8.0 7.0 5.0
100.0 133.0 110.0 101.0 15.0 29.0 17.0 14.0 9.0 6.6 10.3 8.7 21.0 24.0 23.0 20.0 5.3 5.7 5.6 5.6 8.0 9.0 8.0 12.0 7.0 6.0 10.0 11.9 11.0 11.0 8.0 8.0 8.0 7.0 6.0 12.0 17.0 12.0 11.0 8.0 8.0 10.0 7.0 15.0 14.0 13.0 12.0
23 21 19 21 23 21 19 21 23 21 19 21 23 21 19 21 23 21 19 21 21 19 21 21 19 21 21 19 20 21 19 19 21 19 21 22 21 19 21 23 21 19 21 22 21 19 21
584
SYSTEMATIC SECTION
Table 38 Continued Characteristics a Anterior body end to anteriormost postoral ventral cirrus, distance
Anterior body end to last postoral ventral cirrus, distance Anterior body end to dorsal kinety 1, distance Anterior body end to dorsal kinety 2, distance Anterior body end to dorsal kinety 3, distance Anterior body end to dorsal kinety 4, distance Anterior body end to end of dorsal kinety 4, distance Anterior body end to anteriormost macronuclear nodule, distance
Nuclear figure, length
Macronuclear nodules, distance in between
Anterior macronuclear nodule, length
Anterior macronuclear nodule, width
Macronuclear nodules, number
Anterior micronucleus, length
Anterior micronucleus, width
species
mean
M
SD
SE
CV
Min
Max
n
are gla qua syl gla qua gla qua syl qua syl qua syl gla
21.5 26.9 21.5 19.3 32.8 27.6 16.7 23.0 18.1 18.7 15.0 5.1 5.3 5.8
22.0 27.0 22.0 19.0 33.0 28.0 17.0 22.0 17.0 19.0 15.0 5.0 5.0 6.0
1.3 2.4 1.8 1.5 3.0 1.8 2.8 3.9 2.2 3.1 2.3 0.8 0.7 1.0
0.3 0.5 0.4 0.3 0.7 0.4 0.6 0.9 0.5 0.7 0.5 0.2 0.1 0.2
5.9 8.8 8.6 7.7 9.2 6.5 16.6 17.0 13.4 16.5 15.5 15.4 12.3 16.9
19.0 23.0 18.0 16.0 27.0 24.0 10.0 16.0 15.0 13.0 11.0 4.0 4.0 4.0
24.0 32.0 24.0 23.0 39.0 30.0 22.0 31.0 23.0 24.0 21.0 7.0 6.0 8.0
23 21 19 21 21 19 21 19 21 19 21 19 21 21
gla
22.1
22.0
3.1
0.7
14.2
16.0
27.0
21
are gla qua syl are gla qua syl are gla qua g syl are gla qua syl are gla qua syl are b gla b qua syl are gla qua syl are gla qua syl
20.7 23.6 20.4 18.8 33.3 47.6 46.6 35.7 3.3 14.2 0.8 3.3 15.0 16.6 12.3 16.9 4.2 6.3 4.0 3.5 2.0 2.0 4.1 2.0 2.8 2.2 2.5 2.7 1.7 2.0 1.8 1.7
21.0 23.0 20.0 18.0 32.0 49.0 46.0 34.0 3.0 15.0 0.0 3.0 14.0 17.0 12.0 17.0 4.0 6.0 4.0 3.0 2.0 2.0 4.0 2.0 2.5 2.2 2.5 2.5 1.5 2.0 1.9 1.5
2.1 3.0 2.4 2.4 5.7 8.8 6.6 5.9 1.9 4.3 – 1.9 3.2 3.5 2.7 3.1 1.9 1.1 0.6 0.8 0.0 0.0 0.5 0.0 0.6 – 0.4 0.9 – – 0.3 0.3
0.4 0.7 0.5 0.5 1.2 1.9 1.5 1.3 0.4 0.9 – 0.4 0.7 0.8 0.6 0.7 0.4 0.2 0.1 0.2 0.0 0.0 0.1 0.0 0.1 – 0.1 0.2 – – 0.1 0.1
10.1 12.8 11.6 12.8 17.0 18.6 14.3 16.4 58.4 30.0 – 57.1 21.2 21.3 22.1 18.4 44.3 17.5 14.4 21.3 0.0 0.0 11.2 0.0 20.6 – 16.7 31.8 – – 13.6 14.7
18.0 17.0 15.0 16.0 22.0 32.0 37.0 28.0 0.0 7.0 0.0 1.0 6.0 12.0 9.0 10.0 3.0 5.9 3.0 3.0 2.0 2.0 3.0 2.0 1.5 1.9 2.0 2.0 1.5 1.5 1.5 1.5
25.0 29.0 24.0 26.0 44.0 64.0 61.0 47.0 8.0 22.0 2.0 9.0 23.0 24.0 20.0 22.0 12.0 9.0 5.0 5.0 2.0 2.0 5.0 2.0 4.0 2.5 3.2 6.0 2.5 2.2 2.5 2.0
23 21 19 21 23 21 19 21 28 21 19 21 23 21 19 21 23 21 19 21 21 23 19 19 22 21 19 21 22 21 19 21
Erimophrya
585
Table 38 Continued Characteristics a
species
Micronuclei, number
Posterior body end to rearmost transverse cirrus, distance Posterior body end to right marginal row, distance Posterior body end to left marginal row, distance Adoral membranelles, number
Frontal cirri, number
Frontoventral cirri, number
Buccal cirri, number
Postoral ventral cirri, number
Transverse cirri, number
Right marginal cirri, number
Left marginal cirri, number
Dorsal kineties, number
Dorsal kinety 1, number of bristles
mean
M
SD
SE
CV
Min
Max
n
are gla qua syl gla qua syl gla qua syl gla
1.7 2.0 2.0 2.0 2.0 [1.0 [1.0 5.5 6.0 8.8 2.2
2.0 2.0 2.0 2.0 2.0
0.6 0.0 – – 0.5
0.1 0.0 – – 0.1
36.5 0.0 – – 24.3
1.0 2.0 1.0 1.0 1.0
3.0 2.0 2.0 2.0 3.0
22 21 21 21 21
5.0 5.0 9.0 2.0
2.6 2.8 2.8 1.3
0.6 0.6 0.6 0.3
46.8 46.1 31.4 60.6
2.0 3.0 4.0 1.0
14.0 12.0 13.0 5.0
21 19 21 21
are gla qua syl are gla qua syl are gla qua syl are gla qua syl are gla c qua syl are d gla qua h syl are gla qua syl are gla qua syl are gla qua syl are
15.0 20.5 15.8 14.2 3.0 3.0 3.0 3.0 4.0 4.0 4.0 4.0 1.0 1.0 1.0 1.0 1.0 2.0 2.8 1.7 1.0 1.1 2.0 2.0 18.6 21.2 21.3 19.0 19.3 19.8 17.7 16.6 3.0 4.0 3.0 3.0 3.0
15.0 20.0 16.0 14.0 3.0 3.0 3.0 3.0 4.0 4.0 4.0 4.0 1.0 1.0 1.0 1.0 1.0 2.0 3.0 2.0 1.0 1.0 2.0 2.0 19.0 22.0 21.0 19.0 19.0 20.0 18.0 17.0 3.0 4.0 3.0 3.0 3.0
0.9 0.8 1.4 0.9 0.0 0.0 0.0 0.0 0.4 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 – – 0.0 – 0.0 0.0 2.4 1.7 2.1 1.6 3.4 1.7 1.4 1.2 0.0 0.0 0.0 0.0 1.5
0.2 0.2 0.3 0.2 0.0 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 – – 0.0 – 0.0 0.0 0.5 0.4 0.5 0.4 0.7 0.4 0.3 0.3 0.0 0.0 0.0 0.0 0.4
5.8 3.7 8.6 6.5 0.0 0.0 0.0 0.0 9.5 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 – – 0.0 – 0.0 0.0 12.9 8.1 10.0 8.6 17.9 8.5 7.7 7.0 0.0 0.0 0.0 0.0 49.4
13.0 19.0 12.0 12.0 3.0 3.0 3.0 3.0 3.0 4.0 4.0 4.0 1.0 1.0 1.0 1.0 1.0 2.0 1.0 1.0 1.0 1.0 2.0 2.0 14.0 17.0 16.0 16.0 14.0 18.0 16.0 13.0 3.0 4.0 3.0 3.0 2.0
17.0 22.0 17.0 16.0 3.0 3.0 3.0 3.0 5.0 4.0 4.0 4.0 1.0 1.0 1.0 1.0 1.0 2.0 3.0 2.0 1.0 2.0 2.0 2.0 24.0 24.0 25.0 23.0 27.0 24.0 20.0 19.0 3.0 4.0 3.0 3.0 7.0
22 21 19 21 22 21 19 21 22 21 19 21 23 21 19 21 23 21 19 21 5 21 19 21 22 21 19 21 23 21 19 21 22 21 19 21 11
586
SYSTEMATIC SECTION
Table 38 Continued Characteristics a
Dorsal kinety 2, number of bristles
Dorsal kinety 3, number of bristles Rightmost dorsal kinety f, number of bristles
Caudal cirri, number
species
mean
M
SD
SE
CV
Min
Max
n
gla qua syl are gla qua syl gla are gla qua syl are gla e qua syl
9.3 2.5 3.0 9.7 12.1 9.4 9.5 9.3 3.3 4.3 3.6 3.0 2.0 2.0 2.0 2.0
9.0 2.0 3.0 10.0 12.0 9.0 9.0 9.0 3.0 4.0 4.0 3.0 2.0 2.0 2.0 2.0
0.7 1.0 – 1.4 1.1 1.0 0.9 1.2 0.6 1.0 0.8 – 0.0 0.0 0.0 0.0
0.2 0.2 – 0.4 0.2 0.2 0.2 0.3 0.2 0.2 0.2 – 0.0 0.0 0.0 0.0
7.8 39.0 – 14.6 8.8 10.2 9.2 12.4 19.1 23.5 21.5 – 0.0 0.0 0.0 0.0
8.0 1.0 2.0 7.0 10.0 8.0 8.0 7.0 2.0 3.0 2.0 3.0 2.0 2.0 2.0 2.0
11.0 5.0 4.0 12.0 15.0 12.0 11.0 12.0 4.0 6.0 5.0 4.0 2.0 2.0 2.0 2.0
21 19 21 11 21 19 21 21 12 21 19 21 22 21 19 21
a
All measurements in µm. CV = coefficient of variation in %, M = median, Max = maximum value, mean = arithmetic mean, Min = minimum value, n = number of individuals investigated, SD = standard deviation, SE = standard error of arithmetic mean. Data based on protargol-impregnated specimens. b
Specimens with three or four nodules occur very rarely.
c
Specimens with one or three postoral ventral cirri occur very rarely.
d
From late dividers.
e
Specimens with only one caudal cirrus are very rare.
f
This is the dorsomarginal kinety.
g
Between central nodules.
h
Rarely only one cirrus present.
mib Desert (24°50'S, 15°20'E), where we discovered it in humous sand under Acacia erioloba (Foissner et al. 2002, site 24); it occurred in great numbers, indicating that many cysts were present. Feeds on bacteria and heterotrophic flagellates (Foissner et al. 2002).
Erimophrya arenicola Foissner, Agatha & Berger, 2002 (Fig. 122a–h, Table 38) 2002 Erimophrya arenicola nov. spec.1 – Foissner, Agatha & Berger, Denisia, 5: 796, Fig. 173a–h, Table 153 (Fig. 122a–h; original description; the holotype slide [accession number 2002/243] and two 1
Foissner et al. (2002) provided the following diagnosis: Size about 90 × 13 µm in vivo; pisciform and slightly twisted about main body axis. On average 2 almost abutting, very elongate ellipsoidal macronuclear nodules, 15 adoral membranelles, 19 cirri each in right and left marginal row, 1 postoral cirrus, 1
Erimophrya
587
paratype slides [2002/244, 245] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; see Foissner et al. 2002, p. 39 and Aescht 2003, p. 380).
Nomenclature: The species-group name arenicola (Latin adjective; living in sand) refers to the preferred habitat (Foissner et al. 2002). Remarks: For separation from congeners, see key and Table 38. In life, Erimophrya arenicola is easily confused with Urosomoida agilis (for review, see Berger 1999, p. 347) which has three postoral ventral cirri (vs. one), four dorsal kineties (vs. three), and colourless to yellowish cortical granules (vs. lacking). Hemiurosoma similis (Foissner, 1982) lacks postoral ventral cirri (p. 633; for review, see Berger 1999, p. 419, as Urosoma similis). In life, Erimophrya arenicola is identified by the following combination of features: body very slender (7:1), two almost abutting macronuclear nodules, and one postoral ventral cirrus. Nevertheless, identifications of such slender specimens should be checked by protargol impregnation (Foissner et al. 2002). Morphology: Body size 70–110 × 10–20 µm, on average about 90 × 13 µm in life; body length:width ratio about 7:1 on average both in life and in protargol preparations (Table 38). Body outline rather constant, usually slenderly fusiform or pisciform, that is, anterior end narrowly rounded and posterior end elongated tail-like (Fig. 122a, b, d–f); occasionally elongate elliptical with both ends rounded or posterior end bluntly pointed (Fig. 122c). Body flexible, but acontractile; usually slightly twisted about main body axis and inconspicuously flattened dorsoventrally. Macronuclear nodules in middle third left of midline, ellipsoidal (length:width ratio 2:1) to very elongate ellipsoidal (4.5:1), on average 4:1, and close together; contain many chromatin bodies. On average two ellipsoidal micronuclei, one attached to each macronuclear nodule in variable positions. Contractile vacuole slightly to distinctly ahead of mid-body at left cell margin. No specific cortical granules. Cytoplasm colourless, with few to many lipid droplets 1–3 µm across and some ordinary cytoplasmic crystals mainly in posterior body portion, which may thus appear dark at low magnification. Moves rather slowly and clumsily, possibly due to the tortuous body. Adoral zone occupies 18–25%, on average 21% of body length, of usual shape and structure, composed of an average of 15 membranelles (Fig. 122a, e, Table 38); bases of largest membranelles 6 µm in life; frontal three membranelles usually separated from ventral membranelles by a more or less distinct gap at left anterior body corner. Buccal cavity narrow and flat; buccal lip angularly projecting and thus prominent, covers posterior third of adoral zone of membranelles and bears paroral. Undulating membranes slightly curved and almost in parallel, endoral about 7–10 µm long, paroral 5–6 µm long, begins about 2 µm ahead of endoral at level of buccal cirrus; exact structure of membranes not clearly recognisable. Pharyngeal fibres distinct in life and in protargol preparations, of ordinary length and structure, extend obliquely backwards (Fig. 122a, e, Table 38).
transverse cirrus, 2 caudal cirri, and 3 dorsal kineties.
588
SYSTEMATIC SECTION
Fig. 122a–h Erimophrya arenicola (from Foissner et al. 2002. a–d, from life; e–h, protargol impregnation). a: Ventral view of a representative specimen, 102 µm. b–d: Ventral views of shape variants. e, f: Infraciliature of ventral and dorsal side and nuclear apparatus of holotype specimen, 79 µm. Long arrow marks inconspicuous gap in adoral zone, anterior short arrow denotes frontoventral cirrus III/2, rear arrows denote very likely the transverse cirrus. Frontal cirri connected by dotted line, broken lines connect cirri originating from same anlage (note that the two cirri [rearmost frontoventral cirrus marked by asterisk and postoral ventral cirrus] formed by anlage IV are not connected; transverse cirrus originates from anlage VI [see Fig. 122h]). g, h: Infraciliature of ventral side of a middle (91 µm) and late (97 µm) divider (new cirri black, parental white). Arrow in (g) marks new right marginal row of opisthe. Long arrow in (h) denotes dorso-
Erimophrya
589
Cirral pattern rather constant, number of cirri of usual variability (Fig. 122a, e, f, Table 38). Most cirri about 8 µm long in life and rather fine, transverse cirri about 12 µm long. Frontal cirri about of same size as other cirri, form slightly oblique pseudorow. Buccal cirrus right of anterior end of paroral. Usually four frontoventral cirri, anteriormost (= cirrus VI/4) about at level of right frontal cirrus. One postoral ventral cirrus behind buccal vertex. Number of transverse cirri not recognisable in morphostatic specimens; cell division shows that one cirrus is formed (Fig. 122g, h); thus, one of the three cirri usually present at rear end is a transverse cirrus (Fig. 122e, f). Right marginal row commences near level or rearmost frontoventral cirrus, extends, like left row, to near rear cell end. Left row begins left of buccal vertex, extends onto dorsolateral surface posteriorly due to body torsion; distance between individual cirri increase distinctly from anterior to posterior in both rows. Dorsal bristles about 2 µm long in life, arranged in three kineties (Fig. 122f, Table 38): kinety 1 composed of 2–7 bristles only, one bristle invariably near corresponding caudal cirrus; kinety 2 slightly shortened anteriorly and also terminating with a caudal cirrus; kinety 3 is a dorsomarginal row and composed of only 2–4 bristles extending in anterior quarter of cell. Caudal cirri about 12 µm long. Cell division (Fig. 122g, h): We found some dividers showing that this part of the life cycle proceeds basically as in Urosomoida (Foissner et a. 2002; for review, see Berger 1999, p. 345). Morphogenesis commences with the formation of an oral primordium left of the single postoral cirrus. Subsequently, five cirral anlagen each develop independently in both filial products, that is, no primary primordia – as in Apourosomoida (Fig. 110d) – are formed. Anlage IV produces the single postoral ventral cirrus (IV/2), while anlage VI forms the single transverse cirrus (Fig. 122g, h). From the cirri present in non-dividers and the anlagen formation one can conclude that anlage V is lacking. This anlage forms the two right/posterior postoral ventral cirri, the left pretransverse ventral cirrus, and a transverse cirrus in 18-cirri hypotrichs. As is usual, two anlagen each are generated within parental dorsal kineties 1 and 2. Each anlage produces a caudal cirrus. Dorsal kinety 3 is a dorsomarginal row, that is, originates from/near the anterior end of the right marginal row primordium (Fig. 122h). Occurrence and ecology: Very likely confined to terrestrial habitats (Foissner et al. 2002, p. 56). Type locality of E. arenicola is the Sossus Vlei of the Namib Desert (24°50'S, 15°20'E), where we discovered it in sand with litter from Nara scrubs (Foissner et al. 2002, site 23); also recorded from a second site (site 20) in the same area. Interestingly, the type species (E. glatzeli) also occurred in the Sossus Vlei. Feeds on bacteria and the heterotrophic flagellate Polytomella (Foissner et al. 2002).
b
marginal kinety of opisthe, short arrow marks new postoral ventral cirrus of proter. Cirri originating from same anlage are connected by broken lines (only shown for opisthe). AZM = distal end of adoral zone of membranelles, CC = caudal cirri, FT = frontoterminal cirri (part of the frontoventral cirri), LMR = left marginal row, MA = posterior macronuclear nodule, P = paroral, RMR = right marginal row, TC = transverse cirrus of proter, I–IV, VI = frontal-ventral-transverse cirri anlagen, 1–3 = dorsal kineties (kinety 3 is a dorsomarginal row). Page 586.
590
SYSTEMATIC SECTION
Erimophrya sylvatica Foissner, Berger, Xu & ZechmeisterBoltenstern, 2005 (Fig. 123a–f, 124a, b, Table 38) 2005 Erimophrya sylvatica nov. spec.1 – Foissner, Berger, Xu & Zechmeister-Boltenstern, Biodiversity and Conservation, 14: 667, Fig. 11a–f, 13a, d, Table 9 (Fig. 123a–f, 124a, b; original description; one holotype slide and two paratype slides are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria).
Nomenclature: The species-group name sylvatic·us, -a, -um (Latin adjective [m; f; n]; belonging to forests) refers to the habitat where the species was discovered (Foissner et al. 2005). Remarks: The present species is most similar to E. arenicola, which, however, is possibly confined to Africa. They differ from each other in the number of postoral ventral cirri (2 vs. 1) and transverse cirri (2 vs. 1). Minor differences are the more slender macronuclear nodules (4.8:1 vs. 3.6:1) and the cortical crystal reticulum (present vs. absent). The type species differs from E. sylvatica in the body length:width ratio (4.8:1 vs. 7:1 in present species), the macronuclear nodules (length:width ratio 2.6:1 vs. 4.8:1; distance between nodules 14.2 µm vs. 3.3 µm), and the number of adoral membranelles (20 vs. 14), dorsal kineties (4 vs. 3), and bristles in dorsal kinety 1 (9 vs. 3). Morphology: Body size 80–115 × 10–20 µm, on average about 95 × 15 µm in life; body length:width ratio about 7:1 on average both in life and in protargol preparations (Table 38). Body outline moderately variable, usually slenderly pisciform or lanceolate with narrowly rounded anterior end and curved, bluntly pointed posterior end; occasionally almost parallel-sided. Body acontractile, invariably twisted about main body axis and up to 2:1 fattened dorsoventrally (Fig. 123a, b, 124a). Macronuclear nodules in middle third of cell left of midline, usually close together and connected by fine thread, ellipsoidal (2:1) to cylindroidal (7:1), on average very elongate ellipsoidal (4.8:1; Table 38); chromatin bodies rather large, globular to ellipsoidal. Two ellipsoidal, inconspicuous micronuclei, one attached to each macronuclear nodule in variable positions. Contractile vacuole ahead of mid-body at left cell margin, surrounded by small vacuoles during diastole. Cortex very flexible, yellowish due to a reticular pattern of 0.5–2.0 µm-sized crystals conspicuously sparkling under interference contrast illumination (Fig. 123e, 124a, b). Specific cortical granules lacking. Cytoplasm colourless, contains some lipid droplets up to 3 µm across and crystals like those found in the cortex. Food vacuoles about 5 µm across. Glides rapidly on microscope slide and soil particles, showing great flexibility. Adoral zone occupies 18–25%, on average 20% of body length, extends along left margin of cell, composed of an average of 14 membranelles, of which the fron1
Foissner et al. (2005) provided the following diagnosis: Size about 95 µm × 15 µm in vivo; slenderly pisciform and slightly twisted about main body axis. On average 2 almost abutting, very elongate ellipsoidal (4.8:1) macronuclear nodules, 14 adoral membranelles, 19 cirri in right and 17 in left marginal row, 2 postoral cirri, 2 transverse cirri, 2 caudal cirri, and 3 dorsal kineties with kinety 1 composed of 3 bristles.
Erimophrya
591
Fig. 123a–f Erimophrya sylvatica (from Foissner et al. 2005. a, e, from life; b–d, f, protargol impregnation). a: Ventral view of a representative specimen, 94 µm. b, c, f: Infraciliature of ventral and dorsal side and nuclear apparatus of holotype specimen, 88 µm. Arrow in (b) marks the two postoral ventral cirri formed from anlage IV. Arrow in (f) marks rearmost bristle of dorsal kinety 1 close to the corresponding caudal cirrus. d: Infraciliature of oral region at high magnification, 26 µm. Long arrow marks gap in adoral zone, short arrow denotes frontoventral cirrus III/2. Asterisk marks rearmost frontoventral cirrus (IV/3). Broken lines connect cirri originating from same anlage. e: Surface view showing cortical crystal pattern in mid-body (cp. Fig. 124a, b). AZM = distal end of slightly bipartite adoral zone of membranelles, CC = caudal cirri on dorsal kineties 1 and 2, E = endoral, FC = right frontal cirrus, FT = frontoterminal cirri, LMR = anterior end of left marginal row, MA = macronuclear nodule, MI = micronucleus, RMR = anterior end of right marginal row, TC = transverse cirri, IV = frontal-ventral-transverse cirri anlage IV, 1–3 = dorsal kineties (kinety 3 is a dorsomarginal row and therefor has no caudal cirrus at the posterior end). Page 590.
592
SYSTEMATIC SECTION
Fig. 124a–d Erimophrya sylvatica (a, b) and Erimophrya quadrinucleata (c, d) (from Foissner et al. 2005. From life). a, b: Dorsal view of E. sylvatica showing the reticular crystal pattern with sparkling individual crystals (dark or bright depending on light refraction). c, d: Dorsal views of squashed specimens of E. quadrinucleata showing crystals (arrows) associated with conspicuous plaques of minute granules (arrowheads). CV = contractile vacuole. Pages 590, 594.
Erimophrya
593
tal three are separated by a minute gap, as in Erimophrya arenicola and E. quadrinucleata. Buccal cavity narrow and flat; buccal lip angularly covering posterior third of adoral zone. Undulating membranes slightly curved and side by side, endoral about 6 µm long, paroral 8 µm long on average and commencing about 3 µm ahead of endoral at level of buccal cirrus; exact structure (dikinetidal?) of membranes not recognisable. Pharyngeal fibres inconspicuous in life and protargol preparations, of ordinary length and structure (Fig. 123b, d, Table 38). Cirral pattern very constant, number of cirri of usual variability (Fig. 123b, d, Table 38). Cirri about 8 µm long in life, most composed of only six cilia, in posterior third of even only four. Frontal cirri of same size as other cirri, arranged in slightly oblique pseudorow forming bow with three frontoventral cirri. Buccal cirrus right of anterior end of paroral, composed of only four cilia. Four frontoventral cirri arranged in V-like pattern, anteriormost cirrus (VI/4) in, or almost in line with frontal cirri. Cirrus IV/3 slightly set off from frontoterminal cirri; frontoventral cirrus III/2 left of this gap, displaced slightly leftwards. One or two, usually two postoral ventral cirri distinctly left of midline and one behind the other, indicating that both cirri (and not only one as in the 18-cirri hypotrichs) are formed from anlage IV. Pretransverse ventral cirri (very likely) lacking. Two transverse cirri side by side at rear end of body, thus difficult to separate from caudal cirri which are in a very similar position on dorsolateral side. Right marginal row commences far off anterior body end about at level of rearmost frontoventral cirrus (in specimen shown in Fig. 123b at 12% of body length). Left row commences left of proximal end of adoral zone, extends onto dorsolateral surface posteriorly. Marginal rows extend slightly obliquely backwards due to body torsion, both distinctly shortened posteriorly. Dorsal bristles 2–3 µm long in life, arranged in three kineties (Fig. 123c, Table 38). Kinety 1 composed of only three bristles on average, namely two in anterior half and, separated by a wide distance, one very near to the corresponding caudal cirrus. Kinety 2 slightly shortened anteriorly and somewhat sigmoidal, last bristle very near to corresponding caudal cirrus. Kinety 3 near right body margin of oral portion, composed of only three bristles on average; very likely it is, as in congeners, a dorsomarginal row. Two caudal cirri one after the other at right margin of dorsal side, each accompanied by a dorsal bristle, producing highly characteristic pattern similar to that found in E. arenicola (Fig. 123c, f). Occurrence and ecology: Very likely confined to terrestrial habitats. Type locality of Erimophrya sylvatica is the same as for E. quadrinucleata, that is, the Stampfltal near Vienna (Austria), where we discovered it in a Pinus nigra forest soil (Foissner et al. 2005). The sympatric occurrence indicates that they are true species and not subspecies. Feeds on bacteria (Foissner et al. 2005).
594
SYSTEMATIC SECTION
Fig. 125a–g Erimophrya quadrinucleata (from Foissner et al. 2005. a, b, d, e, from life; c, f, g, protargol impregnation). a: Ventral view of representative specimen slightly twisted about main body axis, 101 µm. b: Outline of a broad specimen with rather abruptly narrowed posterior body portion. Note the four macronuclear nodules, the main feature of the species. c: Infraciliature of oral region at high magnification, 47 µm. Long arrow marks gap in adoral zone, short arrow denotes cirrus III/2, asterisk denotes cirrus IV/3. The latter two cirri form, together with the frontoterminal cirri, the frontoventral cirri. Frontal cirri connected by dotted line. Cirri originating from same anlage connected by broken line. d, e: Surface views showing cortical crystals and plaque pattern (arrows) in two specimens. f, g: Infraciliature of ventral and dorsal side and nuclear apparatus of holotype specimen, 100 µm. Arrow marks postoral ventral cirri. However, note that the postoral ventral cirri in E. quadrinucleata are formed only from anlage IV, whereas in 18-cirri hypotrichs, which also have three postoral ventral cirri, they are formed from anlagen IV (cirrus IV/2) and V (V/3, V/4), that is, not all postoral ventral cirri of E. quadrinucleata and the 18-cirri hypotrichs are homologous. AZM = distal end of adoral zone of membranelles, CC = caudal cirri, CV= contractile vacuole, FC = right frontal cirrus, FT = frontoterminal cirri, LMR = left marginal row, MA = macronuclear nodules, MI = micronucleus, P = paroral, PF = pharyngeal fibres, RMR = right marginal row, TC = transverse cirri, I, IV = frontal- ventral cirri anlagen, 1–3 = dorsal kineties (kinety 3 is a dorsomarginal row). Page 595.
Erimophrya
595
Erimophrya quadrinucleata Foissner, Berger, Xu & ZechmeisterBoltenstern, 2005 (Fig. 124c, d, 125a–g, Table 38) 2005 Erimophrya quadrinucleata nov. spec.1 – Foissner, Berger, Xu & Zechmeister-Boltenstern, Biodiversity and Conservation, 14: 671, Fig. 12a–g, 13b, e, Table 9 (Fig. 124c, d, 125a–g; original description; one holotype slide and two paratype slides are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria).
Nomenclature: The species-group name quadrinucleat·us, -a, -um (Latin adjective [m; f; n]; having four nuclei) is a composite of the Latin numeral quadri (four) and the Latin adjective nucleatus (with a nucleus), referring to the four macronuclear nodules, the main feature of the species (Foissner et al. 2005). Remarks: We found this species together with E. sylvatica from which it can be easily distinguished by the number of macronuclear nodules (Foissner et al. 2005). Since it is very similar to the other three species we did not provide a full description, but explained only the differences and some details (see next chapter). Erimophrya quadrinucleata is the sole Erimophrya species with three postoral ventral cirri. For explanation of the non-homology of the postoral ventral cirri in the present species and in the 18-cirri hypotrichs see remarks in genus section. Morphology: Erimophrya quadrinucleata is very similar to the other species, except in the number of macronuclear nodules (four vs. two) and postoral ventral cirri (three vs. one or two). For characterisation, see diagnosis in footnote to list of synonyms, Fig. 124c, d, 125a–g, Table 38, and the following details: (i) The shape is occasionally distinctly oblanceolate due to the rather abruptly narrowed posterior body region (Fig. 125b); (ii) The rather distinctly wrinkled, abutting macronuclear nodules appear as a rod in life (Fig. 125a, b); (iii) Like in E. sylvatica, the cortex contains a crystalline reticulum supplemented by some conspicuous plaques composed of 0.3 µm-sized granules, which are possibly precursors of the crystals (Fig. 124c, d, 125d, e); (iv) The postoral ventral cirri are arranged one behind the other, and thus form a short row (Fig. 125c, f; see remarks above and at genus section); (v) There are no dorsal bristles near to the caudal cirri, which are arranged side by side (Fig. 125g); (vi) The species occurred together with E. sylvatica, but was less numerous. Occurrence and ecology: Very likely confined to terrestrial habitats. Type locality of Erimophrya quadrinucleata is the Stampfltal near the city of Vienna (Austria), where we discovered it in a Pinus nigra forest soil (Foissner et al. 2005).
1
Foissner et al. (2005) provided the following diagnosis: Size about 105 µm × 15 µm in vivo; slenderly oblanceolate and slightly twisted about main body axis. On average 4 elongate ellipsoidal macronuclear nodules, 16 adoral membranelles, 21 cirri in right and 18 in left marginal row, 3 postoral cirri, 2 transverse cirri, 2 caudal cirri, and 3 dorsal kineties with kinety 1 composed of 2 bristles.
596
SYSTEMATIC SECTION
Vermioxytricha Foissner, Agatha & Berger, 2002 2002 Vermioxytricha nov. gen.1 – Foissner, Agatha & Berger, Denisia, 5: 749 (original description). Type species (by original designation): Vermioxytricha arenicola Foissner, Agatha & Berger, 2002.
Nomenclature: Vermioxytricha is a composite of the Latin noun vermis (worm) and the Greek genus-group name Oxytricha (pointed hair l cirrus), referring to the slender body and the relationship with Oxytricha Bory de Saint-Vincent in Lamouroux, Bory de Saint-Vincent & Deslongchamps, 1824 (Foissner et al. 2002). Characterisation (A = supposed apomorphy): Dorsomarginalia with bipartite adoral zone of membranelles. Undulating membranes almost straight, short, and staggered. Three frontal cirri. Buccal cirrus present. Frontoventral cirri arranged in V-shaped pattern. Frontal-ventral-transverse cirri anlage V lost (A?). Postoral ventral cirri, pretransverse ventral cirri, and transverse cirri are lacking (A?). One right and one left marginal row. Dorsomarginal kinety present. Dorsal kinety fragmentation lacking. Caudal cirri absent. Terrestrial. Additional characters: The two species assigned are basically identical in all features, except for the cortical granules. Consequently, they also agree in many “non-diagnostic” features, for example, flexible, very slender (body length:width ration about 10:1) body; body length about 170 µm on average; 13–16 macronuclear nodules left of midline; contractile vacuole ahead of mid-body, with distinct collecting canals during diastole; adoral zone composed of 13–16 membranelles on average; dorsal bristles 2–3 µm long, arranged in two kineties; dorsomarginal kinety very short, near anterior end. Remarks: We assigned Vermioxytricha to the Oxytrichidae because the cirral pattern and its formation are basically as in this taxon (Foissner et al. 2002). Furthermore, we offered the urosomoid dorsal kinety formation as indicator for the inclusion in the oxytrichids which have been characterised, inter alia, by the presence of 18 characteristically arranged frontal-ventral-transverse cirri by Berger & Foissner (1997) and Berger (1999). Now I suppose that the 18-cirri pattern originating from six anlagen, a feature still retained in many Oxytrichidae, is an apomorphy of the Hypotricha, and therefore a plesiomorphy for the Oxytrichidae (Berger 2006, p. 33). The presence of a dorsomarginal row (Urosomoida pattern, see Berger 1999, p. 73)2 assigns Vermioxytricha to the Dorsomarginalia (Berger 2006, p. 38, Fig. 14a), while 1
Foissner et al. (2002) provided the following diagnosis: Oxytrichidae Ehrenberg, 1838 with bipartite adoral zone of membranelles, undulating membranes side by side, and frontoventral cirri in V-shaped pattern. Postoral, pretransverse, transverse, and caudal cirri lacking. Frontal and frontoventral cirri originate from 5 anlagen, the rightmost of which is a primary primordium. Dorsal morphogenesis in Urosomoida pattern. 2 Earlier we assumed that the Urosomoida pattern (presence of dorsomarginal row, absence of dorsal kinety fragmentation) evolved via the Oxytricha pattern (dorsomarginal row plus dorsal kinety fragmentation present) due to the loss of the fragmentation (Berger & Foissner 1997, Berger 1999). Now I tend to assume that the Urosomoida pattern evolved earlier, that is, the Oxytricha pattern evolved from the Urosomoida pattern. Of course, one cannot exclude that an “Urosomoida pattern” evolved twice.
Vermioxytricha
597
Table 39 Morphometric data on Vermioxytricha arenicola (from Foissner et al. 2002; ar1, type population from Namibia; ar2, population from Tunisia) and Vermioxytricha muelleri (mue, from Foissner 1986) Characteristics a
species
Body, length
Body, width
Body length:width, ratio Adoral zone of membranelles, length
Body length:length of adoral zone, ratio Anterior body end to paroral, distance Anterior body end to endoral, distance Paroral, length Endoral, length Anterior body end to right marginal row, distance Anterior body end to rearmost frontoventral cirrus, distance Anterior body end to buccal cirrus, distance Anterior body end to anteriormost macronuclear nodule, distance Nuclear figure, length Macronuclear nodule, length
Macronuclear nodule, width
Macronuclear nodules, number
Micronuclei, number
Micronucleus, length
Micronucleus, width
Adoral membranelles, total number
Frontal adoral membranelles, number
ar1 ar2 mue ar1 ar2 mue ar1 ar2 ar1 ar2 mue ar1 ar2 ar1 ar1 ar1 ar1 ar1
mean
M
162.7 162.0 163.6 160.0 118.9 120.0 15.7 15.0 15.5 15.0 10.9 11.0 10.5 10.3 10.7 10.7 21.2 21.0 21.8 22.0 15.5 15.0 7.7 7.8 7.5 7.5 8.9 9.0 10.8 10.5 5.3 5.0 5.9 6.0 11.5 11.0
SD
SE
CV
Min
Max
n
15.3 24.9 14.1 2.7 2.2 0.5 1.3 2.3 1.9 1.3 1.8 1.0 1.0 1.6 1.2 0.9 – 1.2
4.0 6.9 4.3 0.7 0.6 0.2 0.3 0.6 0.5 0.4 0.5 0.2 0.3 0.4 0.4 0.3 – 0.3
9.4 138.0 192.0 15.2 132.0 215.0 11.9 100.0 140.0 17.1 12.0 22.0 14.1 12.0 20.0 4.9 10.0 12.0 12.8 7.9 12.8 21.2 8.0 16.5 8.8 18.0 26.0 6.0 20.0 24.0 11.4 14.0 20.0 12.4 6.4 9.6 12.6 6.1 9.1 18.0 7.0 13.0 10.8 9.0 12.0 17.2 4.0 7.0 – 5.0 6.5 11.6 8.0 12.0
15 13 11 15 13 11 15 13 15 13 11 15 13 13 8 11 7 15
ar1
15.6
16.0
1.7
0.4
10.8
11.0
18.0
15
ar1 ar1
10.1 26.9
10.0 27.0
1.2 3.5
0.3 0.9
11.6 13.0
8.0 19.0
12.0 33.0
15 15
ar1 ar1 d ar2 d mue ar1 d ar2 d mue ar1 ar2 mue ar1 ar2 mue ar1 d ar2 d mue ar1 d ar2 d mue ar1 ar2 mue ar1 ar2
97.3 9.7 7.8 6.4 2.9 2.8 2.4 12.8 15.6 13.8 3.6 2.6 2.0 1.8 1.9 2.8 1.6 1.9 1.4 17.3 17.5 15,6 3.0 3.0
93.0 9.0 7.0 5.6 2.5 3.0 2.5 13.0 16.0 15.0 4.0 2.0 2.0 1.6 1.9 2.8 1.6 1.9 1.4 17.0 18.0 16.0 3.0 3.0
13.7 2.4 1.5 1.7 0.6 0.5 0.4 2.4 2.0 2.9 0.9 1.2 0.0 – – 0.5 – – 0.4 1.2 1.1 0.7 0.0 0.0
3.5 0.6 0.4 0.5 0.2 0.1 0.1 0.7 0.6 0.9 0.2 0.3 0.0 – – 0.2 – – 0.1 0.3 0.3 0.2 0.0 0.0
14.1 25.2 19.1 27.3 21.1 18.1 16.0 18.4 13.0 20.9 24.1 45.6 0.0 – – 18.5 – – 25.4 7.1 6.4 4.3 0.0 0.0
78.0 118.0 5.0 14.0 6.0 10.0 4.2 10.0 2.4 4.0 2.0 4.0 1.7 3.0 8.0 16.0 13.0 18.0 9.0 17.0 2.0 5.0 1.0 5.0 2.0 2.0 1.6 2.5 1.7 2.0 1.8 4.0 1.0 1.6 1.7 2.0 1.0 2.4 16.0 20.0 15.0 19.0 14.1 16.0 3.0 3.0 3.0 3.0
15 15 13 11 15 13 11 13 13 11 13 13 11 15 13 11 15 13 11 15 13 11 15 13
598
SYSTEMATIC SECTION
Table 39 Continued Characteristics a Ventral adoral membranelles, number Frontal cirri, number
Buccal cirrus, number
Frontoventral cirri, number c
Cirri formed by anlage III, number Left marginal cirri, number
Right marginal cirri, number
Dorsal kineties, number
Dorsal kinety 1, number of bristles Dorsal kinety 2, number of bristles
species ar1 ar2 ar1 ar2 mue b ar1 ar2 mue ar1 ar2 mue ar1 ar2 ar1 ar2 mue ar1 ar2 mue ar1 ar2 mue ar1 ar1
mean
M
SD
SE
CV
Min
Max
n
14.3 14.5 3.0 3.0 4.0 1.0 1.0 1.0 2.9 3.0 3.0 1.1 1.2 39.2 39.2 38.2 43.3 46.4 47.3 2.0 2.0 2.0 16.5 1.8
14.0 15.0 3.0 3.0 4.0 1.0 1.0 1.0 3.0 3.0 3.0 1.0 1.0 39.0 37.0 38.0 44.0 44.0 48.0 2.0 2.0 2.0 17.0 2.0
1.2 1.1 0.0 0.0 0.0 0.0 0.0 0.0 – 0.0 0.0 – – 4.9 7.2 4.1 5.4 10.1 2.8 0.0 0.0 0.0 1.6 –
0.3 0.3 0.0 0.0 0.0 0.0 0.0 0.0 – 0.0 0.0 – – 1.3 2.0 1.2 1.4 2.8 0.8 0.0 0.0 0.0 0.4 –
8.6 7.7 0.0 0.0 0.0 0.0 0.0 0.0 – 0.0 0.0 – – 12.6 18.4 10.6 12.4 21.9 5.9 0.0 0.0 0.0 9.4 –
13.0 12.0 3.0 3.0 4.0 1.0 1.0 1.0 1.0 3.0 3.0 1.0 0.0 30.0 30.0 32.0 29.0 36.0 40.0 2.0 2.0 2.0 14.0 1.0
17.0 16.0 3.0 3.0 4.0 1.0 1.0 1.0 3.0 3.0 3.0 3.0 2.0 48.0 56.0 44.0 50.0 78.0 50.0 2.0 2.0 2.0 19.0 2.0
15 13 15 13 11 15 13 11 15 13 11 15 13 15 13 11 15 13 11 15 13 11 13 15
a All measurements in µm. CV = coefficient of variation in %, M = median, Max = maximum value, mean = arithmetic mean, Min = minimum value, n = number of individuals investigated, SD = standard deviation, SE = standard error of arithmetic mean. Data based on protargol-impregnated specimens. b
Cirrus III/2 included.
c
Cirrus III/2 not included (counted as fourth frontal cirrus in original description in V. muelleri; counted separately in V. arenicola, as cirri formed by anlage III).
d
Anteriormost macronuclear nodule, respectively, micronucleus.
the lack of a dorsal kinety fragmentation – the main apomorphy of the Oxytrichidae (Berger 1999, 2006) – prevents the classification in the Oxytrichidae, a large subgroup of the Dorsomarginalia. Vermioxytricha is included in the present review because one species assigned (V. muelleri) was previously classified in Hemisincirra (p. 387), which was excluded from the Oxytrichidae by Berger (1999, p. 893), inter alia, because of the lack of dorsal kinety fragmentation. In addition, Hemisincirra obviously lacks a dorsomarginal row so that an inclusion in the Dorsomarginalia is also not justified. In the present book, Hemisincirra is transferred to the amphisiellids (as incertae sedis), because the frontoventral row – although short in many species – is formed from two anlagen. By contrast, Vermioxytricha is (very likely) not an amphisiellid, because it produces a
Vermioxytricha
599
dorsomarginal kinety, a feature lacking in Amphisiella, type of the amphisiellids. I am unable to give the exact phylogenetic position of Vermioxytricha in the Dorsomarginalia (see chapter 2 in the general section and brief introduction on p. 560). As mentioned above, we assigned two species to Vermioxytricha, namely Hemisincirra muelleri Foissner, 1986 and V. arenicola, a species occurring in several samples from Namibia and Tunisia (Foissner et al. 2002). Vermioxytricha muelleri, which was also recorded from Namibian samples by Foissner et al. (2002, p. 63), differs from V. arenicola only in the absence of cortical granules. The sympatric distribution (admittedly we did not find them together in the same sample, but in two localities around a guest farm; Foissner et al. 2002) indicates that they are species and not subspecies. We checked all other Hemisincirra species to see whether or not some of them belong to the present genus (Foissner et al. 2002). However, most of them have, like the type species H. buitkampi, transverse cirri and have thus not been included in Vermioxytricha. The absence of transverse cirri in H. vermicularis is uncertain and H. rariseta very likely lacks a buccal cirrus. Thus, we retained both species in Hemisincirra, which comprises mostly slender soil species that do not fit into any other genus well (Foissner et al. 2002). The dorsal kinety pattern of Vermioxytricha is a strongly reduced Urosomoida pattern (for details, see review by Berger 1999, p. 73) because only one bipolar kinety is present, the dorsomarginal row is very short, and caudal cirri are lacking. Erimophrya also lacks, like Vermioxytricha, anlage V and has a dorsomarginal kinety (p. 577). However, it has postoral ventral cirri, transverse cirri, and caudal cirri (vs. lacking), and a higher number of dorsal kineties (3–4 vs. 2). Hemiurosoma (p. 614) has, inter alia, (i) the frontoventral cirri arranged in the Urosoma-pattern (that is, cirrus III/2 is ahead of the level of the remaining three cirri IV/3, VI/3, and VI/4 [see Fig. 130h; Fig. 16a in Berger 1999] vs. in V-shaped pattern [Fig. 126k]); (ii) postoral ventral and transverse cirri (vs. absent); (iii) caudal cirri (vs. absent); (iv) four dorsal kineties (vs. two); and (v) four primary primordia (vs. one). Circinella Foissner, 1994 lacks – like Vermioxytricha – transverse cirri and caudal cirri, but has a rather long frontoventral row originating from only one anlage (vs. short and formed from two or three [when cirrus III/2 is included] anlagen). Since the frontoventral row of Circinella originates from one anlage it is not included in the amphisiellids, which form the row from two or three anlagen. Species included in Vermioxytricha (alphabetically arranged basionyms are given): (1) Hemisincirra muelleri Foissner, 1986; (2) Vermioxytricha arenicola Foissner, Agatha & Berger, 2002.
Key to Vermioxytricha species If you cannot identify your specimen/population with the key below, see also Hemisincirra (transverse cirri present; p. 387) or Hemiurosoma (transverse and caudal cirri present, four dorsal kineties; see p. 614 or Foissner et al. 2002, p. 834).
600
SYSTEMATIC SECTION
1 Cortical granules lacking (Fig. 129a–f). . . . . . . Vermioxytricha muelleri (p. 607) - Cortical granules present (Fig. 129a–p). . . . . . Vermioxytricha arenicola (p. 600)
Vermioxytricha arenicola Foissner, Agatha & Berger, 2002 (Fig. 126a–z, 127a, b, 128a–n, Table 39) 2002 Vermioxytricha arenicola nov. spec.1 – Foissner, Agatha & Berger, Denisia, 5: 751, Fig. 164a–z, 165a, b, 397a–n, Table 146 (Fig. 126a–z, 127a, b, 128a–n; original description; the holotype slide [accession number 2002/234] and five paratype slides [2002/235–239] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; see Foissner et al. 2002, p. 43 and Aescht 2003, p. 380).
Nomenclature: The species-group name arenicola (Latin adjective; living in sand) refers to the preferred habitat (Foissner et al. 2002). Type species of Vermioxytricha. Remarks: The present species differs from the sympatric V. muelleri (both were found in Namibian soils, but not in the same sample; Foissner et al. 2002, p. 63) only by the presence of cortical granules, distinct organelles whose presence/absence we use as a specific feature. Hemisincirra interrupta has only one dorsal kinety, but more macronuclear nodules (about 30 vs. 14). Hemisincirra rariseta (Fig. 86a–g), which has the same body size and shape and nuclear apparatus, lacks cortical granules and (very likely) a buccal cirrus, has more cirri in the frontal field (12 vs. 8), fewer marginal cirri (around 26 per row vs. around 40), and two bipolar kineties (vs. one bipolar and one strongly reduced kinety 2). In life, Vermioxytricha arenicola is recognisable by the vermiform body, the strand-like nuclear figure, the cortical granules, and the two dorsal bristles, one of which is strongly reduced. However, identification of worm-shaped species should be checked in protargol preparations. Morphology: We studied two populations from Africa, namely one from Namibia (type population) and the other from Tunisia (Foissner et al. 2002). They agree almost perfectly, so that conspecificity is beyond reasonable doubt (Table 39). Thus, the diagnosis and the description is from both populations. Body size in life 120–220 × 12–25 µm, usually about 170 × 17 µm; body length:width ratio around 10:1 on average both in life and in protargol preparations (Table 39). Body outline vermiform with anterior portion slightly narrowed and inconspicuously bulged where the adoral zone enters the buccal cavity; posterior region usually tail-like and more or less distinctly curved with slight, but typical subterminal notch at right side (Fig. 126a–g, 128a, g). Body dorsoventrally flattened only in oral area; acontractile, occasionally slightly twisted about main body axis. Nuclear apparatus in central 1
Foissner et al. (2002) provided the following diagnosis: Size about 170 × 17 µm in vivo; vermiform. Usually 13–16 macronuclear nodules in single strand left of midline. Cortical granules around cirri and dorsal bristles and scattered in short rows, about 1 µm across and yellowish. On average 43–46 right and 39 left marginal cirri. Adoral zone occupies 12–14% of body length, comprises 3 frontal and 14–15 ventral membranelles. Invariably 1 bipolar and 1 very short dorsal kinety near anterior end.
Vermioxytricha
601
quarters of cell. Macronuclear nodules in single strand left of midline, individual nodules usually elongate ellipsoidal, rarely globular or ellipsoidal; chromatin bodies minute. Usually 2–4 globular micronuclei along macronuclear strand (Fig. 126a, m, p, 128g). Contractile vacuole ahead of mid-body and, as is usual, near left cell margin with distinct collecting canals during diastole (Fig. 126c, e). Cortex thin and flexible, contains granules around cirral and dorsal bristle bases, but also scattered in unciliated areas; individual granules about 1 µm across and yellowish, usually impregnate distinctly with the protargol method used; when methyl-green pyronin is added, they become red, but are not released; underneath cortex ellipsoidal, about 2 µm-sized, pale structures, likely mitochondria (Fig. 126h, i, o, p, 128g, i, j). Cytoplasm colourless, packed with lipid droplets 1–2 µm across and some vacuoles with a single crystal each; food vacuoles 5 µm across. Movement rapid, winding on microscope slide and between soil particles, showing great flexibility; if numerous, the whirling cells produce a curious spectacle. Adoral zone inconspicuous because occupying only 12–14% of body length, composed of three frontal and 14–15 ventral membranelles of ordinary fine structure separated by a distinct gap at left anterior body corner. Buccal field narrow and flat; buccal lip covers proximal adoral membranelles. Paroral and endoral almost straight, close together, staggered, and likely composed of monokinetids. Pharynx without peculiarities (Fig. 126a, b, e, j–l, o, p, 128a–e, g, h, Table 39). Cirral pattern and number of cirri of usual variability except for the number of marginal cirri, which varies distinctly in the Tunisian specimens (Fig. 126a, j–l, o–q, 128a, f, g, h, Table 39). Most cirri only about 8–10 µm long and fine, each usually consisting of only two or four cilia. Frontal cirri of about same size as other cirri, arranged in oblique pseudorow, with left cirrus (= cirrus I/1) invariably in or near gap between frontal and ventral adoral membranelles. Buccal cirrus in corner of undulating membranes, usually composed of two cilia only. Frontoventral cirri in V-shaped pattern, majority of specimens with the ordinary number of four (detailed designation and variability, see Fig. 126j, k, q and Table 39). Postoral ventral cirri and pretransverse ventral cirri as well as transverse cirri lacking. Right marginal row slightly shortened and extending dorsolaterally anteriorly, ends subterminally. Left row begins left of proximal end of adoral zone, in Tunisian specimens it usually terminates with three narrowly spaced cirri (Fig. 126l). Dorsal bristles about 2 µm long in life, arranged in two kineties. Kinety 1 slightly shortened anteriorly and posteriorly. Kinety 2 usually composed of two basal body pairs near anterior right body corner; note that this kinety is a dorsomarginal kinety. Caudal cirri lacking (Fig. 126m, p, 128a, c–e, Table 39). Cell division (Fig. 126o–z, 127a, b, 128k–n): This part of the life cycle was studied in specimens from the type population (Foissner et al. 2002). The process is similar to that in the oxytrichids (for review, see Berger 1999), but simpler due to the lack of anlage V and therefore a reduced number of cirri. In addition, the remaining anlagen do not form postoral ventral cirri, pretransverse ventral cirri, and transverse cirri.
602
SYSTEMATIC SECTION
Fig. 126a–k Vermioxytricha arenicola (from Foissner et al. 2002. a–d, g–k, Tunisian population; e, f, Namibian site 64 population. a–i, from life; j, k, protargol impregnation). a–g: Ventral view of representative specimen (a, 160 µm) and shape variants (b–g). Arrows in (a, e) mark notch at right posterior margin. h, i: Cortical granulation and mitochondria underneath cortex. j, k: Infraciliature of ventral side of anterior body portion (j = 48 µm, k = 41 µm). Short arrows mark cirrus (= cirrus III/2) behind right frontal cirrus, asterisks mark cirrus IV/3. Long arrow in (j) denotes gap in adoral zone. Frontal cirri connected by dotted
Vermioxytricha
603
Ontogenesis commences with the de-novo formation of a long, narrow oral primordium slightly ahead of mid-body (Fig. 126n, o). Then, four cirral anlagen in forked pattern arise from the anterior end of the oral primordium, where adoral membranelles are already formed (Fig. 126r–t, 128k, l). Next, the rightmost cirral anlage splits transversely to produce an anlage each for the frontoterminal cirri of the proter and the opisthe; thus this streak is a primary primordium and homologous to anlage VI of the 18-cirri hypotrichs. In addition, the parental undulating membranes begin to reorganise, and the buccal cirrus, cirrus III/2, and the rear frontoventral cirrus modify to proter’s anlagen I–IV. This resembles the Steinia-pattern shown in Berger (1999, Fig. 23e in this book), except that anlage V is not formed. In middle dividers, five frontal-ventral cirri anlagen (I–IV, VI) are recognisable in both filial products (Fig. 126v, x, y, 128l). In late dividers, frontal-ventral cirri begin to segregate and arrange to the species-specific pattern, showing that neither postoral and pretransverse ventral cirri nor transverse cirri are formed (Fig. 126z, 127a, 128m). On average, eight cirri are formed, namely, anlage I forms – as is usual – the left frontal cirrus (= cirrus I/1); II forms the middle frontal cirrus (II/3) and the buccal cirrus (II/2); III forms the right frontal cirrus (III/3) and one cirrus (= III/2) behind it (rarely two cirri behind of it are formed; e.g., Fig. 126k); IV forms the rearmost cirrus of the frontoventral cirri (= IV/3); anlage V is lacking in Vermioxytricha; VI forms the two frontoterminal cirri (VI/3, VI/4) which are part of the frontoventral cirri (see Fig. 6a in Berger 1999 for homologisation and detailed designation of cirri). Consequently, Vermioxytricha is an 18-cirri hypotrich without the three postoral ventral cirri, the two pretransverse ventral cirri, and the five transverse cirri. Nuclear division and formation of the remaining ventral ciliature occur in the usual manner (Fig. 126w–z, 127a, b, 128n). Ontogenesis of the dorsal ciliature commences with the formation of two anlagen within parental kinety 1 (Fig. 126w). Later, a very short dorsomarginal kinety (= kinety 2) originates at/near the anterior end of the right marginal row anlagen (Fig. 126z, 127a). No caudal cirri are produced. This pattern corresponds the Urosomoida pattern (Berger 1999, p. 73), except that not three bipolar kineties are present, but only one (see remarks under genus section). Occurrence and ecology: Very likely confined to terrestrial habitats (Foissner et al. 2002, p. 56). Type locality of V. arenicola is the Sossus Vlei (24°50'S, 15°20'E) of the Namib Desert, where Foissner et al. (2002; site 24) discovered it in humous sand under Acacia erioloba trees (Camel thorn). It occurred with high abundance in the non-flooded Petri dish cultures from the type locality and areas nearby, indicat-
b
line (j), broken lines connect cirri originating from same anlage, and frontoventral cirri circled in (j). Dotted line in (k) connects frontoventral cirri arranged in V-shaped pattern. Arrowhead in (k) denotes a second cirrus behind the right frontal cirrus. AZM = adoral zone of membranelles, BC = buccal cirrus, CG = cortical granules, CV = contractile vacuole, E = endoral, FC = left (a) and right (j) frontal cirrus, FT = frontoterminal cirri, LMR = left marginal row, MO = mitochondria, P = paroral, PF = pharyngeal fibres, RMR = right marginal row, I, II = frontal-ventral cirri anlagen. Page 600.
604
SYSTEMATIC SECTION
Vermioxytricha
605
Fig. 126q–s Vermioxytricha arenicola (from Foissner et al. 2002. Type population. Early dividers after protargol impregnation). q: Infraciliature of oral region of type specimen (see also Fig. 126o, p). Short arrow marks gap in adoral zone of membranelles, long arrow denotes cirrus IV/3 which is the rearmost frontoventral cirrus. Broken lines connect cirri originating from same anlage. r, s: Early dividers showing the cuneate oral primordium and three dikinetidal cirral anlagen in forked pattern (arrowheads). AZM = distal end of bipartite adoral zone of membranelles, OP = oral primordium, PF = pharyngeal fibres, RMR = right marginal row. Page 600.
ing the presence of many resting cysts and optimal growth conditions. The Tunisian sample (collected by Thomas Peer, University of Salzburg), was an alkaline (pH 8.1), cyanobacteria-covered crust soil mixed with red sand and some litter from drifting sand of the eastern part of the Grand Erg (33°N 09°E). The present species is – like V. muelleri which occurred in the same Namibian area – well adapted to the sandy habitat by its worm-shaped body, and is therefore rather common in the dunes of the Namib Desert (Foissner et al. 2002).
b
Fig. 126l–p Vermioxytricha arenicola (from Foissner et al. 2002. l, m, Tunisian population; n–p, type population. Protargol impregnation). l, m: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 193 µm. Frontal cirri connected by dotted line. Arrowhead possibly marks a cirrus originating, like cirrus III/2, from anlage III. Asterisk denotes, very likely, cirrus IV/3. Arrow denotes narrowly spaced cirri near body end, likely the rear end of the left marginal row. n: Part of very early divider showing the oral primordium as a short series of single basal bodies slightly ahead of mid-body between the marginal cirral rows. o, p: Infraciliature of ventral and dorsal side, cortical granulation, and nuclear apparatus of holotype specimen (192 µm) which is a very early divider. C = cirrus, CG = cortical granules, LMR = left marginal row, MA = macronuclear nodule, MI = micronucleus, OP = oral primordium, RMR = right marginal row, 1, 2 = dorsal kineties. Page 600.
606
SYSTEMATIC SECTION
Fig. 126t–w Vermioxytricha arenicola (from Foissner et al. 2002. Type population. Early and middle dividers after protargol impregnation). t: Four cirral anlagen are recognisable in the opisthe (arrowheads); the rightmost splits transversely to form anlage VI for proter (anterior portion; arrow) and opisthe (posterior portion). Length of part visible = 114 µm. u: The parental undulating membranes, the buccal cirrus, the cirrus behind the right frontal cirrus (arrowhead), and the rearmost frontoventral cirrus (short arrow) are modified to anlagen I–IV of the proter; anlage VI is the front portion
Vermioxytricha
607
Vermioxytricha arenicola feeds on rod-shaped bacteria, yeast cells, short fungal hyphae, and debris (Foissner et al. 2002).
Vermioxytricha muelleri (Foissner, 1986) Foissner, Agatha & Berger, 2002 (Fig. 129a–f, Table 39) 1986 Hemisincirra muelleri nov. spec.1 – Foissner, Zool. Jb. Syst., 113: 47, Abb. 1a–f, Tabelle 2 (Fig. 129a–f; original description; the holotype slide and one paratype slide are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria [Foissner 1986, p. 47]; however, slides not listed by Aescht 2003, p. 391). 2001 Hemisincirra muelleri Foissner, 1986 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 30 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Vermioxytricha muelleri (Foissner, 1986) nov. comb. – Foissner, Agatha & Berger, Denisia, 5: 750 (combination with Vermioxytricha).
Nomenclature: Foissner (1986) dedicated this species to Hans Joachim Müller, editor of the journal “Zoologische Jahrbücher”. Remarks: The present species is very similar to V. arenicola so that we transferred it to Vermioxytricha, which differs from Hemisincirra (p. 387), inter alia, by the presence of a dorsomarginal kinety (further details see genus section and V. arenicola). Hemisincirra interrupta has only one dorsal kinety, but more macronuclear nodules (about 30 vs. 14). Hemisincirra rariseta (Fig. 86a–g), which has the same size, shape, and nuclear apparatus, also lacks cortical granules, but has more cirri in the frontal field (12 vs. 8), likely lacks a buccal cirrus, has fewer marginal cirri (around 26 per row vs. around 40), and two bipolar kineties (vs. one bipolar and one strongly reduced kinety 2). Engelmanniella mobilis (Engelmann, 1862) Foissner, 1982 – which has a rather similar habitus – has cortical granules and four marginal rows (Foissner 1982, p. 66; Wirnsberger-Aescht et al. 1989, 1990). Morphology: Body size in life 140–200 × 10–20 µm, body length:width ratio 10.9:1 on average (Table 39). Body outline slender, vermiform, anterior end narrowly
b
(long arrow) of anlage VI of the opisthe, that is, it is a primary primordium. Length of part visible = 104 µm. v, w: Ventral and dorsal side of same specimen (170 µm) showing five anlagen each in proter and opisthe. Arrow marks some surplus basal bodies from the primary primordium (see above), arrowheads in (w) denote dorsal kinety anlagen within parental kinety 1. FT = frontoterminal cirri, LMR = left marginal row, MI = micronucleus, OP = oral primordium, RMR = right marginal row, I–IV, VI = frontal-ventral cirri anlagen (note that anlage V is lacking in Vermioxytricha!), 1, 2 = dorsal kineties. Page 600. 1
Foissner (1986) provided the following diagnosis: In vivo etwa 140–200 × 10–20 µm große, postoral drehrunde Hemisincirra, deren Frontalreihe kürzer ist als die adorale Membranellenzone. Durchschnittlich 16 adorale Membranellen, von denen die 3 vorderen durch eine sehr breite Lücke von den hinteren getrennt sind. Links der Frontalreihe 1 isolierter Cirrus. Makronucleus kettenförmig, besteht aus durchschnittlich 15 ellipsoiden Nodien. 2 Dorsalkineten, von denen die rechte stark verkürzt ist.
608
SYSTEMATIC SECTION
Vermioxytricha
Fig. 127a, b Vermioxytricha arenicola (from Foissner et al. 2002. Type population. Very late divider after protargol impregnation). Parental cirri white, new black. Both in proter and opisthe the ordinary set of eight cirri (three frontal, one buccal, four frontoventral) has been formed. Broken lines connect cirri originating from the same anlage. Frontoventral cirri of opisthe circled. Arrows in (a) mark new dorsomarginal rows (= kineties 2), which are usually composed of two basal body pairs only. Arrows in (b) mark rear end of dorsal kinety 1 anlagen, showing that no caudal are formed. FT = parental frontoterminal cirri, 1, 2 = dorsal kineties. Page 600.
b
Fig. 126x–z Vermioxytricha arenicola (from Foissner et al. 2002. Type population. A middle and a late divider after protargol impregnation). Parental structures white, new black. x, y: Ventral and dorsal side of a specimen with five anlagen each in proter and opisthe. Arrow marks some surplus basal bodies from primary primordium, arrowheads denote marginal row primordia. z: Late divider with nine new cirri in frontal field of proter and the usual set of eight cirri in the opisthe. Arrow denotes new buccal cirrus of proter. Arrowhead denotes new dorsomarginal kinety (= kinety 2) of opisthe. Broken lines connect cirri originating from same anlage. FT = parental frontoterminal cirri, I–IV, VI = frontal-ventral cirri anlagen, 1 = dorsal kinety 1. Page 600.
609
610
SYSTEMATIC SECTION
Fig. 128a–f Vermioxytricha arenicola (from Foissner et al. 2002. Scanning electron micrographs of type population). a: Dorsoventral view showing vermiform, slightly twisted body and the minute adoral zone. Arrow marks two supernumerary (parental?) marginal cirri. b: Ventral view showing the narrow, flat buccal cavity (asterisk). c: Left anterior portion. d, e: Dorsolateral and dorsal view of anterior body region showing the gap (arrow in d) between frontal and ventral adoral membranelles and the two dorsal kineties (arrowed numbers). f: Ventral view of pointed posterior region lacking transverse and caudal cirri. Explanation of original labelling: AZM = adoral zone of membranelles, BL = buccal lip, BU = buccal cirrus, FC1 = left frontal cirrus, FM = frontal adoral membranelles, FS = frontal scutum, FT = frontoterminal cirri, LMR = left marginal row, PM = paroral, RMR = right marginal row, VM = ventral adoral membranelles, 1, 2 = dorsal kineties. Page 600.
Vermioxytricha
611
Fig. 128g–n Vermioxytricha arenicola (from Foissner et al. 2002. Specimens of a Namibian population after protargol impregnation). g, j: Ventral view showing oral apparatus (arrowheads) and cortical granules (arrows; dots are scattered cortical granules). h: Ventral view of anterior body half. Small arrow marks frontoventral cirrus III/2, large arrow marks gap in adoral zone. i: Dorsal view showing cortical granules (arrowheads). k: Early divider developing two primordia (arrowheads). l: Middle divider showing the five frontal-ventral cirri anlagen (arrowheads). m, n: Late and very late divider. Arrowheads in (m) mark left and right frontal cirrus. Explanation of original labelling: AZM = adoral zone of membranelles, BU = buccal cirrus, C = cirri, DB = dorsal bristles, EM = endoral, FC3 = right frontal cirrus, FM = frontal adoral membranelles, FVR = frontoventral cirri, LMR = left marginal row, MA = macronuclear nodules, MI = micronucleus, OP = oral primordium, PM = paroral, RMR = right marginal row, UM = undulating membrane, VM = ventral adoral membranelles. Page 600.
612
SYSTEMATIC SECTION
rounded, posterior portion pointed and usually curved leftwards (Fig. 129a–c); often sigmoidal. Body very flexible, and only anterior portion slightly flattened dorsoventrally. Macronuclear nodules in middle body portion behind adoral zone of membranelles left of midline; individual nodules with many small chromatin bodies. One ellipsoidal micronucleus each in anterior and posterior third of nuclear apparatus. Contractile vacuole somewhat ahead of mid-body near left cell margin; during diastole with two long collecting canals (Fig. 129b). Pellicle colourless, very flexible, close underneath many, about 3.5 × 1.3 µm-sized ellipsoidal mitochondria. Cortical granules (termed subpellicular granules in original description) not present. Cytoplasm colourless, densely granulated and therefore opaque; in posterior body portion some colourless, cuboid-shaped crystals. Food vacuoles only about 5 µm across, usually they seem to be empty. Movement fast gliding or winding like a worm between soil particles. Adoral zone occupies about 13% of body length on average (Fig. 129a, b, d, f, Table 39), composed of 16 membranelles of ordinary fine structure; distalmost three membranelles set off from proximal portion of adoral zone by distinct gap. Buccal field very small, rather shallow, anteriorly bordered by buccal cirrus, on right side by the inconspicuous undulating membranes. Cirral pattern and number of cirri of usual variability (Fig. 129a, d, f, Table 39). Cirri very fine, about 10 µm long. Frontal cirri almost of same size as remaining cirri, left one (= cirrus I/1) in gap of adoral zone. Almost invariably four frontoventral cirri arranged in V-shaped pattern (Fig. 129f). Postoral and pretransverse ventral cirri and transverse cirri lacking. Buccal cirrus ahead of undulating membranes, composed of two basal bodies (cilia) only. Right marginal row commences about at level of buccal cirrus (note that in the original description the right marginal row is, par lapsus, labelled as left marginal row), terminates – like left row – at rear cell end; left marginal row begins near rear end of adoral zone. Distance between individual cirri in posterior third slightly larger than in anterior portion of marginal rows. Rear body end with 4–5 spread cirri; according to original description these are transverse or caudal cirri, while according to ontogenetic data on V. arenicola neither transverse nor caudal cirri are present, indicating that these cirri are marginal cirri. Dorsal bristles about 2–3 µm long, invariably arranged in two kineties (Fig. 129e); kinety 1 more or less distinctly shortened anteriorly and posteriorly. Kinety 2 composed of two basal body pairs near anterior cell end; likely it is a dorsomarginal row (cp. V. arenicola which has exactly the same pattern). Caudal cirri absent (see also end of previous paragraph). Occurrence and ecology: Very likely confined to terrestrial habitats (Foissner 1987, 1998). Type locality of V. muelleri is the Pisang Peak, a mountain in the Annapurna area, Nepal. Foissner (1986) found it there in a soil sample (0–5 cm) collected by Eduard Vierthaler (a fellow student of mine, now head of the alpine school in Filzmoos, Salzburg) at a steep slope (altitude about 3300 m); the slope was grown with shrubs and grazed by yaks. Further records: soil from the Apsheron peninsula,
Vermioxytricha
613
Fig. 129a–f Vermioxytricha muelleri (from Foissner 1986. a–c, from life; d–f, protargol impregnation). a: Ventral view of representative specimen, 188 µm. Arrow marks spread cirri at rear end (likely left marginal cirri). b: Dorsal view, 184 µm. c: Lateral view. d, e: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 104 µm. f: Infraciliature of anterior body portion in ventral view and nuclear apparatus, 36 µm. Long arrow marks gap in adoral zone, short arrow denotes cirrus III/2. Frontal cirri connected by dotted line, frontoventral cirri circled (cirrus IV/3 marked with asterisk). Broken lines connect cirri originating from same anlage. AZM = distal end of adoral zone, BC = buccal cirrus, CV = contractile vacuole with collecting canals, FT = frontoterminal cirri (originate from anlage VI), LMR = anterior end of left marginal row, MA = macronuclear nodule, MI = micronucleus, PF = pharyngeal fibres, RMR = anterior end of right marginal row, 1, 2 = dorsal kineties (kinety 2 is very likely a dorsomarginal row; see type species). Page 607.
Azerbaijan (Alekperov & Musayev 1988, p. 1909); in three out of 73 soil samples from Namibia, that is, occurs sympatric with V. arenicola (Foissner et al. 2002, p. 63). Vermioxytricha muelleri feeds on bacteria (Foissner 1986). Biomass of 106 specimens about 11 mg (Foissner 1987, p. 124).
614
SYSTEMATIC SECTION
Hemiurosoma Foissner, Agatha & Berger, 2002 2002 Hemiurosoma nov. gen.1 – Foissner, Agatha & Berger, Denisia, 5: 834 (original description). Type species (by original designation): Hemiurosoma terricola Foissner, Agatha & Berger, 2002.
Nomenclature: Hemiurosoma is a composite of the Greek word hemi (half) and the genus-group name Urosoma (tailed body), referring to the similarity with Urosoma Kowalewskiego, 1882. Neuter gender (ICZN 1999, Article 30.1.2; see note on p. 635). Characterisation (A = supposed apomorphy): Dorsomarginalia with oral apparatus in Gonostomum pattern. Three frontal cirri. Buccal cirrus present. Frontoventral cirri arranged in Urosoma pattern. Frontal-ventral-transverse cirri anlage V lost (A?). Anlagen II–IV and VI originate from primary primordia, form less than the plesiomorphic number of 17 cirri so that number of postoral ventral cirri, pretransverse ventral cirri, and transverse cirri is reduced or groups are even lacking (A?). One right and one left marginal row. Four dorsal kineties. Dorsomarginal kinety present. Dorsal kinety fragmentation lacking. Caudal cirri present. Terrestrial. Additional characters: The four species are have a rather uniform ventral and dorsal infraciliature. In addition to the features mentioned above, they are characterised, inter alia, by the following plesiomorphies: distinct mitochondria underneath pellicle (lacking in H. similis). Dorsal bristles short, that is, 2–4 µm long. Three caudal cirri. Remarks: Hemiurosoma was established for two species occurring in Namibian soils (Foissner et al. 2002). In addition, we added two Hemisincirra species which I have transferred to Urosoma in the review on oxytrichids (Berger 1999). However, I already stated that they very likely form a distinct clade within Urosoma. Urosoma octonucleata Berger & Foissner, 1989 has the three postoral ventral cirri arranged more or less perfectly in line, whereas in other Urosoma species, and in the 18-cirri hypotrichs in general, the front cirrus (= cirrus IV/2) is shifted leftwards (for review, see Berger 1999, p. 396). Perhaps the cirri of U. octonucleata do not originate from anlagen IV and V as in Urosoma, but exclusively from anlage IV, as the postoral ventral cirri of Erimophrya quadrinucleata (Fig. 125c, f). In addition, Urosoma octonucleata has, like H. goertzi, only 3–4 transverse cirri which is a further hint that one anlage is lacking. When ontogenetic data prove this hypothesis, then U. octonucleata has to be transferred to Hemiurosoma. Hemiurosoma comprises Urosoma-relatives with a distinctly reduced number of postoral ventral cirri, pretransverse ventral cirri, and transverse cirri, mainly due to the loss of cirral anlage V (Foissner et al. 2002). Because of the loss of all postoral ventral cirri in H. terricola (and H. similis), all cirral anlagen of the opisthe originate de novo via the de novo-developing oral primordium (Fig. 130g). Urosoma and 1
Foissner et al. (2002) provided the following diagnosis: Oxytrichidae with oral apparatus in Gonostomum pattern and frontoventral cirri in Urosoma pattern. 2 or less postoral ventral cirri. 5 or less pretransverse and transverse cirri. 1 right and 1 left row of marginal cirri. 4 dorsal kineties. Caudal cirri present. Fronto-ventral-transverse cirri originate from 5 anlagen, four of which are primary primordia. Dorsal ontogenesis in Urosomoida pattern.
Hemiurosoma
615
Hemiurosoma have the same frontoventral cirri pattern with cirrus III/2 somewhat left and ahead of the remaining cirri which form a line (Fig. 130f, 131h, 132a, b). In addition to this Urosoma pattern, in both taxa (i) the oral apparatus is Gonostomumlike (see Berger 1999, p. 56), including the undulating membranes; (ii) the frontalventral-transverse cirri anlagen originate from primary primordia; and (iii) the subcortical mitochondria are very distinct. All these features strongly indicate a sistergroup relationship of Hemiurosoma and Urosoma with the loss of anlage V as apomorphy of Hemiurosoma. Foissner et al. (2002) described some other Dorsomarginalia which have also lost anlage V, namely, Erimophrya, Vermioxytricha, and Apourosomoida. However, they differ from Hemiurosoma in the features mentioned above, strongly indicating that the loss of anlage V occurred independently in Hemiurosoma and these taxa (see introduction on p. 560). Originally, we classified Hemiurosoma in the oxytrichids (Foissner et al. 2002) because of the obvious relationship to Urosoma, which was assigned to the oxytrichids, inter alia, by Berger & Foissner (1997) and Berger (1999). In these papers, we considered Urosoma and Gonostomum as sister-groups because both have a Gonostomum-like oral apparatus. The distinct differences in the dorsal kinety pattern were explained by a simplification of the Oxytricha pattern (dorsomarginal row and kinety fragmentation present) via the Urosomoida pattern (also present in Urosoma; with dorsomarginal row, but loss of fragmentation) to the Gonostomum pattern (dorsomarginal row also lost). However, this is a somewhat complicated, that is, not very parsimonious explanation, but not impossible as indicated by some (not all!) molecular analyses which classify Hemiurosoma in a subcluster, inter alia, containing Oxytricha granulifera and Onychodromopsis flexilis1, both of which have the Oxytricha pattern (Foissner et al. 2004, Foissner & Stoeck 2006, Gong et al. 2006). However, since some years I suppose that the last common ancestor of the Hypotricha had three dorsal kineties (like Gonostomum) which divided by intrakinetal proliferation (see ground pattern of hypotrichs, chapter 2.2 of general section), a mode still present, for example, in the urostyloids (for review, see Berger 2006). In a nonurostyloid branch the dorsomarginal rows evolved, a curious apomorphy of the Dorsomarginalia (Berger 2006). Later, in one member of the Dorsomarginalia a dorsal kinety fragmentation occurred (probably in kinety 3) and this species was very likely the last common ancestor of the oxytrichids (Fig. 9a). According to this hypothesis, the Urosoma/Hemiurosoma group belongs to the Dorsomarginalia (dorsomarginal kinety present), but not to the oxytrichids because a kinety fragmentation is lacking. Note that the 18-cirri pattern in Urosoma and its morphogenesis is not a proof that it belongs to the Oxytrichidae because the 18-cirri pattern corresponds the ground pattern of the Hypotricha (Berger 2006, p. 33; chapter 2.2 of the general section of present book). As mentioned above, Berger & Foissner (1997) and Berger (1999) assumed a close relationship of Urosoma and Gonostomum because of the similar oral apparatus. Foissner et al. (2004) and Gong et al. (2006) found a relatively close relation1
This is Allotricha antarctica according to Berger (1999, p. 268).
616
SYSTEMATIC SECTION
Table 40 Morphometric data on Hemiurosoma goertzi (go1, from non-flooded Petri dish method; go2, from flooded conditions; both from Foissner et al. 2002) and Hemiurosoma terricola (ter, from Foissner et al. 2002) Characteristics a
species
Body, length
Body, width
Body length:width, ratio
Adoral zone of membranelles, length
Body length:length of adoral zone, ratio
Anterior body end to paroral, distance Paroral, length Anterior body end to endoral, distance Endoral, length Anterior body end to frontmost frontoventral cirrus, distance Anterior body end to rearmost frontoventral cirrus, distance Anterior body end to buccal cirrus, distance Anterior body end to right marginal row, distance Anterior body end to postoral ventral cirrus, distance Anterior body end to frontmost macronuclear nodule, distance Macronuclear nodules, distance in between Anterior macronuclear nodule, length
Anterior macronuclear nodule, width
Macronuclear nodules, number Anterior micronucleus, length Anterior micronucleus, width
go1 go2 ter go1 go2 ter go1 go2 ter go1 go2 ter go1 go2 ter go1 ter go1 ter go1 ter go1 ter go1 ter go1 ter go1 ter go1 ter go1 go1 ter go1 go1 go2 ter go1 go2 ter go1 ter go1 ter go1 ter
mean
M
132.3 130.0 235.5 238.0 138.5 138.0 22.3 22.0 41.5 43.0 19.2 20.0 6.0 5.9 5.8 5.4 7.3 7.3 28.9 29.0 41.0 41.0 24.5 25.0 4.6 4.5 5.8 5.9 5.7 5.6 14.1 14.0 11.8 12.0 8.1 8.0 3.6 4.0 16.7 17.0 16.4 16.0 9.8 10.0 5.6 6.0 10.5 10.0 9.0 9.0 24.0 24.0 19.1 19.0 14.9 15.0 11.7 12.0 9.1 9.0 7.4 7.0 32.0 32.0 31.7 31.7 14.5 15.3 16.6 8.5 5.9 6.7 4.3 2.0 4.1 4.2 3.3 2.8 1.8
32.0 31.0 16.0 16.0 17.0 9.0 6.0 6.0 4.0 2.0 4.0 4.0 3.0 3.0 2.0
SD
SE
CV
Min
Max
n
16.4 30.7 10.5 3.2 5.6 2.1 0.5 1.0 0.8 3.0 2.5 2.1 0.6 0.8 0.7 1.7 1.5 1.1 0.6 1.7 2.3 1.1 0.6 0.8 0.8 2.6 1.3 1.7 1.6 2.0 1.0 3.3
4.2 7.9 2.4 0.8 1.4 0.5 0.1 0.3 0.2 0.8 0.6 0.5 0.2 0.2 0.2 0.4 0.4 0.3 0.1 0.4 0.5 0.3 0.1 0.2 0.2 0.7 0.3 0.4 0.4 0.5 0.2 0.9
12.4 95.0 157.0 13.0 184.0 284.0 7.6 117.0 161.0 14.5 16.0 27.0 13.5 30.0 48.0 11.1 15.0 22.0 8.5 5.0 6.8 18.0 4.6 8.8 10.9 5.3 8.7 10.4 23.0 33.0 6.0 36.0 45.0 8.8 19.0 28.0 13.9 3.8 5.8 14.2 4.1 7.1 11.6 4.7 7.3 12.2 10.0 16.0 13.0 9.0 15.0 13.3 6.0 10.0 17.0 3.0 5.0 10.1 13.0 18.0 14.0 14.0 21.0 11.2 8.0 12.0 10.6 5.0 7.0 8.0 9.0 12.0 9.1 7.0 10.0 10.8 18.0 28.0 7.0 17.0 22.0 11.3 11.0 17.0 13.4 8.0 14.0 22.2 6.0 13.0 13.7 6.0 9.0 10.4 26.0 36.0
15 15 19 15 15 19 15 15 19 15 15 19 15 15 19 15 19 14 19 15 19 13 19 15 19 14 19 15 19 15 19 15
3.4 1.9 5.2 2.3 1.5 1.8 0.5 1.0 0.6 0.0 – 1.1 – 0.6 –
0.9 0.4 1.3 0.6 0.4 0.4 0.1 0.3 0.1 0.0 – 0.3 – 0.1 –
10.7 6.1 35.7 15.4 9.1 21.2 8.8 14.6 13.5 0.0 – 27.1 – 20.3 –
15 19 15 15 15 19 15 15 19 15 30 15 19 15 19
25.0 28.0 7.0 10.0 14.0 6.0 5.0 5.0 3.0 2.0 4.0 2.5 3.0 1.5 1.5
36.0 35.0 24.0 18.0 20.0 13.0 7.0 8.0 5.0 2.0 5.0 6.0 4.0 4.0 2.0
Hemiurosoma
617
Table 40 Continued Characteristics a Micronuclei, number
Nuclear figure, length Posterior body end to rear transverse cirrus, distance Posterior body end to front transverse cirrus, distance Adoral membranelles, number
Frontal cirri, number Buccal cirri, number Frontoventral cirri, number Postoral ventral cirri, number Transverse cirri, number Right marginal cirri, number
Left marginal cirri, number Dorsal kineties, number Caudal cirri, number
species
mean
M
SD
SE
CV
Min
Max
n
go1 go2 ter go1 ter go1 go2 ter go1
1.7 1.7 2.1 45.7 45.0 11.1 22.0 1.3 15.6
2.0 2.0 2.0 45.0 45.0 10.5 22.0 1.0 14.0
– – – 6.2 4.8 2.6 4.9 – 3.8
– – – 1.6 1.1 0.7 1.3 – 1.0
– – – 13.7 10.7 23.9 22.1 – 24.5
1.0 0.0 2.0 37.0 35.0 8.0 10.0 1.0 10.0
2.0 3.0 3.0 58.0 51.0 16.0 30.0 2.0 22.0
15 15 19 15 19 14 15 19 14
go1 go2 ter go1 ter go1 ter go1 ter go1 ter go1 ter go1 go2 ter go1 ter go1 ter go1 ter
27.3 25.3 21.1 3.0 3.0 1.0 1.0 4.0 4.1 1.0 0.0 3.1 2.0 38.6 39.1 37.0 34.5 31.8 4.0 4.0 3.0 3.0
27.0 26.0 21.0 3.0 3.0 1.0 1.0 4.0 4.0 1.0 0.0 3.0 2.0 39.0 39.0 37.0 35.0 31.0 4.0 4.0 3.0 3.0
2.1 1.8 0.9 0.0 0.0 0.0 0.0 0.0 – 0.0 0.0 – 0.0 2.4 3.1 1.8 2.7 2.9 0.0 0.0 0.0 0.0
0.6 0.5 0.2 0.0 0.0 0.0 0.0 0.0 – 0.0 0.0 – 0.0 0.6 0.8 0.4 0.7 0.7 0.0 0.0 0.0 0.0
7.9 6.9 4.4 0.0 0.0 0.0 0.0 0.0 – 0.0 0.0 – 0.0 6.3 8.0 4.7 7.8 9.2 0.0 0.0 0.0 0.0
25.0 22.0 19.0 3.0 3.0 1.0 1.0 4.0 4.0 1.0 0.0 3.0 2.0 34.0 34.0 35.0 28.0 28.0 4.0 4.0 3.0 3.0
30.0 28.0 23.0 3.0 3.0 1.0 1.0 4.0 5.0 1.0 0.0 4.0 2.0 43.0 46.0 41.0 39.0 40.0 4.0 4.0 3.0 3.0
11 15 19 15 19 15 19 14 19 15 30 14 19 15 15 19 15 19 8 19 7 19
a All measurements in µm. CV = coefficient of variation in %, M = median, Max = maximum value, mean = arithmetic mean, Min = minimum value, n = number of individuals investigated, SD = standard deviation, SE = standard error of arithmetic mean. Data based on protargol-impregnated specimens (H. goertzi was fixed in Stieve's solution amended with some drops of osmium tetroxide for better preservation; Foissner et al. 2002).
ship of Hemiurosoma and Gonostomum using the small subunit rRNA. By contrast, Hemiurosoma and Gonostomum are not closely related according to Foissner & Stoeck (2006) and Schmidt et al. (2007; their Fig. 2), indicating that the gonostomoid oral apparatus evolved twice independently. Unfortunately, Gonostomum clusters rather differently in these two Hypotricha trees. The data by Schmidt et al. (2007) suggest that it branches off rather near the base of the tree, that is, outside the cluster comprising the Dorsomarginalia (ranging from Oxytricha lanceolata to Uroleptus piscis in their Fig. 2). This position supports the hypothesis that Gonostomum
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SYSTEMATIC SECTION
has the plesiomorphic dorsal kinety pattern composed of three bipolar rows. As just mentioned, this hypothesis requires the assumption that the gonostomoid oral apparatus evolved convergently in Urosoma/Hemiurosoma and Gonostomum, as also indicated by some phylogenetic analyses based on SSrRNA (see above). According to the sequence analysis by Foissner et al. (2004), Hemiurosoma terricola (type species) clusters with Engelmanniella mobilis (Engelmann, 1862) Foissner, 1982, a species of uncertain position according to molecular and morphological data. A more or less similar position was estimated by Foissner & Stoeck (2006) and Schmidt et al. (2007). Engelmanniella has one bipolar and one short kinety, which, however, does not originate dorsomarginally, but – like the bipolar row – intrakinetally1 (Wirnsberger-Aescht et al. 1989). According to this pattern, Engelmanniella does not belong to the Dorsomarginalia, that is, it must branch off very basally. Such a position was estimated by Dalby & Prescott (2004) using the micronuclear gene encoding actin I. The close relationship of “Paraurostyla viridis” (basionym Urostyla viridis) in their tree must not be overinterpreted for two reasons: (i) this species is little-known and was never redescribed using modern methods, that is, the cirral and dorsal kinety pattern are basically not known. For that reason, Berger (2006) classified it in Urostyla, as originally proposed by Stein (1859); (ii) the population used for molecular analysis was identified by a non-taxonomist, and since morphological data have not been provided the identification of this population cannot be verified. For further details, see my review on the urostyloids (Berger 2006, p. 1106). Species included in Hemiurosoma (alphabetically arranged basionyms are given): (1) Hemisincirra polynucleata Foissner, 1984; (2) Hemiurosoma goertzi Foissner, Agatha & Berger, 2002; (3) Hemiurosoma terricola Foissner, Agatha & Berger, 2002; (4) Perisincirra similis Foissner, 1982.
Key to Hemiurosoma species If you know that your specimen/population belongs to Hemiurosoma (frontoventral cirri in Urosoma pattern), then the identification is rather simple and needs only live observation because the number of macronuclear nodules and postoral ventral cirri (lacking or one cirrus present) are relevant. If you cannot identify your hypotrich with the key below, see also Hemisincirra (dorsomarginal kinety lacking; p. 387), Vermioxytricha (postoral ventral cirri, pretransverse ventral cirri, transverse cirri, and caudal cirri absent; p. 596), and Erimophrya (frontoventral cirri arranged V-like; p. 577). 1 Three postoral ventral cirri present . . . . . . . . Urosoma (see Berger 1999, p. 396) - One or no postoral ventral cirrus (e.g., Fig. 132a, b). . . . . . . . . . . . . . . . . . . . . . 2 1
Because kinety 2 is very short, the anlage for the opisthe originates de novo at the same level as the anlagen for kinety 1 and the marginal rows occur (Wirnsberger-Aescht et al. 1989).
Hemiurosoma 2 3 4 -
619
Two macronuclear nodules (e.g., Fig. 132b). . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Four (rarely five) or about eight macronuclear nodules. . . . . . . . . . . . . . . . . . . . 4 One postoral ventral cirrus (Fig. 131a–m). . . . . . . Hemiurosoma goertzi (p. 628) No postoral ventral cirrus (Fig. 132b). . . . . . . . . . . Hemiurosoma similis (p. 633) (2) Four macronuclear nodules (Fig. 130a–i). . . . Hemiurosoma terricola (p. 619) 6–9, usually eight macronuclear nodules (Fig. 132a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemiurosoma polynucleatum (p. 634)
Hemiurosoma terricola Foissner, Agatha & Berger, 2002 (Fig. 130a–v, Table 40) 2002 Hemiurosoma terricola nov. spec.1 – Foissner, Agatha & Berger, Denisia, 5: 835, Fig. 182a–u, 381q, 400a–j, Table 160 (Fig. 130a–v; original description; the holotype slide [accession number 2002/127] and three paratype slides [2002/128–130] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; see Foissner et al. 2002, p. 39 and Aescht 2003, p. 398). 2004 Hemiurosoma terricola Foissner et al., 2002 – Foissner, Moon-van der Staay, van der Staay, Hackstein, Krautgartner & Berger, Europ. J. Protistol., 40: 267, 273 (analysis of small subunit ribosomal RNA gene sequence of Zambezi population described by Foissner et al. 2002; GenBank accession number AY498651). 2006 Hemiurosoma terricola – Foissner & AL-Rasheid, Acta Protozool., 45: 11, Fig. 23, 37 (detailed study about oral apparatus of hypotrichs [= stichotrichines in their paper]).
Nomenclature: The species-group name terricola is a composite of the Latin noun terr·a (soil), the thematic vowel ·i-, and the Latin verb colere (to live) and means living in soil, the habitat where the species was discovered (Foissner et al. 2002). Usually, species-group names ending with -cola are considered as appositive substantives and are thus not changed when transferred to a genus of different gender (Werner 1972, p. 138). Type species of Hemiurosoma. Remarks: Hemiurosoma terricola has four macronuclear nodules and is thus clearly separated from H. similis, which has only two. However, all other features are very similar and would not justify separation at species level. Generally, the present species is easily confused with the congeners and several Urosomoida species (for review see Berger 1999, p. 345). Thus, the following combination of features is important for live identification (Foissner et al. 2002): body long and slender (about 150 × 20 µm), four macronuclear nodules, frontoventral cirri in Urosoma pattern, no postoral ventral cirri, transverse cirri very near to rear body end, buccal field very flat and narrow. For some notes on the phylogenetic position of Hemiurosoma based on morphological and molecular data see remarks in genus section. Morphology: For scanning electron micrographs of the Zambezi population, see Foissner et al. (2002, p. 1339, 1340). Body size 120–180 × 15–25 µm, on average about 150 × 20 µm in life; body length:width ratio 5.3–8.7:1, on average 7.3:1 in 1
Foissner et al. (2002) provided the following diagnosis: Size about 150 × 20 µm in vivo; slenderly lanceolate. 4 macronuclear nodules and 21 adoral membranelles on average.
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SYSTEMATIC SECTION
Fig. 130a–g Hemiurosoma terricola (from Foissner et al. 2002. a–f, from life; g, protargol impregnation). a: Ventral view of a representative specimen, 150 µm. b, c: Shape variants. Arrow marks dorsal furrow. d: Optical section showing aggregates of minute fat globules in the cytoplasm. e: Surface showing subcortical mitochondria. f: Ventral side of anterior body portion showing, inter alia, the broad buccal lip, covering the minute buccal cavity (asterisk) and part of the adoral zone. Arrow marks anteriormost frontoven-
Hemiurosoma
Fig. 130h–j Hemiurosoma terricola (from Foissner et al. 2002. Protargol impregnation). h, i: Infraciliature of ventral and dorsal side and nuclear apparatus of holotype specimen, 147 µm. Arrow in (h) marks anteriormost frontoventral cirrus (= cirrus III/2). Arrows in (i) denote dorsal kinety 4, which is a dorsomarginal row. j: Infraciliature of ventral side of a very early divider, 155 µm. A Yshaped cirral anlage grows out from the oral primordium and extends to the parental buccal vertex. CC = caudal cirri, OP = oral primordium, TC = transverse cirri, 1–4 = dorsal kineties. Page 619.
b
tral cirrus (= cirrus III/2). g: Very early divider with oral primordium in anterior body half. AZM = distal (f), respectively, proximal (g) end of adoral zone of membranelles, BL = buccal lip, Cr = crystal, FC = right frontal cirrus, FG = fat globules, FT = frontoterminal cirri, FV = food vacuole, MA = macronuclear nodules, MO = mitochondria, OP = oral primordium, P = paroral, PF = pharyngeal fibres, RMR = right marginal row, IV/3 = rearmost frontoventral cirrus. Page 619.
621
622
SYSTEMATIC SECTION
Fig. 130k–m Hemiurosoma terricola (from Foissner et al. 2002. Protargol impregnation). k, l: Four cirri anlagen develop by lateral budding from the stem of the Yshaped anlage present in specimen shown in Fig. 130j and generate primary primordia. Arrow in (l) marks dikinetids migrating anteriorly. Adoral membranelles are formed in anterior portion of oral primordium. m: The primary primordia commence to split transversely and the anterior portions migrate anteriorly (arrows). Broken lines connect anterior and posterior portion of primary primordia as well as prospective proter anlagen (see Fig. 130n). OP = oral primordium, II–IV, VI = frontal-ventraltransverse cirri anlagen. Page 619.
life, scanning electron micrographs, and protargol preparations. Body outline very elongate lanceolate with anterior end usually transversely truncate and posterior portion distinctly narrowed; specimens from Namibian site (29) almost parallel-side and narrowed tail-like posteriorly, very much like H. similis (Fig. 130a–c, h, i, v, Table 40). Body slightly flattened dorsoventrally, very flexible, but acontractile. Cells difficult to preserve, usually inflated and/or wrinkled in ordinary protargol preparations (Foissner’s method), while well-preserved with Dieckmann’s technique (Fig. 130h, i). Macronuclear nodules left of midline in middle third of cell, ellipsoidal, contain many minute chromatin bodies; one or two nodules incompletely separated in about 25% of specimens, quadrinuclear pattern, however, always recognisable; nodules
Hemiurosoma
623
sometimes in two more or less distinct pairs, especially in Costa Rican and Namibian site (29) specimens; cells with five or six nodules more frequent in Costa Rican than Namibian population. Usually two ellipsoidal micronuclei, one each in anterior and posterior portion of nuclear figure, frequently not impregnated with protargol. Contractile vacuole near mid-body left of midline, during diastole with fine collecting canals. Cortical granules lacking; subcortical mitochondria as conspicuous as in Urosoma spp., about 2–3 × 1–2 µm in size. Cytoplasm colourless, contains many sand-like crystals 1–3 µm across, mainly in rear body quarter, and innumerable, about 1 µm-sized fat droplets forming short rows and reticular structures (Fig. 130d). Cells usually packed with 4–6 µm-sized food vacuoles. Movement slow, glides on microscope slide and soil particles. Resting cysts of a population from the Zambezi floodplain globular with smooth, colourless wall (Fig. 400j in Foissner et al. 2002). Adoral zone short, occupies only 14–21%, on average 18% of body length, roughly in Gonostomum pattern (see Berger & Foissner 1997, Berger 1999, and H. goertzi), composed of an average of 21 membranelles, bases of largest membranelles 5–6 µm wide in life. Buccal cavity very flat and narrow, right margin forms hyaline lip bearing paroral and covering buccal cavity and proximal third of adoral zone. Undulating membranes almost straight and in series or slightly overlapping, both minute and likely dikinetidal; paroral with a small fibre bundle anteriorly and cilia about 7 µm long in life. Pharyngeal fibres distinct in life and protargol preparations, of ordinary length and structure, extend obliquely backwards (Fig. 130a, f, h; Fig. 400a, c, f, h, i in Foissner et al. 2002; Table 40). Cirral pattern constant, number of cirri of usual variability (Fig. 130a, h, i, v; Fig. 400a–i in Foissner et al. 2002; Table 40). Frontal cirri slightly enlarged, right cirrus, as is usual, behind distal end of adoral zone. Buccal cirrus right of anterior half of paroral. Frontoventral cirri right of midline, arranged in Urosoma pattern, that is, in slightly oblique row with anterior cirrus (= cirrus III/2) somewhat dislocated to left and indistinctly enlarged. No postoral ventral cirri and pretransverse ventral cirri. Transverse cirri of about same size as marginal cirri, but 15–20 µm long, very near to body end and thus distinctly projecting. Marginal cirri about 10 µm long in life, likely composed of 3–4 × 2 basal bodies, except for rear cirri comprising only four cilia. Right marginal row commences almost at level of right frontal cirrus, terminates, like left row, slightly ahead of or about at level of transverse cirri; left row begins left of rear end of adoral zone. Dorsal bristles 3–4 µm long in life, much more closely spaced in anterior than posterior half of cell, arranged in four kineties; kinety 1 slightly shortened anteriorly, kineties 2 and 3 bipolar; kinety 4, a dorsomarginal row, consists of only 3–4 bristles in anterior body third. One caudal cirrus each at rear end of kineties 1–3, 15–20 µm long, near posterior body end, and thus difficult to distinguish from transverse cirri (Fig. 130i; Fig. 400e in Foissner et al. 2002). Cell division (Fig. 130g, j–u): This part of the life cycle was studied by Foissner et al. (2002) in specimens from the type locality (Namibia) and from Costa Rica
624
SYSTEMATIC SECTION
Fig. 130n–q Hemiurosoma terricola (from Foissner et al. 2002. Protargol impregnation). n, o: Infraciliature of early dividers, n = 164 µm, o = 138 µm. The anterior portion of the primary primordia II and III migrate to the proter, where they unite with anlagen formed by the parental buccal cirrus and the anteriormost frontoventral cirrus (= cirrus III/2; a broken line connects the individual parts of an anlage). Thus, five anlagen are recognisable in proter and opisthe. Arrows in (o) mark right marginal row anlagen. p, q:
Hemiurosoma
625
(Fig. 130n, u). Ontogenesis of H. terricola is similar to that of Urosoma because the frontal-ventral-transverse cirri of the proter are entirely or at least partially produced by the opisthe via long primary primordia (term introduced by Foissner 1983a), a highly characteristic feature uniting Urosoma and Gonostomum according to Berger & Foissner (1997) and Berger (1999, p. 68). From the interphasic cirral pattern and the cell division one can conclude that frontal-ventral-transverse cirri anlage V is lost in H. terricola. Consequently, I designate the five remaining anlagen as I–IV and VI, and not I–V as in the original description, that is, anlage V in Foissner et al. (2002) corresponds anlage VI in present review. Cell division begins with the formation of a long, narrow oral primordium extending anteriorly from mid-body (Fig. 130g). Subsequently, two cirral anlagen arise from the anterior end of the oral primordium and extend to the parental buccal vertex in characteristic Y-pattern (Fig. 130j). Next, two further anlagen develop from the stem of the Y. Thus, four more or less long cirral anlagen are recognisable in this stage (Fig. 130k, l). These anlagen are primary primordia, which split transversely, producing a cirral anlagen set each in both filial products (Fig. 130m, n): the posterior half of the leftmost anlage becomes the opisthe’s anlage I which forms the undulating membranes and the left frontal cirrus; in addition, this streak gives rise to anlage II of the opisthe; the anterior portion unites with proter’s anlage II formed by the parental buccal cirrus; anlage I of the proter originates, as is usual, from the reorganising parental undulating membranes (Fig. 130o, p, r); the anterior portion of the primary primordium III unites with the anlage III of the proter produced by the disorganised cirrus III/2; the anlagen IV and VI of the proter are the anterior portion of the primary primordia IV and VI (Fig. 130n, o), because the other parental frontoventral cirri (IV/3, VI/3, VI/4) are ontogenetically inactive and will be resorbed in very late dividers (Fig. 130r, s, u). When the macronuclear nodules have formed a single, globular mass and micronuclear division commences, five widely separated cirral anlagen and three dorsal primordia are recognisable in both the proter and the opisthe. Furthermore, the new adoral zone is almost complete (Fig. 130p, r). Subsequently, cirri begin to segregate in the anlagen and the last cirrus of anlagen IV and VI migrate posteriorly to form the two transverse cirri usually present in this species (Table 40). In late dividers, the undulating membranes are completed, the cirri migrate to their specific sites, and supernumerary and parental cirri are resorbed. Caudal cirri originate at the end of the new kineties 1–3 and the globular macronuclear mass divides twice to produce the species-specific four nodules (Fig. 130s–u). In addition, a very short dorsomarginal kinety (= dorsal kinety 4) develops at/near anterior end of right marginal row anlage (Fig. 130s).
b
Infraciliature of ventral and dorsal side and nuclear apparatus of a middle divider, 140 µm. Cirri form within anlagen and two primordia develop within each bipolar dorsal kinety. Arrow in (p) marks opisthes’s left frontal cirrus (= cirrus I/1) which forms at the anterior end of the undulating membrane anlage. MA = condensed macronucleus, MI = micronuclei, I–IV, VI = frontal-ventral-transverse cirri anlagen. Page 619.
626
SYSTEMATIC SECTION
Fig. 130r–t Hemiurosoma terricola (from Foissner et al. 2002. Protargol impregnation. Parental structures white, new black). r: Infraciliature of ventral side of late divider, 141 µm. s, t: Infraciliature of ventral and dorsal side and nuclear apparatus of very late divider, 160 µm. Arrows in (t) mark caudal cirri of opisthe. CC = parental caudal cirri, MA = macronuclear nodule, MI = micronucleus, TC = new transverse cirri, I, IV, VI = frontal-ventral-transverse cirri anlagen, 1–4 = dorsal kineties (kinety 4 is a dorsomarginal row). Page 619.
Hemiurosoma
Fig. 130u, v Hemisincirra terricola (from Foissner et al. 2002. Protargol impregnation). u: Infraciliature of ventral side and nuclear apparatus of a late divider, 158 µm. Note that in the opisthe distinctly more cirri have been formed than usually present in interphasic specimens (parental structures white, new black). v: Infraciliature of ventral side and nuclear apparatus of a paratype specimen. Arrow marks cirrus III/2. FC = right frontal cirrus (= cirrus III/3), MA = macronuclear nodule, TC = transverse cirri transverse cirri (of opisthe in u). Page 619.
627
628
SYSTEMATIC SECTION
Molecular data: The complete 18S rRNA gene sequence of Hemiurosoma terricola is 1773 nucleotides long (GenBank accession number AY498651; Foissner et al. 2004). The present species is included in phylogenetic analyses by Gong et al. (2006, p. 71), Foissner & Stoeck (2006), and Schmidt et al. (2007). For a discussion about the phylogenetic position of the type species of Hemiurosoma, see remarks in genus section. Occurrence and ecology: Hemiurosoma terricola is very likely confined to terrestrial habitats (Foissner et al. 2002, p. 53). Type locality is the Etosha National Park (19°S, 15°40'E), Namibia, where we discovered it in soil from the ghost tree forest (Moringa ovalifolia; site 56 in Foissner et al. 2002). It occurred in several other sites in Namibia, but also (i) in a desert soil from Arizona, USA (sample collected by Klaus Hausmann, Berlin); (ii) in a soil sample from Costa Rica, namely a horse pasture near peak of Monte Verde (pH 5.5; originally a fog rain forest; Fig. 130n, u); and (iii) in a sample from the Zambezi River floodplain in Botswana, indicating a broad ecological (mud of ephemeral polls to true soils, inter alia, saline ones) and geographic range (Foissner et al. 2002, p. 842). Species of the Zambezi population could be cultivated in Eau de Volvic enriched with some wheat grains, and the scanning electron micrographs resulting from this population match the description based on live observations and protargol preparations (Fig. 400a–j in Foissner et al. 2002). Hemiurosoma terricola is well adapted to the terrestrial habitat by its long and slender body. Feeds on bacteria and, likely, heterotrophic flagellates (Foissner et al. 2002).
Hemiurosoma goertzi Foissner, Agatha & Berger, 2002 (Fig. 131a–m, Table 40) 2002 Hemiurosoma goertzi nov. spec.1 – Foissner, Agatha & Berger, Denisia, 5: 843, Fig. 183a–i, 381a–d, Table 161 (Fig. 131a–m; original description; the holotype slide [accession number 2002/102] and four paratype slides [2002/103–106] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; see Foissner et al. 2002, p. 39 and Aescht 2003, p. 387).
Nomenclature: We dedicated this new species to Hans-Dieter Görtz, Stuttgart University (Germany), for his scientific excellence and editorial activities. Remarks: The present species differs from H. polynucleatum, the sole congener with one postoral ventral cirrus, mainly by the macronuclear pattern (2 vs. 8 nodules in line), an excellent feature because of its high stability. Minor differences occur in the number (3 vs. 2) and location (subterminal vs. terminal) of the transverse cirri and the number of adoral membranelles (25–30 vs. 19–24). In life, Hemiurosoma goertzi is characterised by the following combination of features (Foissner et al.
1 Foissner et al. (2002) provided the following diagnosis: Size about 150 × 20 µm in vivo, under certain circumstances about 250 × 40 µm. Slenderly lanceolate with short tail. 2 macronuclear nodules, a single postoral cirrus, and 27 adoral membranelles on average.
Hemiurosoma
629
Fig. 131a–f Hemiurosoma goertzi (from Foissner et al. 2002. a–d, from life; e, f, protargol impregnation). a: Ventral view of a representative specimen, 145 µm. b, c: Ventral and right lateral view of shape variant showing, inter alia, contractile vacuole with collecting canals. d: Mitochondria are 2–3 µm long and form a conspicuous layer under the cortex. This feature separates H. goertzi from similar species, for instance Erimophrya spp. e, f: Infraciliature of ventral and dorsal side and nuclear apparatus of paratype specimen, 120 µm. Arrow marks postoral ventral cirrus IV/2. For a more detailed labelling see Fig. 131g, h. AZM = adoral zone of membranelles, BL = buccal lip, CC = caudal cirri, MA = macronuclear nodule, MI = micronucleus, MO = mitochondria, P = paroral, TC = transverse cirri (possibly, the smallest cirrus is a pretransverse ventral cirrus), 1–4 = dorsal kineties. Page 628.
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SYSTEMATIC SECTION
Fig. 131g–i Hemiurosoma goertzi (from Foissner et al. 2002. Protargol impregnation). Large specimens from flooded cultures (details see first paragraph of morphology chapter). g: Detail of marginal and transverse cirri. h: Infraciliature of oral region showing fine structure of cirri and adoral membranelles. Cirri III/2 and IV/3 as well as frontoterminal cirri (FT) form the frontoventral cirri which are arranged in the so-called Urosoma pattern, that is, all cirri are in line except of cirrus III/2, which is slightly displaced leftwards. i: Infraciliature of ventral side and nuclear apparatus of holotype specimen, 231 µm. AZM = distal end of adoral zone of membranelles, BC = buccal cirrus, E = endoral, FC = right frontal cirrus (= cirrus III/3), FT = frontoterminal cirri (= cirri VI/3, VI/4), LMR = left marginal row, MA = macronuclear nodule, MI = micronucleus, P = paroral, PF = pharyngeal fibres, RMR = right marginal row, TC = transverse cirri (possibly one [or two] of them is actually a pretransverse ventral cirrus; otherwise all anlagen [II, III, IV, VI] must form a transverse cirrus). Page 628.
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2002): slender, slightly tailed body with anterior end transversely truncate, two macronuclear nodules, single postoral ventral cirrus, and distinct mitochondria. Morphology: Hemiurosoma goertzi showed an extraordinary variability of body size (Foissner et al. 2002). When it was studied and prepared from an about 10 days old, ordinary non-flooded Petri dish culture, it measured about 150 × 20 µm. After three weeks, the Petri dish was flooded causing strong reproduction and increase in size to about 250 × 40 µm within two weeks (Fig. 130g–i). Interestingly, all other main features, for example, the number of adoral membranelles and marginal cirri as well as the size of the macronuclear nodules, remained stable (Fig. 130i, Table 40). A similar size variability has been reported for Urosoma caudata (Ehrenberg, 1833) Berger, 1999, which was considerably smaller in protargol slides from terrestrial (Foissner 1982) than limnetic material (Foissner 1984; named U. cienkowskii in Foissner’s papers; for review, see Berger 1999, p. 398). Thus, we confined the description to material first obtained from the ordinary Petri dish culture (Foissner et al. 2002). Body size 115–180 × 15–30 µm, on average about 150 × 20 µm in life; body length:width ratio about 7:1 (range 6–8:1) in life, on average 6:1 in protargol preparations (Table 40). Body outline elongate lanceolate with anterior end usually transversely truncate and posterior end tail-like and curved to right. Body flattened about 2:1 dorsoventrally, very flexible, but acontractile. Cells very fragile and thus difficult to preserve, tail often inflated in silver preparations (Fig. 131a, b, e, i, l, m). Macronuclear nodules slightly left of midline, about 17 × 8 µm in life, ellipsoidal, with many medium-sized chromatin bodies. Micronuclei about 6 × 3 µm in life, ellipsoidal, one usually attached to front end of anterior macronuclear nodule, the other to rear end of posterior nodule. Contractile vacuole above mid-body at left cell margin, during diastole with distinct collecting canals (Fig. 131a, f, i, j, m). Cortical granules lacking; subcortical mitochondria, however, as conspicuous as in most Urosoma-like hypotrichs. Cytoplasm colourless, contains some ordinary crystals mainly in posterior third and food vacuoles 4–11 µm across. Glides rather rapidly on microscope slide and soil particles. Adoral zone occupies 17–26%, on average 22% of body length, roughly in Gonostomum pattern, that is, extends straight along left body margin, performing right bend and slight clockwise rotation to plunge into buccal cavity; composed of an average of 27 membranelles, bases of largest membranelles about 4 µm wide in life. Buccal cavity very flat and narrow, right margin forms hyaline lip bearing paroral and covering buccal cavity and proximal adoral membranelles. Undulating membranes almost straight and side by side, endoral on average 10 µm long, paroral 8 µm long and commencing about 3 µm ahead of endoral at level of buccal cirrus; both membranes likely composed of dikinetids. Pharyngeal fibres distinct in life and protargol preparations, of ordinary length and structure, extend obliquely backwards (Fig. 131a, b, e, h, j, k, Table 40). Cirral pattern and number of cirri of usual variability (Fig. 131a, e, g–m, Table 40). Frontal cirri frequently slightly enlarged, right one behind distal end of adoral
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SYSTEMATIC SECTION
Fig. 131j–m Hemiurosoma goertzi (from Foissner et al. 2002. Protargol impregnation). These specimens from the flooded culture are more than 200 µm long. j, k: Ventral view of anterior portion. Arrowhead marks cirrus III/2, arrow denotes single postoral ventral cirrus. l, m: Overviews showing the slender body outline and the subterminal location of the transverse cirri. Explanation of original labelling: AZM = adoral zone of membranelles, BU = buccal cirrus, EM = endoral, LMR = left marginal row, MA = macronuclear nodule, PF = pharyngeal fibres, PM = paroral, RMR = right marginal row, TC = transverse cirri. Page 628.
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zone. Buccal cirrus right of anterior portion of paroral, of variable size, that is, composed of 4 × 3, 4 × 2, or fewer basal bodies. Frontoventral cirri in typical Urosoma pattern, that is, cirri IV/3, VI/3, and VI/4 longitudinally arranged and anterior cirrus (= cirrus III/2) somewhat shifted leftwards and slightly enlarged (Fig. 131h, j; for details, see Berger 1999, p. 66 and Fig. 19a). Postoral ventral cirrus at about 24% of body length and slightly left of midline, composed of 3 × 2, 4 × 2, or 3 × 3 basal bodies. Transverse cirri about 20 µm long in life and frequently slightly larger than marginal cirri, subterminal and thus not projecting posteriorly; usually arranged in triangular pattern, so that one cannot exclude that one cirrus is a pretransverse ventral cirrus. Right marginal row usually commences right of area between right frontal cirrus and cirrus III/2, terminates, like left row, slightly ahead of rear body end. Left marginal row commences left of proximal end of adoral zone. Marginal cirri about 12 µm long in life, composed of 4 × 2 basal bodies, except for posterior cirri comprising only four cilia. Dorsal bristles about 3 µm long in life, arranged in four kineties (Table 40); kineties 1–3 about of body length, each with a single caudal cirrus; kinety 4 lacks a caudal cirrus and is slightly shortened anteriorly and distinctly so posteriorly because it is (very likely) a dorsomarginal row (Fig. 131f). Occurrence and ecology: Probably confined to terrestrial habitats (Foissner et al. 2002, p. 53). Type locality of H. goertzi is a highly saline soil from the Sporobolus zone around the Etosha Pan (19°10'S, 15°55'E), Namibia (Foissner et al. 2002). We found it also in three other sites from Namibia, ranging from non-saline mud and soil from rock-pools on an Inselberg to the highly saline soil of the type locality. Feeds on bacteria, golden-coloured fungal conidia (11 × 6 µm), and, rarely, small ciliates (Foissner et al. 2002).
Hemiurosoma similis (Foissner, 1982) Foissner, Agatha & Berger, 2002 (Fig. 132b) 1982 Perisincirra similis nov. spec. – Foissner, Arch. Protistenk., 126: 94, Abb. 24a–f, 66, 72, Tabelle 21 (Fig. 132b; original description; the holotype slide [accession number 1981/92] is deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; Aescht 2003, p. 396). 1984 Hemisincirra similis (Foissner, 1982) nov. comb. – Foissner, Stapfia, 12: 119 (combination with Hemisincirra, see nomenclature). 1999 Urosoma similis (Foissner, 1982) comb. nov. – Berger, Monographiae biol., 78: 397, 419, Fig. 131a–g, Table 24 (Fig. 132b; combination with Urosoma and detailed review). 2001 Urosoma similis (Foissner, 1982) Berger, 1999 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 72 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Hemiurosoma similis (Foissner, 1982) nov. comb. – Foissner, Agatha & Berger, Denisia, 5: 835, Fig. 183k (Fig. 132b; combination with Hemiurosoma).
Nomenclature: No derivation of the name is given in the original description and the review by Berger (1999). The species-group name simil·is, -is, -e (Latin adjec-
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SYSTEMATIC SECTION
tive [m; f; n]; similar, identical) alludes to the similarity with the congeners (Perisincirra spp. sensu Foissner 1982), Urosoma, and Urosomoida species. Foissner (1984) transferred it to Hemisincirra before this genus was published by Hemberger (1985; see p. 388). In the Catalogue of ciliates names (Berger 2001) I did not correct this obviously invalid act because this combination was – due to the transfer to Urosoma (Berger 1999) – no longer relevant. Remarks: Foissner (1982) established this species in Perisincirra Jankowski, 1978; later, he transferred it to Hemisincirra (Foissner 1984), a heterogeneous group preliminarily classified in the amphisiellids (p. 387). In 1999, I transferred the species Urosoma (further details, see remarks at Hemiurosoma polynucleatum). For detailed review, see Berger (1999, p. 419).
Hemiurosoma polynucleatum (Foissner, 1984) Foissner, Agatha & Berger, 2002 (Fig. 132a) 1984 Hemisincirra polynucleata nov. spec. – Foissner, Stapfia, 12: 119, Abb. 63a–h, Tabelle 30 (Fig. 132a; original description; according to Aescht 2003, p. 394, two slides with syntypes [accession numbers 1984/95, 96] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria). 1999 Urosoma polynucleata (Foissner, 1984) comb. nov. – Berger, Monographiae biol., 78: 397, 419, Fig. 130a–i, Table 24 (Fig. 132a; combination with Urosoma and detailed review). 2001 Urosoma polynucleata (Foissner, 1984) Berger, 1999 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 30 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Hemiurosoma polynucleata (Foissner, 1984) nov. comb. – Foissner, Agatha & Berger, Denisia, 5: 835, Fig. 183j, Table 153 (Fig. 132a; combination with Hemiurosoma).
Nomenclature: No derivation of the name is given in the original description and the review by Berger (1999). The species-group name polynucleatum (having many macronuclear nodules) is a composite of the Greek quantifier polys and the Latin adjective nucleat·us, -a, -um (m; f; n; [nut]nucleus-like) and refers to the increased number (usually eight) of macronuclear nodules. Hemisincirra is of feminine gender. By contrast, Urosoma and Hemiurosoma are neuter (ICZN 1999, Article 30.1.2) so that the species-group name polynucleatum has to be used in combination with Urosoma or Hemiurosoma (nomen corrigendum). Remarks: Foissner (1984) established this species in Hemisincirra, a still insufficiently defined group including mainly very slender soil species that do not fit any other genus well (p. 387). I transferred Hemisincirra polynucleata to Urosoma because of the oral apparatus, the Urosoma-like arrangement of the frontoventral cirri, the dorsal kinety pattern (mainly because of the dorsomarginal kinety), and the distinct mitochondria underneath the pellicle (Berger 1999). I already stated that this species and U. similis very likely form a distinct clade in Urosoma because of the reduced number of frontal-ventral-transverse cirri behind the buccal vertex (Berger 1999, p. 397, 419). Somewhat later, we therefore transferred both species to Hemi-
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urosoma (Foissner et al. 2002; see remarks at genus section). For review of H. polynucleatum, see Berger (1999, p. 419).
Fig. 132a Hemiurosoma polynucleatum (from Foissner 1984. Protargol impregnation). Infraciliature of ventral side and nuclear apparatus, 145 µm. Arrow marks postoral ventral cirrus. Page 634. Fig. 132b Hemiurosoma similis (from Foissner 1982. Protargol impregnation). Infraciliature of ventral side and nuclear apparatus, 120 µm. Arrow marks postoral ventral cirrus. Page 633.
Note to nomenclature of Hemiurosoma (p. 614; added at proof-reading): Foissner et al. (2002) wrote that Hemiurosoma is of feminine gender. However, according to Article 30.1.2 of the ICZN (1999), a genus-group name that is or ends in a Greek word transliterated into Latin without other changes takes the gender given for that word in standard Greek dictionaries. And according to the examples presented in the ICZN (1999, p. 35), the word -soma is of neuter gender.
Supplement to the Urostyloidea The revision of Hemisincirra – a heterogeneous group of uncertain phylogenetic position provisionally classified as incertae sedis in the amphisiellids in the present review (p. 387) – showed, that the two species reviewed below have a more or less distinct midventral complex originating from more than the ordinary six (I–VI) frontal-ventral-transverse cirri anlagen (has to be confirmed for A. verrucosa). Thus, they are transferred from Hemisincirra to Anteholosticha, the corresponding urostyloid genus characterised by three frontal cirri and a midventral complex composed of cirral pairs only (for review and terminology of the urostyloids, see Berger 2006). In addition, the supplemented key to Anteholosticha species is provided. Erratum: In the monograph of urostyloids, the designation of the frontal cirri in Pseudourostyla nova is incorrect (Fig. 153b in Berger 2006): cirrus II/1 (false) = cirrus II/3 (correct) and III/1 = III/3. Possibly, the same mistake was also made in a few other cases.
Supplemented Key to Anteholosticha species If you know that your specimen/population belongs to the urostyloid genus Anteholosticha, identification is still rather difficult because many species are not described in detail. Key features are the nuclear apparatus, body size, cortical granulation, ventral and dorsal ciliature, and habitat. Thus, very detailed life observation and protargol impregnation is needed for a reliable identification of many species. Also check the Caudiholosticha key if you are uncertain about the presence or absence of caudal cirri (Berger 2006, p. 234). Note: All references to figures and pages in the following key refer to the Monograph of the Urostyloidea (Berger 2006) unless otherwise indicated! 1 2 3
Two macronuclear nodules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 More than two macronuclear nodules. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Single micronucleus between macronuclear nodules (Fig. 51a, 69a, 82a, 83a). . 3 Different nucleus pattern (e.g., Fig. 96a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Transverse cirri displaced very far anteriorly, that is, inserted at end of second body third (Fig. 51a; caudal cirri likely present). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caudiholosticha navicularum (p. 274) - Transverse cirri displaced not so far anteriorly. . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4 About 10 enlarged frontal cirri (Fig. 83a). . . . . . Anteholosticha alpestris (p. 403) - Three enlarged frontal cirri (Fig. 69a, 82a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5 Body length:width ratio about 2:1; midventral complex extends to near transverse cirri; limnetic (Fig. 69a). . . . . . . . . . . . . . . . . Anteholosticha brevis (p. 360) 636
Anteholosticha 6 7 8 9 10 11
12 13 14 15
16 17
637
Body length:width ratio about 3:1; midventral complex terminates slightly behind mid-body; terrestrial (Fig. 82a). . . . . . . . . Anteholosticha muscicola (p. 401) (2) One conspicuous ring-shaped structure each in anterior and posterior body portion; marine (Fig. 95a, e). . . . . . . . . . . . . . . . . Anteholosticha oculata (p. 448) No such structures in this arrangement; marine or limnetic. . . . . . . . . . . . . . . . . 7 Ectoplasm with alveolar seam; marine (Fig. 96a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha arenciola (p. 452) Alveolar seam lacking; limnetic or slightly saline waters. . . . . . . . . . . . . . . . . . . 8 Body length below 100 µm (Fig. 72a). . . . . . Anteholosticha vuxgracilis (p. 369) Body length more than 100 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Body length:width ratio around 6–8:1; adoral zone occupies about 30% of body length (Fig. 67a). . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha violacea (p. 342) Body length:width ratio around 2–3:1; adoral zone occupies about 40% or more of body length (Fig. 70a, 71a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 About 11 midventral cirral pairs; 9–15 transverse cirri (Fig. 70a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha camerounensis (p. 361) About 5 (very loosely arranged) midventral cirral pairs; 4–6 transverse cirri (Fig. 71a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha macrostoma (p. 365) (1) Four (rarely 3) longitudinally arranged pairs of macronuclear nodules, each with a single micronucleus in between; dorsal bristles about 10 µm long (Fig. 92a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha extensa (p. 438) No such nuclear pattern; dorsal bristles distinctly shorter (in A. violacea bristles are about 8 µm long). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Many small macronuclear nodules more or less scattered throughout cytoplasm (e.g., Fig. 61j). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 About 4 to about 20 macronuclear nodules, often in more or less distinct row in left body portion (e.g., Fig. 57c). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 (41) Cytoplasm diffuse yellow (Fig. 68a). . Anteholosticha xanthichroma (p. 345) Cytoplasm not distinctly yellow. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Cortical granules (e.g., Fig. 78b) or extrusomes (e.g., Fig. 55q, r) present. . . . 15 Cortical granules or extrusomes lacking. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 Cortical granules/extrusomes colourless, rod-shaped (about 2.0–3.0 × 1.0–1.5 µm), form (indistinct) seam, ejected when methyl-green pyronin is added (Fig. 55w, x, 75b–e). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 Cortical granules more or less globular to slightly ellipsoidal, colourless or coloured (e.g., Fig. 77h, i, m). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 Seven to about 12 transverse cirri; usually 6 dorsal kineties; mainly limnetic (Fig. 55b, t, v). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha monilata (p. 297) 3–6 transverse cirri; 4 dorsal kineties; terrestrial (Fig. 75a–g). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha australis (p. 382) (15) Midventral complex extends beyond mid-body; cortical granules in longitudinal rows, colourless; on average 20 or more adoral membranelles (Fig. 77a–m). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha sigmoidea (p. 387)
638 -
SYSTEMATIC SECTION
Midventral complex terminates ahead of mid-body; cortical granules around dorsal bristles, colourless, yellowish, orange, or pink; on average about 15 adoral membranelles (e.g., Fig. 78a–e). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 18 (42) Cortical granules pink or orange, globular; right marginal row commences about at level of proximal end of adoral zone (Fig. 78a–j). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha bergeri (p. 393) - Cortical granules yellowish, ellipsoidal; right marginal rows commences about at level of buccal cirrus (number of macronuclear nodules 28–38; Fig. 80a–g). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha brachysticha (p. 397) 19 (14) Three dorsal kineties (Fig. 64a–c). . . . . . Anteholosticha mancoidea (p. 336) - 4 dorsal kineties (Fig. 65a, 66a, 76a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 20 Four distal adoral membranelles distinctly set off from remaining part of adoral zone (Fig. 65a). . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha randani (p. 338) - Adoral zone of membranelles continuous (Fig. 66a, 76a; separation of following 2 species difficult). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 21 About 16 macronuclear nodules; 30–33 adoral membranelles; midventral complex terminates at about 80% of body length; usually two enlarged transverse cirri; terrestrial (Fig. 76a). . . . . . . . . . . . . . . . . . . . Anteholosticha distyla (p. 385) - 8–11 macronuclear nodules; 24–25 adoral membranelles; midventral complex terminates at about 60% of body length; usually 5 transverse cirri; limnetic (Fig. 66a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha sphagni (p. 340) 22 (12) Cortical granules red (dark-red, yellow-reddish). . . . . . . . . . . . . . . . . . . . . 23 - Cortical granules not red or lacking. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 23 Body length 200–300 µm; dark-red cortical granules around cirri and dorsal bristles (Fig. 90a, b). . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha pulchra (p. 433) - Body length around 120 µm; yellowish-red granules arranged in rows (Fig. 88a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha gracilis (p. 426) 24 (22) Cytoplasm diffuse yellow (colour is not due to yellow cortical granules!) (e.g., Fig. 68a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24a - Cytoplasm not distinctly yellow (yellow cortical granules are present in some species!). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 24a Limnetic; buccal cirri present; body not distinctly tailed (Fig. 68a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha xanthichroma (p. 345) - Saltwater; buccal cirri lacking; posterior body end narrowed tail-like (Fig. 97a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha azerbaijanica (p. 454) 25 (24) More than 1 buccal cirrus present (e.g., Fig. 74a, b). . . . . . . . . . . . . . . . . . 26 - Buccal cirrus present or lacking (e.g., Fig. 85a, g). . . . . . . . . . . . . . . . . . . . . . . 30 26 More than 3 frontal cirri (Fig. 84a–c). . . . . . . Anteholosticha multistilata (p. 405) - 3 enlarged frontal cirri (e.g., Fig. 61a, f, i). . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 27 Marine (salt water) (Fig. 86a). . . . . . . . . . . . . . . . Anteholosticha estuarii (p. 420) - Limnetic or terrestrial. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 28 Body length above 200 µm in life; transverse cirri hardly project beyond rear body end (Fig. 73a–h). . . . . . . . . . . . . . . . . . . Anteholosticha antecirrata (p. 370)
Anteholosticha -
639
Body length below 200 µm in life; transverse cirri project distinctly beyond rear body end (e.g., Fig. 61f). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 29 Marginal rows overlapping posteriorly (Fig. 61i, l, n). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha intermedia (p. 317) - Marginal rows distinctly separated posteriorly (Fig. 74b, g, i). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha adami (p. 376) 30 (25) Buccal cirrus lacking (Fig. 97a). . . . . Anteholosticha azerbaijanica (p. 454) - Buccal cirrus present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30a 30a Terrestrial. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 - Limnetic or marine (saltwater). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 31 Cortical granules around dorsal bristles; about 28–38 macronuclear nodules (Fig. 80a–g). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha brachysticha (p. 397) - Cortical granules lacking (Fig. 81a). . . . . . Anteholosticha plurinucleata (p. 399) 32 (30a) Limnetic (see also marine, if your sample site is slightly brackish). . . . . 33 - Marine (saltwater; see also limnetic, if the sample site is brackish). . . . . . . . . . 36 33 Four distalmost adoral membranelles distinctly set off from remaining membranelles; 16–20 macronuclear nodules (Fig. 65a) . Anteholosticha randani (p. 338) - Adoral zone continuous; more than 20 macronuclear nodules . . . . . . . . . . . . . . 34 34 Body length 140–250 µm; body rather slender (Fig. 62a, 67b). . . . . . . . . . . . . 35 - Body length 100–140 µm; body elliptical (Fig. 63a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha thononensis (p. 334) 35 Body 140–180 µm long; dorsal bristles about 3 µm long (Fig. 62a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha grisea (p. 332) - Body 180–250 µm long; dorsal bristles about 8 µm long (Fig. 67b). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha violacea (p. 342) 36 (32) Body length 200–300 µm; body very slender (length:width ratio about 10:1; Fig. 93a, b). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha fasciola (p. 441) - Body length below 200 µm; body length:width ratio distinctly below 10:1. . . . 37 37 Cortical granules yellow-greenish or yellow-red (Fig. 88a, e). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha gracilis (p. 426) - Cortical granules(?) colourless or lacking (note: for A. longissima the presence or absence of cortical granules is not known). . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 38 Cortical granules shaped like mammalian erythrocytes (Fig. 85a–g). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha warreni (p. 412) - Cortical granules of other shape. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 39 Midventral complex extends to near transverse cirri (Fig 91a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha longissima (p. 437) - Midventral complex terminates at about 2/3 of body length (e.g., Fig. 87a, 94f)40 40 Body length about 100–130 µm; about 5 transverse cirri (Fig. 87a–g). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha manca (p. 422) - Body length below 100 µm (around 50–80 µm); about 7–8 transverse cirri (Fig. 94f). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha scutellum (p. 443)
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41 (12) Two dorsal kineties1 (Fig. 133a, b, r, s in present book). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha heterocirrata (p. 640 in present book) - Three or more dorsal kineties. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 42 (17) Three dorsal kineties (Fig. 78i). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 - Four dorsal kineties (Fig. 134a–k in present book). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anteholosticha verrucosa (p. 647 in present book)
Supplement to Anteholosticha The following two species are transferred from Hemisincirra to Anteholosticha because they show a more or less distinct midventral complex.
Anteholosticha heterocirrata (Hemberger, 1985) comb. nov. (Fig. 133a–s, Table 26) 1982 Perisincirra heterocirrata n. spec.2 – Hemberger, Dissertation3, p. 207, Abb. 36a–k (Fig. 133c–s; description of morphology and morphogenesis). 1985 Hemisincirra heterocirrata n. spec. – Hemberger, Arch. Protistenk., 130: 408, Abb. 14 (Fig. 133a, b; original description; no formal diagnosis provided; type slides are deposited in the Institut für landwirtschaftliche Zoologie und Bienenkunde, University of Bonn, Germany). 2001 Hemisincirra heterocirrata Hemberger, 1985 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 30 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs).
Nomenclature: The species-group name heterocirrat·us, -a, -um (adjective [m; f; n]; having different cirri) is a composite of the Greek word heteros (a different one; different, in compositions), the Latin noun cirr·us (curl; cirrus in present case), the suffix -at (provided with a certain feature or organ), and the inflectional ending -a. It alludes to the feature that the marginal cirri of the anterior body half are stronger than those of the posterior portion (Hemberger 1985). Since both Hemisincirra and Fig. 133a–h Anteholosticha heterocirrata (a, b, from Hemberger 1985; c–h, from Hemberger 1982. a–h, infraciliature of ventral side and nuclear apparatus after protargol impregnation; body outline of (a) from life). a, b: Interphasic specimen, 120 µm. Cirri which originate from same anlage connected by broken line. Arrow marks cirrus whose origin is uncertain. c, d: Very early divider, 120 µm. Arrow marks oral primordium which originates at the rearmost cirrus of the midventral complex (see, however, explanation of arrow in (a)). e, f: Early divider, size not indicated. g, h: Middle divider, size not indicated. Arrow marks the two rightmost anlagen, which are likely primary primordia. AZM = adoral zone of membranelles, CV = contractile vacuole, LMR = left marginal row, MI = micronucleus, OP = oral primordium, RE = replication band, RMR = anterior end of right marginal row, TC = transverse cirri, I, III, V, VII = frontal-ventral-transverse cirral anlagen. Page 640. 1
The presence/absence of cortical granules is not known for A. heterocirrata; thus, the number of dorsal kineties has to be used rather early to determine this species. 2 This name is disclaimed for nomenclatural purposes (ICZN 1999, Article 8.3). 3 See same footnote at Uroleptoides binucleatus.
d
Anteholosticha
641
642
SYSTEMATIC SECTION
Anteholosticha
643
Anteholosticha are feminine, the ending of the species-group name does not change. Remarks: Hemisincirra buitkampi, type of Hemisincirra, has a short amphisiellid median cirral row composed of only four cirri, plus one cirrus left of it. This strongly indicates that in total only five or six frontal-ventral-transverse cirri anlagen are present. Of course, this assumption has to be confirmed by ontogenetic data. By contrast, in the present species the frontal-ventral-transverse cirri are formed from seven, and not from the ordinary six anlagen. Furthermore, the anlagen IV–VII form cirral pairs so that the cirri of the frontoventral row are arranged in a more or less distinct zigzag pattern, that is, they form a midventral complex (Fig. 133a, j, p). Dorsomarginal kineties and a dorsal kinety fragmentation are lacking (Fig. 133n, p, r, s). All these features indicate that H. heterocirrata belongs to the Urostyloidea (for review, see Berger 2006). Probably it is a member of Anteholosticha Berger, 2003 whose midventral complex is, however, usually distinctly longer. In spite of that, it is preliminarily assigned to this, likely non-monophyletic genus (for review, see Berger 2006, p. 292). Periholosticha-species are also similar, but have caudal cirri and lack a buccal cirrus (for review, see Berger 2006, p. 498). Redescription (mainly investigation of live data like presence/absence of cortical granulation) recommended. Morphology: Body size (in life?) 100–140 × 15–25 µm; body length:width ratio 5–6:1. Body outline elongate elliptical to almost parallel-sided; anterior and posterior portion somewhat narrowed (Fig. 133a). Body flexible. 12–14 macronuclear nodules; position of nuclear apparatus not mentioned, likely, as is usual, in central body portion slightly left of midline. 2–3 micronuclei attached to macronuclear figure (Fig. 133b). Contractile vacuole at left cell margin about in mid-body; in specimen illustrated about at 45% of body length (Fig. 133a). Presence/absence of cortical granules, details of cytoplasm (inclusions, colour), and movement not described. Adoral zone occupies about 25% of body length, composed of about 15 membranelles of ordinary fine structure. Undulating membranes optically not intersecting in specimen illustrated; rear membrane (paroral?) slightly curved, anterior membrane (endoral?) almost straight (Fig. 133a). Cirral pattern as shown in Fig. 133a. Frontal cirri slightly enlarged, arranged in somewhat oblique pseudorow with middle cirrus behind distal end of adoral zone. Buccal cirrus small (two basal bodies only?), ahead of anterior end of rear undulating membrane (paroral?). One cirrus (= cirrus III/2) behind right frontal cirrus. Midventral complex composed of three cirral pairs arranged in distinct zigzag pattern (see cell division); complex terminates at 26% of body length in specimen illustrated and thus only slightly longer than adoral zone. Two frontoterminal cirri (= anteriormost two cirri of anlage VII; Fig. 133a). Postoral ventral cirri lacking. Two (rarely
b
Fig. 133i–o Anteholosticha heterocirrata (from Hemberger 1982. Infraciliature of ventral side and nuclear apparatus after protargol impregnation). Middle, late, and very late dividers, sizes not indicated. Arrow in (j) marks left marginal row primordium for proter. In (n) the frontoterminal cirri of the proter and the transverse cirri of the opisthe are circled. I, VII = first (leftmost) and last (rightmost) frontal-ventraltransverse cirri primordium. Page 640.
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Fig. 133p–s Anteholosticha heterocirrata (from Hemberger 1982. Protargol impregnation). p, q: Infraciliature of ventral side and nuclear apparatus of very later divider, size not indicated. Cirri which originate from the same anlage are connected by a broken line. One of the two anlagen marked by an arrow is an additional anlage so that the present species has not the ordinary six anlagen, but seven. Anlage VII of the present species is, of course, homologous with anlage VI of the 18-cirri hypotrichs or the rightmost anlage of the urostyloids because it forms the frontoterminal cirri. r, s: Division of dorsal kineties. No dorsomarginal kineties occur and dorsal kinety fragmentation and caudal cirri are lacking. I, VII = frontalventral-transverse cirri primordia, 1, 2 = dorsal kineties. Page 640.
three) small, 14–18 µm long transverse cirri near rear cell end. Right marginal row somewhat shortened anteriorly, commences at 14% of body length in specimen illustrated, terminates – like left row – at level of transverse cirri; marginal rows thus widely separated posteriorly. Left row begins left of proximal end of adoral zone. Marginal cirri 7–12 µm long; cirri of anterior body half invariable stronger (that is, composed of six basal bodies) than those of rear portion (four basal bodies). Dorsal bristles 3–5 µm long, arranged in two kineties, likely roughly of body length (Fig. 133r). Caudal cirri lacking. Cell division (Fig. 133c–s): This part of the life cycle is described in detail by Hemberger (1982). It commences with the proliferation of some basal bodies at the rearmost cirrus of the midventral complex (Fig. 133c). Due to the formation of new basal bodies a long oral primordium with a short process at the anterior end is
Anteholosticha
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formed (Fig. 133e). Somewhat later the formation of new membranelles in the oral primordium of the opisthe begins (Fig. 133g). Two anteriorly extending processes (primary primordia for the two right anlagen) are recognisable right of the first membranelles. Several anlagen have been formed by the modification of parental cirri on the frontal field. The parental undulating membranes are modified to primordia. As is usual, the three frontal cirri and the two frontoterminal cirri are not involved in primordia formation (Fig. 133i). In total, seven anlagen (I–VII) are produced both in the proter and the opisthe (Fig. 133j). The anlagen I–III of the proter originate from the undulating membrane anlage (anlage I), from the buccal cirrus (II), and cirrus III/2, that is, the cirrus behind the right frontal cirrus (III). The anlagen IV–VII originate from modified cirri of the midventral complex (exact origin not known). I suppose that the anlagen VI and VII are the anterior portion of the two primary primordia mentioned above. The anlagen I–VII form the following number of cirri (Fig. 133a, l, n, p): anlage I – cirrus I and undulating membranes; anlage II – middle frontal cirrus and buccal cirrus; anlage III – right frontal cirrus and cirrus behind (= cirrus III/2); anlage IV – two cirri forming anterior pair of midventral complex; anlage V – two cirri forming middle pair of midventral complex; anlage VI – two cirri forming rear pair of midventral complex plus left transverse cirrus; anlage VII – two frontoterminal cirri plus right transverse cirrus. The slightly increased number of cirral anlagen, the cirral pairs formed by the anlagen IV–VI as well as the lack of dorsomarginal kineties and kinety fragmentation strongly indicate that the present species is a urostyloid. Thus, it is removed from Hemisincirra and transferred to Anteholosticha. The marginal cirri and the dorsal kineties divide in the ordinary manner, that is, each two anlagen occur within the parental structures to form the new row and kineties of the proter and opisthe (Fig. 133j, l, n, p). No caudal cirri are formed (Fig. 13r, s). Prior to division, replication bands are formed in the macronuclear nodules (Fig. 133b). The nodules fuse to a single mass during cell division and later divide into the species-characteristic number of nodules (Fig. 133d, f, h, k, m, o, q). Occurrence and ecology: Very likely confined to terrestrial habitats (Foissner 1987, 1998). Type locality of Anteholosticha heterocirrata not unequivocally fixed; Hemberger (1985, p. 408) mentioned infusions of snail excrement and garden soil. Unfortunately, neither the excrement nor the garden soil are from Peru, the sole country mentioned in the original description (Hemberger 1985, p. 397). According to Hemberger (1982, p. 2) the excrement of the snail Deroceras reticulatum is from the mouth of the river Sieg (Germany) and the garden soil is either from the University of Bonn area (Institut für landwirtschaftliche Zoologie und Bienekunde) or from the village of Bornich (about 50°10'N 7°48'E; Germany). The neotropic distribution in Foissner (1998, p. 204) is therefore incorrect. No further records published, that is, so far only recorded from Europe (Hemberger 1982; Foissner 2000a, p. 258). Food not known. Biomass of 106 specimens about 26 mg (Foissner 1987, p. 124; 1998, p. 204).
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SYSTEMATIC SECTION
Anteholosticha
647
Anteholosticha verrucosa (Foissner & Schade in Foissner, 2000) comb. nov., stat. nov. (Fig. 134a–k, Table 26) 1982 Perisincirra gellerti nov. spec. – Foissner, Arch. Protistenk., 126: 90, Abb. 22i, not Abb. 22a–h (Fig. 134l; misidentification, see remarks). 2000 Hemisincirra gellerti verrucosa Foissner & Schade nov. subspec.1 – Foissner, Europ. J. Protistol., 36: 278, Fig. 84–91, Table 8 (Fig. 134a–k; original description; the holotype slide [accession number 2000/80] and two paratype slides [2000/81, 82] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria; Aescht 2003, p. 386).
Nomenclature: The species-group name verrucos·us, -a, -um (Latin adjective [m; f; n]; warty) refers to the wart-like accumulations of cortical granules (Foissner 2000). I suppose that H. gellerti verrucosa is not conspecific with H. gellerti gellerti as described by Foissner (1982) and therefore raise it to species rank (Hemisincirra verrucosa Foissner & Schade in Foissner, 2000). Furthermore, I transfer it to Anteholosticha (see heading; details, see remarks). Since both Hemisincirra and Anteholosticha are feminine, the ending of the species-group name does not change. Remarks: According to Foissner (2000) Hemisincirra gellerti verrucosa differs from H. gellerti gellerti as described by Foissner (1982) only by the arrangement of the cortical granules, which is in small clusters around the cirri and dorsal bristles in the former, but in narrowly spaced, longitudinal rows in the latter. By contrast, I consider both taxa as valid species which differ from each other, besides the cortical granulation, in the frontoventral row. In H. gellerti it is more or less continuous (Fig. 134a–h), whereas it is rather irregular and obviously composed of cirral pairs in A. verrucosa (Fig. 134c, i, k). Foissner (1982) already recognised this difference (Fig. 134l in present book and Fig. 79a in Berger 2006), and in my review on the urosty-
b
Fig. 134a–k Anteholosticha verrucosa (a–k, from Foissner 2000; l, from Foissner 1982. a–f, k, type population from Tenerife; g–j, Berlin population. a, b, e–h, from life; c, d, i–l, protargol impregnation). a: Ventral view of representative specimen, 70 µm. b: Dorsal view showing cortical granules around dorsal bristles. c, d: Infraciliature of ventral and dorsal side and nuclear apparatus of a specimen of the type population, 60 µm. Arrow in (d) marks short dorsal kinety 5 (often lacking). Dorsal bristles of individual kineties connected by dotted line. e, f: Cortical granules (0.3–0.7 µm across) occur only around cirri and dorsal bristles. g, h: Right lateral and dorsal view of shape variant. i, j: Infraciliature of ventral and dorsal side and nuclear apparatus of a specimen of the Berlin population, 59 µm. k: Anterior portion of (c) enlarged to show supposed origin of frontoventral cirri. I suppose that this species has at least nine (I–IX) frontalventral-transverse cirri anlagen; the cirrus behind the right frontal cirrus (= cirrus III/2) and the pairs IV–VIII form the midventral complex. l: Perisincirra gellerti from the Tullnerfeld area is likely identical with A. bergeri or A. verrucosa. AZM = distal end of adoral zone, FC = right frontal cirrus (connected with middle and left cirrus by dotted line), FT = frontoterminal cirri, LMR = left marginal row, MA = macronuclear nodule, MI = micronucleus, RMR = right marginal row, TC = transverse cirri (pretransverse ventral cirri likely present), I, III, VI, VIII, IX = frontal-ventral-transverse cirri anlage, 1–4 = dorsal kineties. Page 647. 1
Foissner (2000) provided the following diagnosis: Cortical granules only around bases of cirri and dorsal bristles.
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SYSTEMATIC SECTION
loids I therefore supposed that the population from the Tullnerfeld area belongs to Anteholosticha (Berger 2006, p. 393). Anteholosticha verrucosa has the same cirral pattern as the Tullnerfeld population of H. gellerti and therefore is also very likely a holostichid. The ventral infraciliature of A. verrucosa is strongly reminiscent of A. bergeri (Foissner, 1987) Berger, 2003 (see Fig. 78d, f, h in Berger 2006) with which Foissner (2000) did not compare the present species. Therefore, I raise Hemisincirra gellerti verrucosa from subspecies rank to species rank and transfer it to Anteholosticha. The main differences between A. verrucosa and A. bergeri are in (i) the colour of the cortical granules (colourless to yellowish vs. shining-pink in typepopulation and orange in populations described by Blatterer & Foissner 1988); (ii) the number of macronuclear nodules (5–16 [8 and 9 on average] vs. 13–34 [15, 16, and 29 on average]); and (iii) the dorsal kineties (4 kineties with widely spaced bristles vs. 3 kineties with moderately wide spaced bristles). Ontogenetic data are needed to confirm my assumption of the presence of midventral complex. Foissner (2000) characterised two populations in detail and mentioned further populations (e.g., from Australia, Madagascar) which might be, according to him, further subspecies of H. gellerti because they have only three or two dorsal kineties, respectively. This means, that the Anteholosticha bergeri/verrucosa-group is possibly a complex of several, rather similar species. Morphology: Foissner (2000) described two populations (Tenerife [type; Fig. 134a–f, k] and Berlin [Fig. 134g–j]) which are very similar in morphology and all main morphometric features (Table 26). He divided the morphology section in “Type population” and “Berlin population”, but obviously the description of the type population also contains some data (body size, colour and size of cortical granules) from the Berlin population. Thus, the description below does not refer exclusively to the type population. Body size in life 50–100 × 15–22 µm, on average about 70 × 17 µm; body length:width ratio 3.6–7.0:1, on average 4.5:1 in protargol preparations (Table 26). Body flattened up to 2:1 dorsoventrally (Fig. 134g). Body outline elongate rectangular, that is, margins straight or only slightly convex and both ends broadly rounded (Fig. 134a, b, h). About nine macronuclear nodules left of midline one behind the other behind adoral zone; individual nodules globular to elongate ellipsoidal (length:width ratio 3:1), on average broadly ellipsoidal (1.6:1), contain many minute chromatin bodies. Usually one broadly ellipsoidal, compact micronucleus each near end of macronuclear apparatus (Fig. 134a, c, i). Contractile vacuole in mid-body near left cell margin, during diastole with two lacunar collecting canals. Cortex thin, fragile, and very flexible. Cortical granulation described in detail by Foissner (2000); accordingly, granules conspicuous – although minute and only present around bases of cirri and dorsal cilia – because compact and thus highly refractive; dorsal clusters appear as minute, bright warts (species-group name!) at a magnification of m 200×, specific arrangement preserved even in heavily squeezed specimens; individual granules of type population colourless and 0.7 µm across (according to Foissner’s original notes), do not stain with methyl green-pyronin or protargol (Fig. 134b, e, f). Cortical granules of Berlin population 0.3–0.7 µm, yellowish,
Anteholosticha
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around and between cirri and usually semicircularly arranged around dorsal bristles. Cytoplasm colourless, contains some fat globules 1–3 µm across and food vacuoles. Glides moderately rapidly on slide surface and between and on soil particles, showing great flexibility. Adoral zone occupies 27% of body length on average (Table 26; 30% in Berlin population), composed of 15 membranelles on average; longest bases about 4 µm wide in life. Distal (frontal) three membranelles separated from proximal membranelles by a distinct gap which is about one membranelle wide (Fig. 134a, c). Buccal cavity narrow, but rather deep, right posterior portion covered by narrow, inconspicuous lip. Paroral and endoral inconspicuous, both possibly composed of monokinetids, curved, and optically intersecting in mid-buccal cavity. Pharyngeal fibres rather distinct, extend backwards. Cirral pattern and number of cirri of usual variability (Table 26). All cirri about 10 µm long. Pattern very similar (virtually identical) to that of Anteholosticha bergeri (Fig. 134a, c, i, k; see remarks and Berger 2006). Frontal cirri slightly enlarged, arranged in transverse pseudorow. Buccal cirrus slightly behind anterior end of front undulating membrane (paroral?). Two distinct frontoterminal cirri right of and between level of right frontal cirrus and anterior end of midventral complex which extends on average to 42% of body length in type population (Table 26; 47% in Berlin population); cirri of midventral complex slightly zigzagging, especially in anterior half; in addition, the cirri are not uniformly oriented, but alternate between longitudinal an oblique orientations; this arrangement strongly proves that the cirri form a midventral complex, the apomorphy of the urostyloids (for details, see Berger 2006); however, ontogenetic data are needed to confirm (or reject) my hypothesis. About four, fine transverse cirri (possibly 1–2 pretransverse ventral cirri included because rather irregularly arranged) near rear cell end, therefore extend distinctly beyond body proper. Right marginal row distinctly shortened anteriorly (commences at 17% of body length in specimen shown in Fig. 134c, i, k), terminates slightly ahead of level of transverse cirri; left row commences about at level of buccal vertex, terminates about at level of transverse cirri; thus, marginal rows distinctly separated posteriorly (Fig. 134c). Dorsal bristles about 3 µm long in life, invariably arranged in four kineties as shown in Fig. 134b, d, j; kinety 1 invariably(?) distinctly shortened anteriorly; rarely a short kinety 5 present at anterior end of cell (Fig. 134b, d). No caudal cirri. Occurrence and ecology: Very likely confined to terrestrial habitats; common in soils throughout the world, thus cosmopolitan, except for the Antarctic region, and euryoecious (Foissner 2000; Foissner et al. 2002, p. 53). Type locality of A. verrucosa is a pine forest (0–5 cm litter and soil layer; 1400 m above sea-level; 17°W 28°N; collected by B. Krassnigg, University of Salzburg) near the village of Arafo, Tenerife, Canary islands. The soil (pH 6.2) was reddish brown, with many grass roots and more or less completely decomposed pine-needles (Foissner 2000). The Berlin population occurred in soil samples from abandoned sewage irrigation fields (for details, see site 19 in Foissner 2000, p. 255). Further records: soil from the
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SYSTEMATIC SECTION
Enns/Danube River floodplain, Upper Austria (Foissner 2004, p. 370); various forest stands in Austria (Foissner et al. 2005, p. 628); Namibia (Foissner et al. 2000, p. 60). Anteholosticha verrucosa feeds on bacteria (Foissner 2000).
Supplement to the Oxytrichidae This chapter comprises three taxa, two of which (Amphisiellides and Pseudouroleptus) were previously assigned to the amphisiellids by some workers (Tables 4, 7–10). However, in the type species of both genera dorsal kinety fragmentation occurs, strongly indicating that they belong to the oxytrichids (Berger 2006, p. 33). Interestingly, dorsomarginal kineties are not present so that one has to assume that they have been lost, because oxytrichids are (very likely) a subgroup of the Dorsomarginalia (Fig. 9a). Whether this loss is a synapomorphy for Amphisiellides and Pseudouroleptus or occurred independently is not known. I doubt that these two genera are closely related because the frontoventral row is formed rather differently, namely, from one anlage in Amphisiellides and from three anlagen in Pseudouroleptus. Since the Oxytrichidae were already reviewed by Berger (1999), Amphisiellides and Pseudouroleptus are included as supplement to the oxytrichids in the present book. Further details, see remarks in genus sections. Ponturostyla was not considered by Berger (1999) because the dorsal kinety pattern was not known. Song’s (2001) data show that an oxytrichid kinety fragmentation is present, demonstrating the P. enigmatica is an oxytrichid. Note that in the last volume of the revision of Hypotricha a key to all genera, including a reference to the corresponding monograph, will be provided.
Amphisiellides Foissner, 1988 1988 Amphisiellides nov. gen.1 – Foissner, Stapfia, 17: 120 (original description). Type species (by original designation): Uroleptoides atypica Hemberger, 1985. 1990 Amphisiellides – Foissner & Blatterer, J. Protozool., 37: 9A, Abstract 53 (brief note). 1994 Amphisiellides Foissner, 1988 2 – Eigner & Foissner, J. Euk. Microbiol., 41: 255 (redefinition). 1996 Amphisiellides Foissner, 1988 3 – Petz & Foissner, Acta Protozool., 35: 277 (redefinition). 1999 Amphisiellides Foissner, 1988 – Shi, Acta Zootax. sinica, 24: 255 (generic revision of hypotrichs). 1999 Amphisiellides Foissner, 1988 – Shi, Song & Shi, Progress in Protozoology, p. 102 (generic revision of hypotrichs). 2001 Amphisiellides Foissner 1988 – Aescht, Denisia, 1: 21 (catalogue of generic names of ciliates).
1
Foissner (1988) provided the following diagnosis: Amphisiellidae mit mehr als 1 Cirrus links der Ventralreihe im Frontalfeld. Transversal- und Caudalcirren vorhanden. 2 Eigner & Foissner (1994) provided the following improved diagnosis: The amphisiellid median cirral row originates from two rightmost anlagen. One dorsal kinety develops from right marginal row. More than one cirrus left of amphisiellid median cirral row. Transverse cirri longitudinally arranged, usually originate from single anlage. Caudal cirri present. 3 Petz & Foissner (1996) provided the following improved diagnosis: The oral primordium originates in close contact with the amphisiellid median cirral row. The amphisiellid median cirral row commences anlagen formation at its posterior end and originates from two rightmost anlagen. One dorsal kinety develops from the right marginal row. More than one cirrus left of amphisiellid median cirral row. Usually two cirri right of amphisiellid median cirral row. Transverse cirri longitudinally arranged, usually originate from single anlage. Caudal cirri present.
651
652
SYSTEMATIC SECTION
2001 Amphisiellides Foissner, 1988 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 8 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Amphisiellides Foissner, 1988 – Lynn & Small, Phylum Ciliophora, p. 453 (guide to ciliate genera). 2005 Amphisiellides – Berger, Int. Congr. Protozool., 12: 106, Abstract S4.1-25 (brief note on amphisiellids)
Nomenclature: Amphisiellides is a composite of the genus-group name Amphisiella (see there for derivation) and the suffix -ides (similar, especially in shape), indicating a similarity with Amphisiella (Foissner 1988). According to Article 30.1.4.4 of the ICZN (1999), a compound genus-group name ending with -ides is to be treated as masculine, unless its author, when establishing the name, stated that it had another gender or treated it as such by combining it with an adjectival species-group name in another gender form. Foissner (1988) retained the masculine gender for Amphisiellides. Characterisation (A = supposed apomorphy): Oxytrichid with flexible body. Adoral zone of membranelles formed like a question mark, undulating membranes curved. Three frontal cirri. Buccal cirrus present. More than one cirrus left of anterior portion of frontoventral row1. Frontal-ventral-transverse cirri originate from five anlagen, that is, (very likely) anlage V lacking. Frontoventral row originates from anlage VI only. Postoral ventral cirrus lacking. Transverse cirri present. One left and one right marginal row. Dorsal morphogenesis very likely with fragmentation. Dorsomarginal kinety lost (A?). Caudal cirri present. Terrestrial. Remarks: Foissner (1988) established Amphisiellides for Uroleptoides atypicus. A few years later, Eigner & Foissner (1994) described Amphisiellides illuvialis and redefined the genus, mainly using the morphological and ontogenetic data of A. illuvialis. Petz & Foissner (1996) slightly modified this improved diagnosis (see corresponding footnotes). Unfortunately, the improved diagnoses are orientated too strongly on A. illuvialis and not on the type species A. atypicus. Amphisiellides, respectively its type species, was classified in the amphisiellids by most previous workers because of the “amphisiellid median cirral row” (Hemberger 1982, Foissner 1988, Eigner & Foissner 1994, Petz & Foissner 1996, Shi 1999, Shi et al. 1999, Lynn & Small 2002). However, the monophyly of Amphisiellides and its classification in the amphisiellids are very questionable, problems not discussed in detail in previous papers. Eigner & Foissner (1994, p. 254) provided a redefinition of the Amphisiellidae (see footnote on p. 79). Accordingly, the amphisiellid median cirral row is formed by the two (or three) rightmost anlagen. Interestingly, in the type species A. atypicus the “amphisiellid median cirral row” originates very likely only from a single anlage, making the assignment of Amphisiellides to the amphisiellids difficult to understand (Fig. 135d). I suppose that the row is formed by anlage VI because it commences close to the distal end of the adoral zone of membranelles. This is reminiscent of the frontoterminal cirri which are formed from anlage VI. By 1
Because Amphisiellides is (very likely) not an amphisiellid, the specific term “amphisiellid median cirral row” is replaced by the neutral term “frontoventral row”.
Amphisiellides
653
contrast, the cirri formed by anlage V never extend to near the distal end of the adoral zone. The two species included in Amphisiellides at present (A. atypicus, A. illuvialis) differ in the following features: (i) formation of the frontoventral row (from rightmost anlage vs. from two rightmost anlagen); (ii) transverse cirri (three slightly enlarged, transversely arranged cirri [forming a pseudorow!] originating from three anlagen, that is, true transverse cirri present vs. true transverse cirri lacking; see descriptions for details); (iii) dorsomarginal kinety (lacking vs. present); and (iv) dorsal kinety fragmentation (present vs. lacking). These distinct differences strongly indicate that A. atypicus and A. illuvialis are not closely related. Further, both the presence of a fragmenting dorsal kinety in the type species and the presence of a dorsomarginal row in A. illuvialis suggest that they do not belong to the amphisiellids because all other taxa now included in this group lack these features. The presence of a dorsal kinety fragmentation (this feature has to be checked in further populations) assigns A. atypicus to the oxytrichids (Berger 1999), and the dorsomarginal kinety in A. illuvialis indicates that it is a member of the Dorsomarginalia Berger, 2006, but likely not of the Oxytrichidae. Since the oxytrichids are a subgroup of the Dorsomarginalia (Berger 2006, p. 33), one has to assume that the dorsomarginal kinety has been lost in A. atypicus, or one of its recent ancestors. As a consequence, Amphisiellides is removed from the amphisiellids and assigned to the oxytrichids using the dorsal kinety fragmentation as key feature (Berger 1999, 2006). Further, Amphisiellides illuvialis is preliminarily transferred to Nudiamphisiella, which has a very similar cirral pattern and also a dorsomarginal kinety. This new classification is more parsimonious, that is, shows a distinctly lower number of contradictions than the original classification. Eigner (1997, p. 555; 1999, p. 46), who eliminated the Amphisiellidae, assigned Amphisiellides illuvialis to the Parakahliellidae Eigner, 1997, a group derived from a computer analysis based on six features only. In my opinion this group is non-monophyletic, for example, because it contains some stylonychines. The taxon Stylonychinae was established by Berger & Foissner (1997) for the rigid oxytrichids which, inter alia, have lost the cortical granules in their last common ancestor. Berger (1999) added some non-18cirri oxytrichids (e.g., Pattersoniella, Laurentiella, Onychodromus) because they also have the stylonychine apomorphies. Later, the Stylonychinae were confirmed by molecular data (e.g., Modeo et al. 2003, Foissner et al. 2004, Schmidt et al. 2007). Pseudouroleptus caudatus is a further “amphisiellid” hypotrich with a dorsal kinety fragmentation (Fig. 136c). Thus, it was transferred to the oxytrichids by Berger (1999, p. 888; see also p. 661 in present book). Pseudouroleptus differs from Amphisiellides, inter alia, in the number of frontal-ventral cirri anlagen (6 vs. 5), the cirral pattern (transverse cirri lacking vs. present; postoral ventral cirrus present vs. absent), and the number of anlagen forming the frontoventral row (3 vs. 1). Species included in Amphisiellides (basionym given): (1) Uroleptoides atypica Hemberger, 1985.
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SYSTEMATIC SECTION
Species misplaced in Amphisiellides: The following species, originally assigned to Amphisiellides, has been removed from the present genus. Amphisiellides illuvialis Eigner & Foissner, 1994. Remarks: Now Nudiamphisiella illuvialis (p. 569).
Key to Amphisiellides atypicus and Amphisiellides-like species 1 About 23 macronuclear nodules; (usually) three transversely arranged transverse cirri (Fig. 135a). . . . . . . . . . . . . . . . . . . . . . . . . . Amphisiellides atypicus (p. 654) - Two macronuclear nodules. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2 Cirri (longitudinally or transversely arranged) between posterior portion of marginal rows present (e.g., Fig. 120b). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 - Cirri between posterior portion of marginal rows absent (Fig. 119g). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nudiamphisiella interrupta (p. 562) 3 Usually two longitudinally arranged “transverse cirri” (Fig. 120a, b, i). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nudiamphisiella illuvialis (p. 569) - Five transversely arranged transverse cirri (Fig. 24a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amphisiella oscensis (p. 133)
Single species Amphisiellides atypicus (Hemberger, 1985) Foissner, 1988 (Fig. 135a–e, Tables 35, 41) 1982 Uroleptoides atypica n. spec.1 – Hemberger, Dissertation2, p. 49, Abb. 7a–c (Fig. 135a–e; see nomenclature). 1985 Uroleptoides atypica n. spec. – Hemberger, Arch. Protistenk., 130: 400, Abb. 4 (Fig. 135a, b; original description; no formal diagnosis provided; type slides are deposited in the Institut für landwirtschaftliche Zoologie und Bienenkunde, University of Bonn, Germany). 1988 Uroleptoides atypicus nom. corr. – Foissner, Stapfia, 17: 120 (corrected name; see nomenclature). 1988 Amphisiellides atypicus (Hemberger, 1985) nov. comb. – Foissner, Stapfia, 17: 120 (combination with Amphisiellides and fixation as type species of Amphisiellides). 1994 Amphisiellides atypicus (Hemberger, 1985) – Eigner & Foissner, J. Euk. Microbiol. 41: 245, Fig. 11 (Fig. 135a, b; comparison with A. illuvialis). 2001 Amphisiellides atypicus (Hemberger, 1985) Foissner, 1988 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 96 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs).
Nomenclature: The species-group name atypic·us, -a, -um (Latin adjective [m; f; n]; atypical) is a composite of the prefix a+ (negation) and the Greek adjective typic·us, 1 2
This name is disclaimed for nomenclatural purposes (ICZN 1999, Article 8.3). See next footnote. See corresponding footnote at Uroleptoides binucleatus.
Amphisiellides
655
-a, -um (m, f, n; typical, normal, true), and refers to the transverse cirri, which are clearly recognisable, as compared to those of the other members of Uroleptoides (Hemberger 1985). Amphisiellides atypicus is the type species of Amphisiellides. Hemberger (1982, 1985) classified the present species in Uroleptoides, which is masculine. Thus, Foissner (1988) introduced the nomen corrigendum Uroleptoides atypicus. Remarks: Amphisiellides atypicus was described by Hemberger (1985) as Uroleptoides atypica mainly after protargol impregnation. Thus, some features, which are only clearly recognisable in life, are not known, for example, the presence/absence of cortical granules. Foissner (1988), who synonymised – like Borror (1972) and Jankowski (1979) – Uroleptoides with Amphisiella, established Amphisiellides for the present species because it has a unique combination of features (see corresponding footnote in genus section). Hemberger (1982) found two dividers showing two important features, namely, (i) that the frontoventral row is (very likely) formed from the rightmost anlage only; and (ii) that (very likely) dorsal kinety fragmentation occurs because the species has four dorsal kineties, but only three within row anlagen are formed per filial product and a dorsomarginal kinety is lacking (further details, see cell division). These two characteristics and the presence of true transverse cirri separate it clearly from Amphisiellides illuvialis, the second species assigned to Amphisiellides. Thus, I transfer A. illuvialis preliminary to Nudiamphisiella (further details see remarks under Amphisiellides and N. illuvialis). To get a better idea about the systematic position of A. atypicus, a redescription, a detailed study of the cell division (ventral and dorsal), and the analyses of meaningful molecular markers are needed. Amphisiella oscensis Fernandez-Leborans, 1984 has a cirral pattern closely resembling that of A. atypicus (Fig. 24a). However, it also resembles Caudiamphisiella antarctica, especially as concerns the true transverse cirri and the dorsal kinety pattern. Thus, Amphisiella oscensis is preliminarily assigned to Caudiamphisiella, however, without formal combination. Morphology: Body size (in life?) about 200 × 30–40 µm, body length:width ratio 5:1 on average (Table 41). Body outline roughly rectangular, that is, margins parallel and both ends broadly rounded. About 23 macronuclear nodules arranged in a stripe from proximal portion of adoral zone to last fifth of body; individual nodules of rather different size (Fig. 135b). At least two micronuclei almost of same size as macronuclear nodules. Contractile vacuole in ordinary position, that is, near left cell margin somewhat behind proximal end of adoral zone; at 28% of body length in specimen illustrated (Fig. 135a). Cytoplasm heavily granulated. Presence/absence of cortical granules and movement not known. Adoral zone occupies about 20% of body length, composed of 23–25 membranelles of ordinary fine structure; proximal portion of zone sigmoidal and membranelles rather wide (Fig. 135a). Undulating membranes distinctly curved. Anterior portion of peristomial lip distinctly curved backwards.
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SYSTEMATIC SECTION
Fig. 135a–e Amphisiellides atypicus (a, b, from Hemberger 1985; c–e, from Hemberger 1982. Protargol impregnation. Outline of specimen shown in [a] likely from life). a, b: Infraciliature of ventral side and nuclear apparatus, 200 µm. Frontalventral cirri which very likely originate from the same anlage are connected by broken lines (only shown for anlagen I–IV). c: Middle divider showing that most parental cirri (buccal cirrus; cirrus cirricirrus; left of cirrus frontoventral row, and middle portion of fronrental cirriIII/2; (buccal III/2; cirri leftposterior of frontoventral row, posterior and portion of frontoventral row) and the undulating membranes are involved of in toventral row) and the middle undulating membranes are involved in primordia formation. d, e: Infraciliature ventral side and nuclear apparatus of a late d, divider. This divider shows that row ofofA.a primordia formation. e: Infraciliature of ventral sidethe andfrontoventral nuclear apparatus atypicus originates from a single (vs. from two in illuvialis, was originates originally late divideranlage This divider shows thatNudiamphisiella the frontoventral row of Awhich atypicus assigned to Amphisiellides). Arrows mark new marginal rows of opisthe. CC = caudal cirri, CV = contractile vacuole, FC = right frontal cirrus, FVR = rear end of frontoventral row, MA = macronuclear nodule, MI = micronucleus, TC = transverse cirri, I–VI = frontal-ventral-transverse cirral anlagen (note that anlage V is very likely lacking in this species). Page 654.
The cirral pattern is reminiscent of amphisiellids; however, the dorsal kinety pattern and the frontoventral row formation show that it does not belong to this group (see remarks and cell division). Three enlarged frontal cirri, right one, as is usual, near distal end of adoral zone. Buccal cirrus in mid-region of paroral. Three cirri left of anterior portion of frontoventral row; the left one is likely cirrus III/2 although it is behind the middle frontal cirrus, the other two are very probably formed by anlage IV, that is, one of them is homologous to cirrus IV/3. Frontoventral row commences near distal end of adoral zone, extends sigmoidally to 55% of body length in specimen illustrated (Fig. 135a). Three slightly enlarged transverse cirri near posterior body end, about 18 µm long and thus likely distinctly projecting beyond rear body margin. Right marginal row obviously more or less distinctly shortened anteriorly,
Amphisiellides
657
Table 41 Morphometric data on Amphisiellides atypicus (aty, from Hemberger 1982, 1985) and Amphisiella oscensis (osc, from Fernandez-Leborans 1984; preliminary classified in Caudiamphisiella) Characteristics a Body, length Body, width Macronuclear nodule, length Macronuclear nodule, width Macronuclear nodules, number Micronuclei, number Adoral membranelles, number Frontal cirri, number Buccal cirri, number Cirri left of anterior portion of frontoventral row, number Frontoventral row, number of cirri Transverse cirri, number Left marginal row, number of cirri Right marginal row, number of cirri Dorsal kineties, number Caudal cirri, number
Species
mean
b
aty 200.0 osc – aty – osc – osc 17.0 osc 9.1 aty 23.0 osc 2.0 aty osc 2.0 aty – osc – aty 3.0 aty 1.0 osc 1.0 aty c 3.0 aty osc aty osc aty osc aty osc aty osc aty osc
– – 3.0 5.0 – – – – 4.0 3.0 3.0 –
M
SD
SE
– – – – – – – –
– – – – – – – –
– – – – – – – –
– – – – – – – – – – – – – – – – – – 7.0
CV
Min
Max
n
– 77.7 30.0 31.0 – – – –
– 81.0 40.0 33.0 – – – –
– – – – – – –
– – – – – – – – at least 2 – – – – – – – – – – – – – –
– 23.0 38.0 – – – –
– 25.0 41.0 – – – –
15 ? 15 ? ? ? 15 ? 15 ? 15 ? 15 15 ? 15
– – – – – – – – – – – –
– – – – – – – – – – – –
27.0 17.0 – – 47.0 28.0 47.0 32.0 – – – 6.0
30.0 28.0 – – 53.0 30.0 55.0 34.0 – – – 7.0
15 ? 15 ? 15 ? 15 ? 15 ? 15 ?
– – – – – – – – – – – –
a All measurements in µm. CV = coefficient of variation in %, M = median, Max = maximum value, mean = arithmetic mean, Min = minimum value, n = number of individuals investigated, SD = standard deviation, SE = standard error of arithmetic mean. Data based on protargol-impregnated specimens. When two values are present, then they are listed as MIN and Max; when only one value is given, it is listed as mean. b
Method (from life, protargol preparations) not clearly indicated.
c
Cirrus III/2 included.
extends – like left row – to level of transverse cirri; left row commences left of proximal end of adoral zone. Marginal cirri 12–15 µm long. Dorsal bristles 3 µm long, arranged in four kineties (see cell division); exact pattern not described, making higher level classification difficult. Three caudal cirri, not elongated, that is, likely of same length as marginal cirri, composed of three cilia only. Cell division: Hemberger (1982) found an early to middle and a late divider (Fig. 135c–e). They show that the middle and posterior portion of the frontoventral
658
SYSTEMATIC SECTION
row, the parental undulating membranes, and some parental cirri (buccal cirrus, cirrus III/2 and other cirri left of anterior portion of frontoventral row) are involved in anlagen formation (Fig. 135c). The later stage shows that in total five frontalventral-transverse cirri anlagen are formed. They produce the following cirri: anlage I forms the undulating membranes and the left frontal cirrus; anlage II forms the middle frontal cirrus, the buccal cirrus, and the leftmost transverse cirrus; anlage III forms the right frontal cirrus, cirrus III/2, and the middle transverse cirrus; anlage IV forms two cirri left of the anterior portion of the frontoventral row and the right transverse cirrus; anlage V or VI forms the frontoventral row, that is, anlage VI or V is lacking in this species. The fact that the frontoventral row commences near the anterior cell end strongly indicates that it is produced by anlage VI, that is, the frontoventral row is likely homologous to the frontoterminal cirri which are invariably arranged near the anterior cell end. By contrast, the cirri formed by anlage V are usually located behind the level of the buccal vertex. Hemberger (1982) described, but did not illustrate the morphogenesis of the dorsal infraciliature. He found only three within-row-anlagen per filial product, strongly indicating that one kinety fragments because non-dividers have four kineties and no dorsomarginal kinety is formed. Of course one cannot exclude that a parental row is retained after cell division, which would also result in the presence of four kineties. However, the parental kineties have widely spaced bristles, a conspicuous feature not described by Hemberger (1982). The formation of marginal rows shows no peculiarities, except that no dorsomarginal kinety, which would be characteristic for the Dorsomarginalia and the oxytrichids, is produced. The many macronuclear nodules fuse, as is usual, to a single mass during cell division (Fig. 135e). Although these data are rather informative, a more detailed investigation of the cell division is recommended. Occurrence and ecology: Amphisiellides atypicus is likely confined to terrestrial habitats (Foissner 1987, 1998). Type locality is the Puerto Maldonado region (69°12’W 12°36’S), Madre de Dios, Peru, where Hemberger (1985) discovered it in a soil sample from a pasture. No further records published. Food not known. Biomass of 106 specimens about 184 mg (Foissner 1987, p. 128; 1998, p. 199).
Pseudouroleptus Hemberger, 1985 1982 Pseudouroleptus n. gen.1 – Hemberger, Dissertation2, p. 36. 1985 Pseudouroleptus n. gen.3 – Hemberger, Arch. Protistenk., 130: 398 (original description). Type species (by original designation): Pseudouroleptus caudatus Hemberger, 1985.
1
This name is disclaimed for nomenclatural purposes (ICZN 1999, Article 8.3). See next footnote. See same footnote at Uroleptoides binucleatus. 3 Hemberger (1985) provided the following diagnosis: Je 1 rechte und linke Marginalreihe; 2 Ventralreihen; keine (oder 2 unscheinbare) Transversalciren; deutlich differenzierte Frontalcirren; Körper terminal zugespitzt; Cirrenentwicklung aus familientypischen Longitudinalanlagen; 3 longitudinale Frontalcirrenanlagen. 2
Pseudouroleptus
659
1987 Pseudouroleptus Hemberger, 1981 – Tuffrau, Annls Sci. nat. (Zool.), 8: 115 (classification of hypotrichs). 1994 Pseudouroleptus Hemberger, 1985 1 – Eigner & Foissner, J. Euk. Microbiol., 41: 260 (generic revision of amphisiellids). 1996 Pseudouroleptus Hemberger, 1985 2 – Petz & Foissner, Acta Protozool., 35: 277 (generic revision of amphisiellids and key to genera). 1999 Pseudouroleptus Hemberger, 1985 – Berger, Monographiae biol., 78: 888 (revision, including transfer from amphisiellids to oxytrichids). 1999 Pseudouroleptus Hemberger, 1985 – Shi, Song & Shi, Progress in Protozoology, p. 106 (generic revision of hypotrichs). 2001 Pseudouroleptus Hemberger 1985 – Aescht, Denisia, 1: 137 (catalogue of generic names of ciliates). 2001 Pseudouroleptus Hemberger, 1985 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 77 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Pseudouroleptus Hemberger, 1981 – Lynn & Small, Phylum Ciliophora, p. 451 (guide to ciliate genera).
Nomenclature: No derivation of the name is given in the original description and the review by Berger (1999). Pseudouroleptus is a composite of the Greek word pseudo- (wrong, lying) and the genus-group name Uroleptus, indicating a similarity to Uroleptus, likely because of the posteriorly narrowed body in both taxa; however, the cirral pattern is rather different! Uroleptus is a composite of ur·o- (Greek noun; tail, rear body portion), lept- (Greek adjective; small, thin, narrow), and the suffix ·us, and refers to the tailed posterior body portion characterising this group. Masculine gender (Aescht 2001, p. 297). Characterisation (A = supposed apomorphy): Oxytrichid with flexible body. Adoral zone of membranelles formed like a question mark. Undulating membranes in Oxytricha pattern. Three enlarged frontal cirri. Buccal cirrus present. One cirrus (= III/2) left of anterior portion of left frontoventral row. Two frontoventral rows. Left frontoventral row composed of cirri from anlagen VI (anterior portion), IV (middle portion), and V (rear portion). Right frontoventral row not involved in primordia formation. Cirrus IV/2 (= postperistomial cirrus) present. Pretransverse ventral and transverse cirri lacking. Dorsal kinety fragmentation in kinety 3. Dorsomarginal kineties lacking. Caudal cirri present. Terrestrial. Remarks: Pseudouroleptus was established for a tailed species with two long frontoventral cirral rows and a postperistomial ventral cirrus. Hemberger (1985) did not propose a higher level classification; however, in his dissertation he assigned it to the Amphisiellidae (Hemberger 1982, p. 36), a classification which was taken 1
Eigner & Foissner (1994) provided the following diagnosis: The amphisiellid median cirral row originates from three rightmost anlagen. Usually one postperistomial cirrus develops from third anlage from right. One cirrus left of amphisiellid median cirral row. Transverse cirral row nearly as long as body, parallels amphisiellid median cirral row, originates from single anlage. Caudal cirri present. 2 Petz & Foissner (1996) provided the following diagnosis: The oral primordium originates in close contact with the amphisiellid median cirral row. The amphisiellid median cirral row commences anlagen formation within-row and originates from three rightmost anlagen. All dorsal kineties develop intrakinetally. Usually one postperistomial cirrus developing from third anlage from right. One cirrus left of amphisiellid median cirral row. Transverse cirral row almost as long as body, parallels amphisiellid median cirral row, originates from single anlage. Caudal cirri present.
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SYSTEMATIC SECTION
over by Eigner & Foissner (1994), Petz & Foissner (1996), and Lynn & Small (2002). By contrast, Tuffrau (1987), Foissner & Foissner (1988, p. 82), and Tuffrau & Fleury (1994, p. 137) classified it in the Kahliellidae Tuffrau, 1979. Eigner (1997, p. 555; 1999, p. 46) assigned it the Oxytrichidae, because it shows neokinetal 3 anlagen development, that is, a large V-shaped primordium produces the anlagen V and VI of both the proter and the opisthe. I put Pseudouroleptus also in the Oxytrichidae, because Hemberger (1982, p. 36) described and illustrated a dorsal kinety fragmentation (Fig. 136c), the main feature of the oxytrichids (Berger 1999, Berger 2006, p. 33). Shi et al. (1999) and Shi (1999, p. 258) classified it in the Keronopsidae Jankowski, 1979, an assignment difficult to follow. In the monograph on the oxytrichids I reviewed only the type species (P. caudatus) in detail because it is the sole species assigned to Pseudouroleptus for which a dorsal kinety fragmentation is described (Berger 1999, 2001). The other four species have a somewhat different cirral pattern and a different number of dorsal kineties (2–4), strongly indicating that Pseudouroleptus is not monophyletic. Thus, I confined Pseudouroleptus to the type species, however, without transferring the other four species to one or more different taxa (Berger 1999). No new data for any of the four species has become available during the last years so that they are still in Pseudouroleptus (see Online catalogue of ciliate names at http://www.protozoology. com). However, this is unsatisfactory and therefore I establish Bistichella for the four Pseudouroleptus species not reviewed by Berger (1999). Since their relationship to other groups is difficult to estimate, I preliminarily classify Bistichella as incertae sedis in the Hypotricha (see p. 532). In addition, I again review the type species of Pseudouroleptus, which we redescribed and divided into two subspecies (Foissner et al. 2002). Pseudouroleptus caudatus is reminiscent of some Hemiamphisiella species which have basically the same cirral pattern. Further, some Hemiamphisiella species have, like P. caudatus, four dorsal kineties, namely, the type population of H. terricola and H. wilberti (Fig. 56f, 60g). Eigner & Foissner (1994) studied the cell division of H. terricola. However, their Australian population had only three dorsal kineties originating intrakinetally so that we do not know the formation-type of kinety 4 (intrakinetally? by fragmentation of kinety 3? dorsomarginal kinety? parental row?) in H. terricola because it is not certain that the Australian population is conspecific with the type material. Thus, I preliminarily consider both Hemiamphisiella (no dorsal kinety fragmentation and thus classified in the amphisiellids) and Pseudouroleptus (type species with dorsal kinety fragmentation and thus classified in the oxytrichids) as valid. However, synonymy of Hemiamphisiella and Pseudouroleptus cannot be excluded. Further populations have to be studied in detail (including dorsal morphogenesis), to clear up the situation finally. Note that the very similar ventral cirral pattern – for example, the long frontoventral row in Pseudouroleptus or the amphisiellid median cirral row in Hemiamphisiella – in two taxa is no guarantee that these two taxa are closely related. A similar convergence is known for the so-called midventral pattern, which evolved several times independently, for example, in the
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661
urostyloids, in Uroleptus, and in the oxytrichid Neokeronopsis (for review, see Berger 2006). Petz & Foissner (1996) wrote, likely par lapsus, that in Pseudouroleptus all dorsal kineties develop intrakinetally. However, this is incorrect for the type species P. caudatus. Uroleptus sp. sensu Martin et al. (1981) possibly belongs to Pseudouroleptus. However, no illustrations are provided, making a redetermination more or less impossible because details of the cirral and dorsal kinety pattern are not recognisable in the micrographs. Species included in Pseudouroleptus (basionym given): Pseudouroleptus caudatus Hemberger, 1985. Species misplaced in Pseudouroleptus: The species listed below were originally assigned or transferred to Pseudouroleptus. However, the dorsal kinety pattern strongly indicates they do not belong to this genus (details see remarks above). Paraurostyla buitkampi Foissner, 1982. Remarks: Now Bistichella buitkampi (p. 535). Pseudouroleptus procerus Berger & Foissner, 1987. Remarks: Now Bistichella procera (p. 547). Pseudouroleptus terrestris Hemberger, 1985. Remarks: Now Bistichella terrestris (p. 554). Uroleptus humicola Gellért, 1956b. Remarks: Now Bistichella humicola (p. 556).
Single species Pseudouroleptus caudatus Hemberger, 1985 (Fig. 136a–i, 137a–c, Table 42) 1982 Pseudouroleptus caudatus n. spec.1 – Hemberger, Dissertation, p. 36, Abb. 4 (Fig. 136a–c; see nomenclature). 1985 Pseudouroleptus caudatus n. spec. – Hemberger, Arch. Protistenk., 130: 399, Fig. Abb. 1 (Fig. 136a, b; original description, see also footnote on previous entry; no formal diagnosis provided; type slides are deposited in the Institut für landwirtschaftliche Zoologie, University of Bonn, Germany). 1999 Pseudouroleptus caudatus Hemberger, 1985 – Berger, Monographiae biol., 78: 889, Fig. 219.1a–j (Fig. 136a–c; revision). 2001 Pseudouroleptus caudatus Hemberger, 1985 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 77 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2002 Pseudouroleptus caudatus Hemberger, 1985 2 – Foissner, Agatha & Berger, Denisia, 5: 649, Fig. 145a–g, 146a–c (Fig. 136a, b, d–i, 137a–c; redescription of nominotypical subspecies and original description of subspecies namibiensis). 1
This name is disclaimed for nomenclatural purposes (ICZN 1999, Article 8.3). For details see footnote 2 under genus Pseudouroleptus. 2 Foissner et al. (2002) provided the following diagnosis: Size about 230 × 50 µm in vivo. Slenderly lanceolate with posterior portion narrowed tail-like. 2 macronuclear nodules with 1 micronucleus each. Cortical granules in closely spaced rows, colourless, about 1.2 µm across. Amphisiellid median cirral row (ACR) extends to near body end and consists of about 56–60 cirri. On average 45–55 adoral
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SYSTEMATIC SECTION
2002 Pseudouroleptus caudatus caudatus Hemberger, 1985 nov. stat. – Foissner, Agatha & Berger, Denisia, 5: 650, Fig. 145a–g, Table 127 (Fig. 136d–i; redescription of nominotypical subspecies; 3 voucher slides [accession numbers 2002/573–575; Foissner et al. 2002, p. 42] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria). 2002 Pseudouroleptus caudatus namibiensis nov. sspec. – Foissner, Agatha & Berger, Denisia, 5: 652, Fig. 146a–c, Table 127 (Fig. 137a–c; original description of subspecies; the holotype slide [accession number 2002/449; Foissner et al. 2002, p. 42] and 4 paratype slides [2002/452–455] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria).
Nomenclature: No derivation of the name is given in the original description. The species-group name caudat·us, -a, -um (Latin adjective [m, f, n]; tailed) obviously refers to the tail-like posterior body portion. Pseudouroleptus caudatus is the type species of Pseudouroleptus. Remarks: The improved diagnosis by Foissner et al. (2002; see relevant footnote) is based on the original description and our investigations on the two subspecies. We found cortical granules in the African (Zanzibar; description see below) and Brazilian population, strongly indicating that they were overlooked by Hemberger (1985), who discovered P. caudatus in Peru. Very likely Hemberger did not study live specimens in detail, although he mentioned “cytoplasm slightly brownish”, a feature probably caused by the dense cortical granulation. We split the species in two subspecies mainly because of a distinct difference in the number of cirri forming the right frontoventral row (Foissner et al. 2002). It is of body length in the nominotypical subspecies P. caudatus caudatus (Fig. 136a, h, i), but commences about at mid-body in P. caudatus namibiensis (Fig. 137a). In life, Pseudouroleptus caudatus is easily confused with Hemiamphisiella granulifera (Fig. 61a–e), which has a similar body size, nuclear pattern, and cortical granulation, but lacks the long right frontal-ventral row. In addition, no dorsal kinety fragmentation is described for Hemiamphisiella. For a comparison with H. wilberti see P. caudatus namibiensis. Amphisiellides atypicus is easily distinguished, inter alia, by the rectangular body outline (vs. pisciform), the true transverse cirri (vs. lacking), and – most important – the many (about 23) macronuclear nodules (vs. two; Fig. 135a–e). Subspecies included in Pseudouroleptus caudatus (alphabetically arranged according to basionyms): (1) Pseudouroleptus caudatus Hemberger, 1985; (2) Pseudouroleptus caudatus namibiensis Foissner, Agatha & Berger, 2002.
Key to the Pseudouroleptus caudatus subspecies 1 Right frontoventral row commences near level of buccal cirrus and extends to rear body end, composed of about 40–50 cirri (Fig. 136a, h, i). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudouroleptus caudatus caudatus (p. 663)
membranelles, 55 cirri in right and near 50 in left marginal row, 1 cirrus left of ACR row, 1 buccal and 1 postperistomial cirrus, 25–50 transverse cirri forming a shortened or unshortened row right of ACR, and 4 dorsal kineties.
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663
Right frontoventral row commences near mid-body and extends to rear body end, composed of about 25 cirri (Fig. 137a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudouroleptus caudatus namibiensis (p. 667)
Pseudouroleptus caudatus caudatus Hemberger, 1985 (Fig. 136a–i, Table 42) 1982 Pseudouroleptus caudatus n. spec.1 – Hemberger, Dissertation, p. 36, Abb. 4a–k (Fig. 136a–c). 1985 Pseudouroleptus caudatus n. spec. – Hemberger, Arch. Protistenk., 130: 399, Fig. Abb. 1 (Fig. 136a, b; original description, see also footnote on previous entry; no formal diagnosis provided; type slides are deposited in the Institut für landwirtschaftliche Zoologie, University of Bonn, Germany). 1999 Pseudouroleptus caudatus Hemberger, 1985 – Berger, Monographiae Biol., 78: 889, Fig. 219.1a–j (Fig. 136a–c; revision). 2002 Pseudouroleptus caudatus caudatus Hemberger, 1985 nov. stat. – Foissner, Agatha & Berger, Denisia, 5: 650, Fig. 145a–g, Table 127 (Fig. 136d–i; redescription of nominotypical subspecies; three voucher slides [accession numbers 2002/573–575; Foissner et al. 2002, p. 42] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria).
Nomenclature: For derivation of the subspecies-group name, see this chapter under P. caudatus. Remarks: The data by Foissner et al. (2002) match those of the original description rather well, except for the cortical granules, which were very likely overlooked by Hemberger (1985), who did not make detailed live observations (see remarks at species). Specifically, body size and shape, the nuclear pattern, and the main morphometric features are very similar (Fig. 136a, d, h, Table 42) so that conspecificity is beyond reasonable doubt. Morphology: For description of type population, see Hemberger (1982, 1985) and the review by Berger (1999, p. 889). The following description is from the Zanzibar population investigated by Foissner et al. (2002; Fig. 136d–i, Table 42). Body size 180–280 × 40–60 µm in life, on average near 230 × 50 µm; length:width ratio about 5:1 in life, while 4.3:1 on average in protargol preparations where specimens are inflated because they are difficult to fix with the usual fixatives (Table 42). Body outline pisciform with tail-like posterior portion, a conspicuous feature preserved even in most protargol-impregnated specimens; invariably slightly twisted about main body axis, marginal rows thus never recognisable in full length if cells are viewed ventrally (Fig. 136d, g). Body acontractile, highly flexible, and dorsoventrally flattened up to 2:1. Nuclear apparatus in middle body third, left of midline. Macronuclear nodules distinctly separate, elongate ellipsoidal on average (3:1), contain many small globular chromatin bodies. Micronuclei highly conspicuous in life and protargol preparations because compact and large, that is, about 10 × 7 µm in life (Fig. 136d, i). Contractile vacuole slightly ahead of mid-body at left cell margin; 1
This name is disclaimed for nomenclatural purposes (ICZN 1999, Article 8.3). For details, see footnote 2 unde genus Pseudouroleptus.
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SYSTEMATIC SECTION
Fig. 136a–c Pseudouroleptus caudatus caudatus (from Hemberger 1982, 1985. Protargol impregnation). a, b: Infraciliature of ventral side and nuclear apparatus of non-dividing specimen, 233 µm. For detailed labelling see Fig. 136h, i. c: Infraciliature of dorsal side of middle divider, size not indicated. Long arrows mark fragmentation of dorsal kinety 3, short arrows mark caudal cirri of proter. The kinety fragmentation unequivocally assigns Pseudouroleptus caudatus, type of the genus, to the oxytrichids. MA = macronuclear nodule, MI = micronucleus, 1–4 = new dorsal kineties of proter (parental kineties not shown). Page 663.
we did not look for collecting canals. Cortex flexible, contains closely spaced rows of about 1.2 µm-sized granules leaving blank only cirral areas; granules provide cells with a brownish colour, although individual granules are colourless, likely due to some light refraction (Fig. 136f); impregnate lightly with protargol. Cytoplasm colourless, packed with food vacuoles, fat globules, minute, protargol-affine granules, and various crystals up to 5 µm in size which make cells dark in posterior half, where they are concentrated (Fig. 136d, e). Glides rapidly on microscope slide and on and between soil particles, showing great flexibility. Adoral zone occupies 21–37%, on average 28% of body length, extends far (18 µm on average) onto right body margin, that is, to about level of buccal cirrus; composed of an average of 54 membranelles of ordinary fine structure; bases of largest membranelles 10 µm wide in life. Buccal cavity deep and of ordinary width, right half and last adoral membranelles covered by a very hyaline lip (Fig. 136d, g, h). Undulating membranes slightly curved and side by side, rarely intersecting optically in posterior third, paroral likely composed of dikinetids. Pharyngeal fibres distinct in live and protargol preparations, extend obliquely backwards.
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665
Cirral pattern and number of cirri of usual variability (Fig. 136d, h, Table 42). Three distinctly enlarged frontal cirri, in life about 20 µm long; left cirrus in body midline, right one behind distal end of adoral zone, that is, about at level of buccal cirrus which is right of anterior portion (0–5 µm distant from anterior end) of paroral. Invariably one enlarged cirrus (= cirrus III/2 or paramalar cirrus) behind right frontal cirrus, that is, left of anterior end of left frontoventral row. Postperistomial cirrus (= cirrus IV/2 of the 18-cirri oxytrichids) slightly enlarged, very near to buccal vertex. Left frontoventral row (termed “amphisiellid median cirral row” by Foissner et al. 2002) conspicuous because composed of an average of 60 cirri, commences at right body margin behind distal end of adoral zone and extends obliquely to left body margin to end subterminally on average. Right frontoventral row1 (termed “transverse cirri” by Foissner et al. 2002) commences at same level as left row, but extends to posterior body end. Pretransverse ventral and transverse cirri lacking. Marginal rows follow body curvature and are thus distinctly spiralled, extend to posterior body end. Right row commences dorsolaterally about at same level as dorsal kineties, left row begins left of posterior portion of adoral zone of membranelles. Frontal-ventral and marginal cirri about 15 µm long in life, closely spaced within rows, distances between individual cirri increase slightly in posterior third of rows. Dorsal bristles 3 µm long; ciliature (number and arrangement of kineties; presence/absence of caudal cirri) of Zanzibar population not known because poorly impregnated (Fig. 136i). Cell division (Fig. 136c). This process is described by Hemberger (1982) and reviewed in the monograph on oxytrichids (Fig. 219.1b–j in Berger 1999). Thus, only the stage with the dorsal kinety fragmentation is shown again in the present book (Fig. 136c) because this fragmentation is substantial for the assignment of Pseudouroleptus to the oxytrichids (for details, see Berger 1999). The formation of the ventral ciliature of P. caudatus caudatus closely resembles that of Gastrostyla (also shows dorsal kinety fragmentation) and Hemiamphisiella because in all three taxa the (left) frontoventral row is composed of cirri of anlage VI (anterior portion), IV (middle portion), and V (rear portion). Further, these taxa have a postperistomial ventral cirrus (= cirrus IV/2 of the 18-cirri hypotrichs), which is lacking in the typical amphisiellids. For Gastrostyla, the oxytrichid relationship has been confirmed by gene sequence data (e.g., Croft et al. 2003, Foissner et al. 2004, Schmidt et al. 2007). Occurrence and ecology: Likely prefers terrestrial habitats, but also recorded from a running water. Type locality of P. caudatus caudatus is the Puerto Maldonada region (12°36'S, 69°12'W) in Peru (details from Hemberger 1982, p. 2), where Hemberger (1982, 1985) found it in a forest soil and in plankton(?)-samples (“Freiwasserproben”) from the Rio Tambopata, possibly due to soil erosion. We 1
This is the posterior portion formed by frontal-ventral cirral anlage VI. Not even the rearmost cirrus of this row should be designated as transverse cirrus because it has the same size and is not set off from the other cirri.
666
SYSTEMATIC SECTION found it in a grey, sandy, circumneutral (pH 7.8), dry rice field soil collected by Hubert Blatterer (Linz, Upper Austria) in the surroundings of the tourist office of the town of Zanzibar (07°S 39°E), Tanzania (Foissner et al. 2002). In addition, we found it in Brazil (Foissner et al. 2002, p. 649). Further record (not substantiated by morphological data): Guadarama River, Spain (Olmo Rísquez 1998, p. 17). Feeds on coccal cyanobacteria, naked amoebae, flagellates, diatoms, and small and medium-sized testate amoebae (Trinema lineare, T. enchelys) and ciliates like Plagiocampa sp. and Metopus hasei (Foissner et al. 2002). Biomass of 106 specimens about 375 mg (Foissner 1987, p. 127); Foissner (1998) did not list P. caudatus in his updated compilation of world soil ciliates.
Fig. 136d–g Pseudouroleptus caudatus caudatus (from Foissner et al. 2002. From life). d: Ventral view of a representative specimen, 236 µm. Note the three large, compact micronuclei. e: Cytoplasmic crystals up to 5 µm in size. f: Surface view showing closely spaced rows of colourless cortical granules about 1.2 µm across. g: Blunt shape variant. CG = cortical granules, CV = contractile vacuole. Page 663.
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Table 42 Morphometric data on Pseudouroleptus caudatus caudatus (ca1, type population from Hemberger 1985; ca2, Zanzibar population from Foissner et al. 2002) and Pseudouroleptus caudatus namibiensis (nam, from Foissner et al. 2002) Characteristics a
Subspecies mean
Body, length
Body, width
Body length:body width, ratio
Adoral zone of membranelles, length Anterior body end to distal end of adoral zone, distance Body length: length of adoral zone, ratio
Anterior body end to postperistomial cirrus, distance Anterior body end to buccal cirrus, distance Anterior body end to right marginal row, distance Anterior body end to left frontoventral row, distance Posterior body end to left frontoventral row, distance Anterior body end to right frontoventral row, distance Anterior body end to first macronuclear nodule, distance Nuclear figure, length Anterior macronuclear nodule, length Anterior macronuclear nodule, width Micronuclei, length Micronuclei, width Macronuclear nodules, number
Micronuclei, number
Adoral membranelles, number
ca1 ca2 nam ca1 ca2 nam ca1 ca2 nam ca2 nam nam
M
250.0 – 201.8 200.0 214.3 208.0 50.0 – 47.0 47.5 45.6 45.0 5.0 – 4.3 4.3 4.7 4.7 56.4 55.0 56.3 58.0 17.8 18.0
SD
SE
– 33.3 32.2 – 5.7 3.9 – 0.2 0.7 8.8 6.3 3.8
– 10.5 7.4 – 1.8 0.9 – 0.1 0.2 2.8 1.4 0.9
CV
Min
Max
n
– – – 16.5 150.0 255.0 15.0 167.0 295.0 – – – 12.1 39.0 55.0 8.5 40.0 55.0 – – – 5.1 4.0 4.7 15.0 3.5 6.7 15.6 42.0 75.0 11.1 46.0 71.0 21.2 10.0 25.0
? 10 19 ? 10 19 ? 10 19 10 19 19
ca1 ca2 nam nam
5.0 3.6 3.8 60.6
– 3.6 3.7 58.0
– 0.2 0.6 9.6
– 0.1 0.1 2.2
– 6.3 15.7 15.8
– 3.2 2.7 51.0
– 3.9 6.5 90.0
? 10 19 19
nam nam
22.6 11.1
22.0 11.0
3.1 2.2
0.7 0.5
13.8 19.9
18.0 7.0
30.0 15.0
19 19
nam
20.7
20.0
4.4
1.0
21.3
13.0
30.0
19
nam
20.5
20.0
4.3
1.0
20.8
15.0
27.0
19
nam
86.2
82.0
11.4
2.6
13.2
70.0 115.0
19
nam
54.8
55.0
6.1
1.4
11.1
46.0
71.0
19
nam ca2 nam ca2 nam ca2 nam ca2 nam ca1 ca2 nam ca1 ca2 nam ca1 ca2 nam
61.8 25.1 22.1 8.5 8.6 8.5 6.2 5.2 4.3 2.0 2.0 2.0 – 3.0 2.4 40.0 53.2 43.9
59.0 26.0 22.0 8.0 8.0 8.5 6.0 5.0 4.5 – 2.0 2.0 – 3.0 2.0 – 54.0 45.0
8.5 3.0 2.4 1.4 1.5 1.1 0.5 – – – 0.0 0.0 – 0.7 0.7 – 6.5 4.7
2.0 0.9 0.6 0.5 0.4 0.3 0.1 – – – 0.0 0.0 – 0.2 0.2 – 2.0 1.1
14.0 11.8 10.9 16.9 17.9 12.7 8.6 – – – 0.0 0.0 – 22.2 28.6 – 12.1 10.8
48.0 20.0 18.0 7.0 6.0 7.0 5.0 5.0 3.5 – 2.0 2.0 2.0 2.0 1.0 33.0 40.0 33.0
80.0 30.0 27.0 11.0 12.0 10.0 7.0 6.0 5.0 – 2.0 2.0 3.0 4.0 4.0 50.0 63.0 51.0
19 10 19 10 19 10 19 10 19 ? 19 10 ? 10 19 ? 10 19
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SYSTEMATIC SECTION
Table 42 Continued Characteristics a
Subspecies mean
Frontal cirri, number
Buccal cirri, number
Cirri behind right frontal cirrus, number
Postperistomial cirri, number
Left frontoventral row, number of cirri
Right frontoventral row, number of cirri
Left marginal cirri, number
Right marginal cirri, number
Dorsal kineties, number Caudal cirri, number
ca1 ca2 nam ca1 ca2 nam ca1 ca2 nam ca1 ca2 nam ca1 b ca2 nam ca1 c ca2 nam ca1 ca2 nam ca1 ca2 nam ca1 nam ca1 nam
3.0 3.0 3.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.3 42.0 59.4 56.5 55.0 50.1 25.5 45.0 46.6 51.5 55.0 55.1 55.4 4.0 4.0 3.0 2.4
M
SD
SE
CV
Min
– 3.0 3.0 – 1.0 1.0 – 1.0 1.0 – 1.0 1.0 – 60.0 57.0 – 50.0 26.0 – 47.5 50.0 – 55.0 55.0 – 4.0 – 2.0
– 0.0 0.0 – 0.0 0.0 – 0.0 0.0 – 0.0 – – 8.6 4.9 – 6.7 2.9 – 5.5 5.3 – 7.5 4.8 – 0.0 – –
– 0.0 0.0 – 0.0 0.0 – 0.0 0.0 – 0.0 – – 2.7 1.1 – 2.1 0.7 – 1.7 1.2 – 2.4 1.1 – 0.0 – –
– 0.0 0.0 – 0.0 0.0 – 0.0 0.0 – 0.0 – – 14.4 8.7 – 13.2 11.2 – 11.8 10.2 – 13.7 8.6 – 0.0 – –
– 3.0 3.0 – 1.0 1.0 – 1.0 1.0 – 1.0 1.0 – 40.0 50.0 – 35.0 20.0 – 35.0 44.0 – 43.0 47.0 – 4.0 – 1.0
Max – 3.0 3.0 – 1.0 1.0 – 1.0 1.0 – 1.0 3.0 – 70.0 67.0 – 60.0 31.0 – 53.0 61.0 – 70.0 70.0 – 4.0 – 4.0
n ? 10 19 ? 10 19 ? 10 19 ? 10 19 ? 10 19 ? 10 19 ? 10 19 ? 10 19 ? 19 ? 19
a All measurements in µm. CV = coefficient of variation in %, M = median, Max = maximum value, mean = arithmetic mean, Min = minimum value, n = number of individuals investigated, SD = standard deviation, SE = standard error of arithmetic mean, ? = sample size not known (if only one value is known, it is listed in the mean column; if two values are available then they are listed as Min and Max). Data based on protargol-impregnated specimens. b
Hemberger (1982, 1985) very likely confused the left and right frontoventral row, that is, this value probably refers to the right row.
c
Hemberger (1982, 1985) very likely confused the left and right frontoventral row, that is, this value probably refers to the left row.
Pseudouroleptus caudatus namibiensis Foissner, Agatha & Berger, 2002 (Fig. 137a–c, Table 42) 2002 Pseudouroleptus caudatus namibiensis nov. sspec. – Foissner, Agatha & Berger, Denisia, 5: 652, Fig. 146a–c, Table 127 (Fig. 137a–c; original description of subspecies; the holotype slide [acces-
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669
Fig. 136h, i Pseudouroleptus caudatus caudatus (from Foissner et al. 2002. Protargol impregnation). Infraciliature of ventral and dorsal side and nuclear apparatus, 197 µm. Dorsal kineties and caudal cirri not shown because too indistinct in preparations. Short arrow denotes buccal cirrus at anterior end of undulating membranes, long arrow marks cirrus III/2, which originates from the same anlage as the right frontal cirrus (connected by broken line). The three frontal cirri are enlarged and connected by a dotted line. Arrowhead denotes the postperistomial ventral cirrus, which is very close to the buccal vertex. In Foissner et al. (2002, Fig. 145e; Fig. 136i in present book) the right frontoventral row was incorrectly designated as amphisiellid median cirral row (= left frontoventral row in present book). AZM = distal end of adoral zone of membranelles, LFVR = left frontoventral row, LMR = left marginal row, MA = macronuclear nodule, MI = micronucleus, PF = pharyngeal fibres, RFVR = right frontoventral row (= transverse cirri in Foissner et al. 2002), RMR = right marginal row. Page 663.
sion number 2002/449; Foissner et al. 2002, p. 42] and four paratype slides [2002/452–455] are deposited in the Oberösterreichische Landesmuseum in Linz [LI], Upper Austria).
Nomenclature: The subspecies-group name namibiensis refers to the country (Namibia) where the subspecies was discovered (Foissner et al. 2002).
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SYSTEMATIC SECTION
Fig. 137a–c Pseudouroleptus caudatus namibiensis (from Foissner et al. 2002. Protargol impregnation). a, b: Infraciliature of ventral and dorsal side and nuclear apparatus of holotype specimen, 192 µm. Long arrow marks cirrus III/2 behind right frontal cirrus (connected by broken line), short arrow denotes buccal cirrus, and arrowhead marks postperistomial ventral cirri. Dotted line connects frontal cirri. Note that kinety 3 is distinctly shortened posteriorly, whereas kinety 4 is shortened anteriorly, which is the result of dorsal kinety fragmentation. c: Shape variant in ventral view. AZM = adoral zone of membranelles, CC = caudal cirri, E = endoral, F = fibres extending from adoral membranelles into buccal ca-
Pseudouroleptus
671
Remarks: The present subspecies differs from P. caudatus caudatus only by the distinctly shortened right row of frontoventral cirri. This feature is constant and changes the overall cirral pattern rather distinctly. However, since the difference is only quantitative we classified it as subspecies. The long right frontoventral row in P. caudatus caudatus is now confirmed for three populations (Peru, Brazil, Zanzibar), strongly indicating that the difference is reliable. Pseudouroleptus caudatus namibiensis is easily confused with Hemiamphisiella wilberti (Fig. 60a–g), which has a similar body shape, body size, as well as cirral and nuclear pattern. However, Hemiamphisiella wilberti lacks cortical granules, whereas these organelles are present in both subspecies of P. caudatus (Fig. 136f). Possibly, Hemiamphisiella is a synonym of Pseudouroleptus. However, for a final decision further ontogenetic data and molecular data should be awaited. Perhaps there are differences in the formation of the dorsal kinety pattern. Morphology (Fig. 137a–c, Table 42): This subspecies differs from P. caudatus caudatus by the length of the right frontoventral row. Thus, we restricted the description of P. caudatus namibiensis to the detailed morphometry and figures and a few minor details, namely (i) body size 180–330 × 40–60 µm, that is, slightly larger than nominotypical subspecies; (ii) body never spiralled, a remarkable difference to P. caudatus caudatus and other large soil hypotrichs; (iii) the large, conspicuous micronuclei are frequently ovate; (iv) cortical granule rows as distinct as in P. caudatus caudatus and occasionally the granules lightly impregnate with protargol; (v) the cytoplasm contains, as in P. caudatus caudatus, innumerable minute, protargolaffine granules, making the analysis of the dorsal infraciliature difficult; (vi) buccal cirrus 5–9 µm, on average 6.3 µm, behind anterior end of paroral; (vii) paroral and endoral frequently intersect optically in posterior half; (viii) an early divider shows a long oral primordium near the left frontoventral row and seven cirral anlagen streaks in the opisthe (possible an unusual specimen or one anlage is resorbed later); (ix) late dividers show that the left frontoventral row is composed of three portions, as described for P. caudatus caudatus by Hemberger (1982); (x) the right frontoventral row does not form primordia. Occurrence and ecology: Pseudouroleptus caudatus namibiensis is possibly confined to terrestrial habitats. Type locality is the Aubs Canyon (23°55'S, 16°15'E), Namibia, where we found it with high abundance in a soil sample from the margin of the so-called Riedloch, a small pond (details, see site 30 in Foissner et al. 2002, p. 21). No further records published.
b
vity, FS = frontal scutum, LFVR = left frontoventral row, LMR = anterior end of left marginal row, MA = macronuclear nodule, MI = micronucleus, P = paroral, PF = pharyngeal fibres, RFVR = right frontoventral row (termed transverse cirri in original description), RMR = right marginal row, 1–4 = dorsal kineties. Page 668.
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Ponturostyla Jankowski, 1989 1986 Mixotricha n. g. – Dragesco & Dragesco-Kernéis, Faune tropicale, 26: 439 (original description; no formal diagnosis provided). Type species (by original designation and by monotypy): Paraurostyla enigmatica Dragesco & Dragesco-Kernéis, 1986. 1989 Ponturostyla nom. n. – Jankowski, Vestnik Zoologii, year 1989: 86 (replacement name because of homonymy). Type species (same as for Mixotricha Dragesco & Dragesco-Kernéis, 1986): Paraurostyla enigmatica Dragesco & Dragesco-Kernéis, 1986. 1999 Ponturostyla – Berger, Monographiae biol., 78: 843 (brief note on exclusion from oxytrichids). 2001 Ponturostyla Jankowski 1989 – Aescht, Denisia, 1: 130 (catalogue of generic names of ciliates). 2001 Ponturostyla Jankowski, 1989 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 75 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2001 Ponturostyla Jankowski, 19891 – Song, Europ. J. Protistol., 37: 194 (detailed redescription and morphogenesis of type species and improved diagnosis).
Nomenclature: No derivation of the names are given in the original descriptions. Mixotricha is a composite of the Greek noun he mixis (the mixing, the mixture) and the Greek noun trichos (hair; cirrus in present case) and likely refers to the various cirri (marginal, frontal, transverse, ventral) which are present. Mixotricha Dragesco & Dragesco-Kernéis, 1986 is a junior homonym of Mixotricha Sutherland, 1933, a termite inhabiting flagellate (Devescovinidae, Parabasalia; e. g., Brugerolle & Lee 2002). Thus, Jankowski (1989) replaced the junior homonym by Ponturostyla which is a composite of the Greek noun pont- (the sea; ocean) and the genus-group name Urostyla (see Berger 2006, p. 1041, for derivation) and obviously refers to the fact that the type species lives in saltwater and has as many cirral rows as a Urostyla. Feminine gender (Aescht 2001, p. 295). Characterisation (A = supposed apomorphy): Flexible 18-cirri Oxytrichidae, that is, dorsomarginal kineties present and kinety 3 with multiple fragmentation. Adoral zone formed like a question mark. Undulating membranes roughly in Cyrtohymena pattern. Frontoventral cirri in V-shaped pattern. Postoral ventral cirri in dense cluster behind buccal vertex. Two pretransverse ventral cirri and five transverse cirri. More than one marginal row per side originate from a single, apokinetal anlage (A). Caudal cirri absent. Saltwater. Remarks: See single species. Species included in Ponturostyla (basionym given): (1) Paraurostyla enigmatica Dragesco & Dragesco-Kernéis, 1986.
1 Song (2001) provided the following diagnosis: Oxytrichidae with more than one marginal row on both sides, derived separately from unique group of marginal row primordia on each side; 8 frontal, 5 ventral and 5 transversal cirri generated from 5 frontoventral-transverse cirral anlagen and the UM-primordium; caudal cirri not present; body flexible.
Ponturostyla
673
Single species Ponturostyla enigmatica (Dragesco & Dragesco-Kernéis, 1986) Jankowski, 1989 (Fig. 138a–w, Table 43) 1986 Paraurostyla enigmatica n. sp. – Dragesco & Dragesco-Kernéis, Faune tropicale, 26: 437, Planche 125 C (Fig. 138a; original description; no formal diagnosis provided; site where type slides deposited not mentioned, likely they are in the private collection of Jean Dragesco). 1989 Ponturostyla enigmatica (D. et D.=K., 1986) comb. n. – Jankowski, Vestnik Zoologii 1989: 86 (combination with Ponturostyla). 1999 Paraurostyla enigmatica Dragesco & Dragesco-Kerneís, 1986 – Berger, Monographiae biol., 78: 843 (brief note on exclusion from oxytrichids; see remarks). 2001 Ponturostyla enigmatica (Dragesco & Dragesco-Kerneis 1986) Jankowski 1989 – Aescht, Denisia, 1: 295 (catalogue of generic names of ciliates). 2001 Ponturostyla enigmatica (Dragesco and Dragesco-Kernéis, 1986) Jankowski, 1989 – Berger, Catalogue of ciliate names 1. Hypotrichs, p. 69 (nomenclator containing all basionyms, combinations, and higher taxa of hypotrichs). 2001 Ponturostyla enigmatica (Dragesco & Dragesco-Kernéis, 1986) Jankowski, 19891 – Song, Europ. J. Protistol., 37: 194, Fig. 1–47, Table 1 (Fig. 138b–w; detailed redescription and morphogenesis of type species and improved diagnosis; site where voucher slides deposited not mentioned, likely in the Ocean University of Qingdao, China).
Nomenclature: No derivation of the name is given in the original description. The species-group name enigmatic·us, -a, -um (Greek adjective ainigmatikos [m, f, n]; enigmatic, mysterious) likely refers to the difficult generic assignment. Dragesco & Dragesco-Kernéis (1986) established the new genus Mixotricha for the present species, but did not formerly combine the new genus-group name and the species-group name. In spite of this they have to be considered as combining authors of the name because they established Mixotricha for this species: Mixotricha enigmatica (Dragesco & Dragesco-Kernéis, 1986) Dragesco & Dragesco-Kernéis, 1986. However, note that this name is no longer relevant because Mixotricha was replaced by Ponturostyla because of homonymy. Paraurostyla enigmatica is type species of Mixotricha Dragesco & Dragesco-Kernéis, 1986 and Ponturostyla. Remarks: Dragesco & Dragesco-Kernéis (1986) found this species in a saline pond in Benin and could not assign it unequivocally to a certain genus. Thus, they preliminarily classified it in Paraurostyla Borror, 1972 because P. weissei, type of this genus, has a similar general appearance (for review see Berger 1999, p. 841). They also compared it with Coniculostomum Njine, 1979 and Pleurotricha Stein, 1859, two further oxytrichids (for review see Berger 1999, p. 606, 698), and with 1
Song (2001) provided the following new diagnosis: large marine Ponturostyla in vivo about 130–280 × 50–90 µm in size with 3–7 (usually 4) macronuclear nodules; 5 to 9 marginal rows of cirri on each side; cortical granules in lines on the ventral and irregularly scattered on the dorsal side; single contractile vacuole in mid-body on left; ca. 42–75 adoral membranelles, basically 4 complete and 3–5 shortened dorsal kineties and 2–4 dorsomarginal kineties.
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SYSTEMATIC SECTION
Ponturostyla
675
the urostyloid Pseudourostyla Borror, 1972 (for review, see Berger 2006, p. 750). Simultaneously, they proposed the new genus Mixotricha for the present species because of the unique cirral pattern. Jankowski (1989) found it at the Crimean shore and made Opalblue preparations and micrographs which were, however, very likely never published. In addition, he replaced Mixotricha by Ponturostyla because of homonymy (see nomenclature of genus section). I excluded Ponturostyla enigmatica from the oxytrichids because the dorsal kinety pattern (dorsal kinety fragmentation present or not) was not known at that time (Berger 1999). A few years later, Song (2001) found P. enigmatica in the Yellow Sea and provided a detailed redescription including a description of the cell division. In addition, he transferred Pleurotricha grandis Stein, 1859a to Ponturostyla because it also has more than one left marginal row. I am convinced that this transfer is incorrect because Pleurotricha grandis has a rigid body, lacks cortical granules, and is limnetic (Stein 1859; for review see Berger 1999). Both morphological features strongly indicate that it is a stylonychine oxytrichid and I therefore propose retaining it in Pleurotricha until a detailed redescription is available. Song (2001) accepted the body consistency (flexible or rigid) to redefine Ponturostyla, Pleurotricha, Coniculostomum, and Allotricha. Simultaneously, he put Pleurotricha grandis, a limnetic, rigid species without cortical granules into Ponturostyla, whose type species is marine and has a flexible body and cortical granules. Architricha Gupta, Komra & Sapra, 2006, with A. indica as sole species has three right and two left marginal rows, which, however, divide individually by intrakinetal proliferation (Gupta et al. 2006), indicating that Architricha and Ponturostyla are not closely related. The original description of Ponturostyla enigmatica is not very detailed and the illustration of the cirral pattern seems not very exact (Fig. 138a). By contrast, the redescription by Song (2001) clearly shows that it is an 18-cirri oxytrichid because it has the ordinary, plesiomorphic number of 18 frontal-ventral-transverse cirri, dorsomarginal kineties, and an oxytrichid fragmentation of kinety 3 (Fig. 138j, k, v, w). In spite of the differences in the cirral pattern (Fig. 138a, 138j) I agree with Song (2001) that the two populations are conspecific. The many marginal rows on each side are formed from a single primordium, similarly to in Pseudourostyla (for review, see Berger 2006, p. 750). Thus, Song (2001) assumed “some evolutionary relationship between oxytrichids and urostylids”. However, the same pattern occurs in Allotricha mollis Sterki, 1878, an 18-cirri oxytrichid with more than one right mar-
b
Fig. 138a–i Ponturostyla enigmatica (a, from Dragesco & Dragesco-Kernéis 1986; b–i, from Song 2001. a, protargol impregnation; b–i, from life). a: Infraciliature of ventral side and nuclear apparatus, 180 µm. b: Ventral view of representative specimen, 196 µm. c: Left lateral view. d, e: Ventral and dorsal view showing, inter alia, arrangement of cortical granules. f: Slender specimen at low magnification. Bright areas are the macronuclear nodules. g: Crystals (short arrow) and lipid droplets (long arrow) in cytoplasm. h: Cortical granules (arrow) along cirral rows. i: Resting cyst, about 100 µm across. Arrowheads mark the central portion (possibly nuclear material), arrow denotes cyst wall. CV = contractile vacuole. Page 673.
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Fig. 138j–p Ponturostyla enigmatica (from Song 2001. Protargol impregnation). j, k: Infraciliature of ventral and dorsal side and nuclear apparatus of same specimen, 158 µm. Arrow in (j) marks rearmost frontoventral cirrus (= cirrus IV/3). Asterisks in (k) denote dorsal kinety fragments originating from posterior portion of kinety 3. Arrow in (k) marks leftmost (longest) dorsomarginal kinety. i–n, p: Frontal (l), right marginal (m), left marginal (n), and transverse cirrus with associated fibrils. o: The paroral is composed of zigzagging basal bodies, while the endoral consists of linearly arranged basal bodies. AZM = distal end of adoral zone of membranelles, BC = buccal cirrus, E = endoral, FC = left frontal cirrus, FT = frontoterminal cirri, LMR = outermost left marginal row, MA = macronuclear nodule, MI = micronucleus, P = paroral, PTVC = pretransverse ventral cirri, PVC = postoral ventral cirri, RMR = outermost right marginal row, TC = transverse cirri, III/2 = cirrus behind right frontal cirrus (= paramalar cirrus), 1–3 = dorsal kineties 1–3. Page 673.
Ponturostyla
677
ginal row (Berger 1999, p. 262), indicating that this pattern evolved independently in various higher taxa. Morphology: The original description is rather short and summarised in Table 43. Thus, the following description is based on the detailed investigations by Song (2001), unless otherwise indicated. Body size rather variable in life, that is, 130–280 × 50–90 µm; freshly sampled specimens usually more than 220 µm long, after several days of culture usually less than 200 µm; in one sample specimens were less than 150 µm long and relatively slender; body length:width ratio mostly 2.5–3.0:1. Body outline usually elliptical with both ends broadly rounded and margins more or less straight (Fig. 138b), sometimes fusiform, or broadly oval to elliptical with slightly narrowed posterior end (Fig. 138f). Shape seems to depend on physiological rather than on nutritional state since differences in size and shape occur in the same culture. Body flexible, about 3:1 flattened dorsoventrally (Fig. 138c). Usually four, rarely more, and only one individual with only three macronuclear nodules; nodules arranged, as is usual, left of midline, globular; in life often easily recognisable as bright, round areas (Fig. 138b, f, k). Chromatin bodies small to medium-sized. Micronuclei about 3 µm across, close to macronuclear nodules (Fig. 138k). Contractile vacuole, as is usual, near left cell margin about in mid-body; during diastole obviously with some collecting vacuoles; pulsates at long intervals, usually more than 30-s (Fig. 138b, c, e); contractile vacuole not observed in Benin population (Dragesco & Dragesco-Kernéis 1986). Pellicle thin. Cortical granules colourless or slightly greenish, about 0.8 µm across, aligned in lines between cirral rows on ventral side (Fig. 138d, h), while densely, but irregularly distributed on dorsal side (Fig. 138e); granules recognisable only in life, that is, do not impregnate with the protargol method used. Presence/absence of cortical granules not described for Benin population (Dragesco & Dragesco-Kernéis 1986). Cytoplasm usually dark-grey; typically containing numerous globular lipid-like inclusions 2–4 µm across and about 2–3 µm long crystals (Fig. 138g), so that cells are dark and opaque especially at low magnification (Fig. 138f). Food vacuoles not recognisable. Continuously crawling on substrate or on bottom of Petri dish, reacting quickly, often slightly contracting when disturbed; not sensitive to cover glass pressure. Adoral zone large and conspicuous, occupies almost 40% of body length on average (Table 43), composed of an average of 48 membranelles of ordinary fine structure. Bases of largest membranelles 10–15 µm wide, cilia up to 25 µm long. Proximal portion of zone covered by buccal lip, distal end of zone extends onto right body margin causing a distinct furrow along right-anterior margin (Fig. 138b, d, j). Undulating membranes long and distinctly curved, arranged roughly in Cyrtohymena pattern (Berger 1999). Endoral likely composed of monokinetids, paroral consists of zigzagging basal bodies (Fig. 138j, o). Cirral pattern rather stable, as is usual for 18-cirri hypotrichs (Fig. 138j). Frontal cirri and transverse cirri very strong and up to 30 µm long; other cirri about 20 µm long. Three frontal cirri along distal portion of adoral zone. Buccal cirrus enlarged, somewhat ahead of optical crossing of undulating membranes. Cirrus III/2 about of
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SYSTEMATIC SECTION
Ponturostyla
679
Fig. 138u Ponturostyla enigmatica (from Song 2001. Protargol impregnation). Infraciliature of ventral side of very late divider, 164 µm. Frontal-ventral-transverse cirri which originate from same anlage are connected by broken line (only shown for opisthe). Parental structures white, new black. I, VI = frontal-ventraltransverse cirri anlagen I and VI. Page 673.
same size as buccal cirrus, behind right frontal cirrus. Frontoterminal cirri close behind distal end of adoral zone. Rearmost frontoventral cirrus (= cirrus IV/3) right behind cirrus III/2. Postoral ventral cirri close together behind buccal vertex as in many other flexible 18-cirri hypotrichs. Two pretransverse ventral cirri in ordinary position. Transverse cirri arranged in subterminal, slightly curved pseudorow and thus almost not projecting beyond rear cell end (Fig. 138j). Usually 6–8 marginal rows per side, leaving only a rather narrow field bearing the frontalventral-transverse cirri, which are larger than the marginal cirri. According to Song (2001) on average 20–30 cirri per marginal row, according to Dragesco & Dragesco-Kernéis (1986) about 14–30 cirri. Innermost right marginal cirri more or less distinctly shortened posteriorly; left rows become successively longer anteriorly from inside to outside. Fibrils
b
Fig. 138q–t Ponturostyla enigmatica (from Song 2001. Protargol impregnation). Infraciliature of ventral side of dividing specimens. q: Very early stage, size not indicated. r: Early divider, 185 µm. Arrows mark marginal row primordia. Note that the marginal rows are formed from a single anlage per side. s: Middle divider (170 µm) showing, inter alia, the six frontal-ventral-transverse cirri primordia, the marginal primordia, and the partial reorganisation of the parental adoral membranelles, and the formation of the membranelles in the opisthe from anterior to posterior. t: Late divider, 176 µm. Cirri originating from same anlage are connected by broken line (only shown for opisthe). Parental structures white, new black. OP = oral primordium, I, VI = leftmost and rightmost frontal-ventral-transverse cirri anlage. Page 673.
680
SYSTEMATIC SECTION
Fig. 138v, w Ponturostyla enigmatica (from Song 2001. Protargol impregnation). Infraciliature of dorsal side and nuclear apparatus of middle and late divider, v = 176 µm (same specimen as shown in Fig. 138t), w = 153 µm. Arrows in (v) mark leftmost dorsomarginal kinety. Long arrows in (w) denote rear fragments of kinety 3 of opisthe, short arrow marks a fragmentation of kinety 1 which was observed only once. Asterisks in (w) mark new dorsomarginal rows of proter. AZM = distal end of adoral zone of membranelles, MA = fused macronucleus, 1–3 = dorsal kineties 1–3. Page 673.
associated with cirri well developed (Fig. 138l–n, p). Marginal rows more or less confluent posteriorly (Fig. 138j). Dorsal bristles about 2–3 µm long, basically arranged as shown in Fig. 138k; number of rows difficult to count because of multiple fragmentation of kinety 3 and variable (2–3) number of dorsomarginal kineties. Rear portion of kinety 3 fragments into 2–4 rows. Short dorsomarginal kineties with about 3–8 basal body pairs. Caudal cirri lacking. Resting cyst (Fig. 138i): According to Song (2001) about 100 µm across, surface covered by hyaline alveolar structures which are often irregularly bubble-like and 3–10 µm in size. Wall dark grey and opaque, about 10–20 µm thick. Central area invariably with a big mass with several lipid-like granules, which possibly represent the nuclear material. Cell division (Fig. 138q–w): Song (2001) studied this part of the life cycle in great detail. In the present review only the most important stages are shown and the major events are described. For a more comprehensive description and documenta-
Ponturostyla
681
Table 43 Morphometric data on Ponturostyla enigmatica (en1, from Dragesco & Dragesco-Kernéis 1986; en2, from Song 2001) Characteristics a
Population mean
Body, length Body, width Adoral zone of membranelles, length Macronuclear nodules, width Adoral membranelles, number Macronuclear nodules, number Micronuclei, number Frontal cirri, number Ventral cirri, number Transverse cirri, number Right marginal rows, number Innermost right marginal row, number of cirri Left marginal rows, number Innermost left marginal row, number of cirri Dorsal kineties, number
M
SD
SE
CV
Min
Max
n
en1 en2 en1 en2 en1 en2 en2 en1 en2 en1 en2 en2 en1 b en2 c en1 en2 d en1 en2 en1 en2 en2
– 171.9 – 73.1 – 65.7 13.7 – 48.3 6.0 4.2 4.0 5.0 8.0 – 5.1 6.0 5.0 – 6.6 24.6
– – – – – – – – – – – – – – – – – – – – –
– 27.6 – 19.8 – 12.4 4.7 – 17.5 – 0.6 1.0 – 0.0 – 0.3 – 0.0 – 1.1 4.0
– 6.0 – 4.7 – 2.9 1.2 – 5.9 – 0.1 0.3 – 0.0 – 0.1 – 0.0 – 0.3 1.1
– 140.0 200.0 16.0 104.0 236.0 – 60.0 105.0 27.1 46.0 112.0 – 80.0 95.0 18.9 44.0 84.0 34.5 9.0 14.0 – 70.0 76.0 36.3 42.0 61.0 – – – 15.0 3.0 7.0 23.8 3.0 6.0 – – – 0.0 8.0 8.0 – 4.0 12.0 5.9 5.0 6.0 – – – 0.0 5.0 5.0 – 6.0 7.0 17.1 5.0 9.0 16.1 18.0 31.0
? 21 ? 18 ? 18 15 ? 18 ? 21 12 ? 17 ? 21 ? 21 ? 21 12
en1 en2 en2
– 7.6 21.6
– – –
– 1.0 4.3
– 0.2 1.2
– 13.6 19.8
9.0 6.0 15.0
10.0 9.0 28.0
? 21 13
en1 en2
– 6.7
– –
– 0.6
– 0.1
– 8.3
5.0 5.0
6.0 7.0
? 21
a All measurements in µm. CV = coefficient of variation in %, M = median, Max = maximum value, mean = arithmetic mean, Min = minimum value, n = number of individuals investigated, SD = standard deviation, SE = standard error of arithmetic mean, ? = sample size not indicated. Data based on protargolimpregnated specimens. b
Buccal cirrus and cirrus III/2 included.
c
Comprising frontal cirri, buccal cirrus, and frontoventral cirri.
d
Comprising postoral and pretransverse ventral cirri.
tion, see Song (2001). Briefly, the ontogenesis is basically as in many other 18-cirri oxytrichids. Stomatogenesis: The oral primordium of the opisthe is formed in the postoral area, obviously without contact to the transverse cirri. However, the postoral ventral cirri are modified to anlagen (Fig. 138q). The membranelles of the opisthe are formed, as is usual, from anterior to posterior. The parental adoral zone is retained for the proter; as is usual, an internal, inconspicuous reorganisation occurs (Fig. 138s).
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SYSTEMATIC SECTION
The parental undulating membranes are completely reorganised and form, also as is usual, the new anlage I for the proter (Fig. 138t). Formation of frontal-ventral-transverse cirri: The anlagen for the proter and opisthe originate independently from each other (Fig. 138r). The exact origin of all anlagen could not be detected. As is usual, the parental undulating membranes, the buccal cirrus, and cirri III/2 and IV/3 are involved in anlagen formation of the proter, while the anlagen of the opisthe are obviously formed from the oral primordium and the postoral ventral cirri. As is usual, the six (I–VI) anlagen form the following cirri both in proter and opisthe (Fig. 138r–u): anlage I forms left frontal cirrus (I/1) and undulating membranes; anlage II forms left transverse cirrus (II/1), buccal cirrus (II/2), and middle frontal cirrus (II/3); anlage III forms transverse cirrus III/1, cirrus behind right frontal cirrus (= paramalar cirrus or cirrus III/2), and right frontal cirrus (III/3); anlage IV forms transverse cirrus IV/1, the anteriormost postoral ventral cirrus (IV/2), and the rearmost frontoventral cirrus (IV/3); anlage V forms transverse cirrus V/1, left (= anteriormost) pretransverse ventral cirrus (V/2), and the middle and rearmost postoral ventral cirri (V/3, V/4); anlage VI forms the rightmost transverse cirrus (VI/1), the right pretransverse ventral cirrus (VI/2), and the two frontoterminal cirri (= VI/3, VI/4, that is, the anteriormost two frontoventral cirri. Marginal row formation: Ponturostyla enigmatica has many marginal rows on both sides (Fig. 138r–u). According to Song (2001) the marginal row anlagen occur de novo, that is, without contact to parental structures. However, according to Fig. 138r, s it is obviously not possible to exclude the involvement of some parental marginal cirri. The marginal rows of each side originate from a single anlage at the rightmost parental row(s). Such a pattern is known, inter alia, from the urostyloid Pseudourostyla (for review, see Berger 2006) and the oxytrichid Allotricha (Berger 1999), indicating that such a mode evolved several times independently. Dorsal kinety formation: Within kineties 1–3 one anlage each is formed for the proter and the opisthe (Fig. 138v, w). Kinety 3 shows a multiple fragmentation, demonstrating that Ponturostyla is an oxytrichid. Very rarely, a fragmentation occurs also in kinety 1 (Fig. 138w). 2–3 dorsomarginal kineties originate from/near the primordium which forms the right marginal rows. No caudal cirri occur. Division of nuclear apparatus: This process proceeds in the plesiomorphic manner, that is, a replication band traverses the macronuclear nodules (Fig. 29 in Song 2001), which later fuse to a single mass and divide again in late dividers. The micronuclei divide mitotically (Fig. 138v, w). Occurrence and ecology: Ponturostyla enigmatica is likely confined to saline waters. Type locality are saline ponds in the city of Cotonou, Benin (Dragesco & Dragesco-Kernéis 1986). Song (2001) found the Chinese population in a sample (21–22 °C, pH 8.0, salinity 32–36‰) from the coastal water off Qingdao. He cultured it at room temperature in boiled seawater (salinity about 32‰) with rice grains to support bacterial growth. I found a small population, which perfectly matched the Chines population, in the northern Adriatic Sea at the city of Lignano, Italy. Food not known.
Addenda The data by Li et al. (2007) about Amphisiella marioni, by Shao et al. (2007) about Trachelostyla pediculiformis, and by Gong et al. (2007) about Protogastrostyla pulchra became available too late for inclusion in the main text. Thus, the major results of these publications are added here at the end of the book.
Amphisiella capitata (Pereyaslawzewa, 1886) Borror, 1972 (see p. 91; Fig. 139a–h, Table 44) 2007 Amphisiella marioni Gourret & Roeser, 18881 – Li, Lin, Shao, Gong, Hu & Song, J. Euk. Microbiol., 54: 364, Fig. 1–23, Tables 1, 2 (Fig. 139a–h; detailed redescription and neotypification; the neotype slide [accession number Lin-04-04-21] is deposited in the Laboratory of Protozoology, Ocean University of China, Qingdao, China).
Nomenclature: Li et al. (2007) fixed a neotype for A. marioni because of the incomplete data and the difficult taxonomy (details see remarks). Remarks: Li et al. (2007) compared A. marioni with A. annulata and A. capitata. They considered all three species as valid whereas I classify, like Hemberger (1982), Amphisiella marioni as junior synonym of A. capitata (details see p. 91ff). Li et al. (2007) distinguished A. capitata and A. marioni, inter alia, by the position of the distal end of the adoral zone. Accordingly, the DE-values2 are rather different, namely, 0.50 against 0.25. However, this value is very difficult to estimate from the illustrations provided by Pereyaslawzewa (1886; Fig. 16a) and Gourret & Roeser (1888; Fig. 16b) and thus should not be over-interpreted. Moreover, the oral apparatus of the specimen shown in Fig. 18 in Li et al. (2007) agrees much better with that of A. capitata (Fig. 16a in present book) than with that of A. marioni (Fig. 16b). Li et al. (2007) emphasised that the adoral zone of their neotype is bipartite with the anterior portion on the ventral side, whereas it is continuous and the anterior portion on the dorsal side of the frontal scutum in A. annulata. Indeed the situation is very similar to that illustrated by Gourret & Roeser (1888). However, details of this meticulous feature can only be seen in good scanning electron micrographs. The position of the contractile vacuole is quoted as second difference by Li et al. (2007). In A. marioni the vacuole is, according to Gourret & Roeser (1888), usually lacking. When it is present, then it is in the posterior dorsal portion. This position 1 Li et al. (2007) provided the following improved diagnosis: Marine Amphisiella with elongated body shape, size about 90–150 × 25–30 µm in vivo; grouped cortical granules, arranged in irregular rows throughout whole body; contractile vacuole located slightly behind the equator; adoral zone with 25–36 membranelles, slightly bipartite in structure with the anterior part ventrally located; about eight frontal cirri left of ACR (amphisiellid median cirral row); ACR with 31–45 cirri, extending to the level of the TC (transverse cirri); closely spaced marginal rows, with 28–42 and 30–43 cirri in the right and left rows, respectively; six TC arranged V-shaped; typically six dorsal kineties; and usually two macronuclear nodules. 2 For explanation of this quotient, see Berger (2006, p. 18).
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agrees with that observed in the neotype population and could be indeed a difference to A. capitata where Pereyaslawzewa (1886) illustrated a vesicle left of the proximal portion of the adoral zone (Fig. 16a). However, it is generally known that the contractile vacuole is a difficult feature in marine hypotrichs. The cirral pattern described by Li et al. (2007) for A. marioni agrees more or less perfectly with that observed by Pereyaslawzewa (1886) in A. capitata (Fig. 16a, 139g). But this does not mean a lot because the infraciliature of Amphisiella species is very similar. All these indistinct differences between A. capitata and A. marioni (preliminary) cause me to follow Hemberger (1982) and to consider them as synonyms. Only studies from the type locality of A. capitata (Black Sea) will show whether the sophisticated differences (mainly the position of the contractile vacuole) between A. capitata and A. marioni can be confirmed or not. Li et al. (2007) discussed the Amphisiella marioni-populations described by Mansfeld (1923; Fig. 16c–e) and Wicklow (1982; Fig. 16m). They inferred from a comparison with the original description of A. marioni and with their own results that both populations are misidentifications. However, they made no comments about the identity of the populations studied by Mansfeld (1923) and Wicklow (1982). Thus, more A. capitata/A. marioni-like populations have to be studied to get a better idea about the taxonomy of this species or species-group. Morphology: Li et al. (2007) described the neotype population of A. marioni in great detail. The population is documented by several micrographs showing live and protargol-impregnated specimens (Fig. 9–23 in Li et al. 2007). Body length usually 100–140 µm in life; length:width ratio 4–5:1. Body outline elongate, anterior end broadly rounded, posterior slightly tapered. Body flexible and only slightly contractile, dorsoventrally flattened 2:1; ventral side flat with three clearly visible grooves along marginal rows and amphisiellid median cirral row (Fig. 139a–d). Usually two macronuclear nodules in central body portion; individual nodules ellipsoidal and 14 × 8 µm in life on average, contain many spherical chromatin bodies. Micronuclei not observed. Full contractile vacuole about 10 µm across, near left cell margin and slightly behind mid-body (about at 56% of body length in specimen illustrated; Fig. 139a). Cytoplasm colourless to greyish, often with many shin-
b
Fig. 139a–h Amphisiella capitata (neotype population of synonym A. marioni; from Li et al. 2007; a–f, from life; g, h, protargol impregnation). a–c: Ventral views of stretched and curved specimens showing great flexibility of cell, a = 128 µm. Note that the distal portion of the adoral zone is obviously on the ventral side (cp. Fig. 16b). d: Left lateral view showing dorsoventral flattening, dorsal bristles, and macronuclear nodules. e: Dorsal view showing contractile vacuole, shape of adoral zone, and cortical granulation, 117 µm. f: Cortical granules (about 0.5 µm across) and mitochondria-like structures (3.0 × 1.5 µm). g, h: Infraciliature of ventral and dorsal side and macronucleus of neotype specimen, 120 µm. Short arrow marks transition of proximal to distal portion of adoral zone; arrowhead denotes buccal cirrus ahead of undulating membranes, and long arrow marks supposed pretransverse ventral cirrus VI/2. Frontal cirri connected by dotted line; right frontal cirrus (III/3) and cirrus III/2 connected by broken line. ACR = amphisiellid median cirral row, AZM = distal end of adoral zone, CG = cortical granules, CV = contractile vacuole, LMR = left marginal row, MA = rear macronuclear nodule, MO = mitochondria-like structures, P = paroral, RMR = right marginal row, 1–6 = dorsal kineties. Page 683.
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Table 44 Morphometric data on the neotype population of Amphisiella marioni, which is preliminary classified as junior synonym of A. capitata in present review (from Li et al. 2007) Characteristics a
mean
M
SD
SE
Body, length Body, width Macronuclear nodule, length Macronuclear nodule, width Macronuclear nodules, number Adoral zone of membranelles, length Adoral membranelles, number Cirri on frontal area, number b Amphisiellid median cirral row, number of cirri Transverse cirri, number c Left marginal cirri, number Right marginal cirri, number Dorsal kineties, number
108.3 32.2 21.6 10.9 2.0 44.1 31.3 8.0 36.2 6.0 32.4 32.1 6.0
– – – – – – – – – – – – –
13.7 8.2 7.7 3.0 0.5 6.2 3.5 0.5 3.9 0.0 4.3 4.2 0.5
– – – – – – – – – – – – –
CV
Min
Max
n
12.7 25.4 26.5 27.7 24.0 14.1 11.0 6.6 10.8 0.0 13.2 13.1 7.5
84.0 140.0 24.0 52.0 9.0 32.0 6.0 16.0 2.0 4.0 33.0 60.0 25.0 36.0 7.0 9.0 31.0 45.0 6.0 6.0 30.0 43.0 28.0 42.0 5.0 6.0
24 24 24 24 24 24 24 22 24 22 24 24 5
a
All measurements in µm. CV = coefficient of variation in %, M = median, Max = maximum value, mean = arithmetic mean, Min = minimum value, n = number of individuals investigated, SD = standard deviation, SE = standard error of arithmetic mean. Data based on protargol-impregnated specimens. b
Includes three frontal cirri, buccal cirrus, cirrus behind right frontal cirrus, and cirri left of anterior portion of amphisiellid median cirral row.
c
Pretransverse ventral cirrus (Fig. 139g, arrow) obviously included.
ing globules 3–5 µm across. Pellicle (cortex?) comparatively thick with dot-like cortical granules; granules about 0.5 µm across, dark red, basically irregularly arranged in longitudinal rows distributed throughout the cell. Very many mitochondria-like structures (about 3.0 × 1.5 µm) just beneath cell surface, conspicuous even at low magnification (Fig. 139e, f). Movement moderately rapid; as is usual, benthic, that is, specimens creeping on debris or bottom of Petri dish frequently making short pauses and then changing direction. Adoral zone occupies 30–40% of body length, composed of 25–36, on average 31 membranelles; 8–10 anterior (distal) membranelles on ventral side (usually they are on the dorsal side of the frontal scutum), cilia about 10–12 µm long; posterior (proximal) portion composed of 23–27 membranelles1 with cilia about 8 µm long; no distinct gap between distal and proximal portion of adoral zone. Paroral and endoral straight and parallel, commence behind buccal cirrus; paroral composed of zigzagging basal bodies, endoral single-rowed (Fig. 139g). Pharyngeal fibres obviously inconspicuous, because neither illustrated nor clearly recognisable in micrographs. Cirral pattern and number of cirri of usual variability (Fig. 139a, g, Table 44). Three frontal cirri rather obliquely arranged; right cirrus, as is usual, behind distal 1 The morphometry is somewhat misleading because the sum of the lowest number of distal membranelles (8) plus proximal membranelles (23) is distinctly higher than the lowest total number (25) of membranelles.
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end of adoral zone; middle cirrus slightly ahead of level of buccal cirrus; frontal cirri of about of same size as remaining cirri. Buccal cirrus ahead of undulating membranes. Usually one cirrus (= cirrus III/2) behind right frontal cirrus and three cirri forming short row left of anterior portion of amphisiellid median cirral row. Amphisiellid median cirral row sigmoidal, commences about at level of cirrus III/2, composed of 36 cirri on average, terminates immediately ahead of transverse cirri; cirri of middle portion larger (wider) and more narrowly spaced than in anterior and posterior portion, exactly as in A. annulata. Invariable six transverse cirri1 form Vshaped pattern, difficult to distinguish from marginal cirri; cirri about 10 µm long in life and thus distinctly projecting beyond rear cell body end. Right marginal row commences about at 30% of body length, ends – like left row – subterminal; left row begins left of proximal portion of adoral zone about at level of cirrus III/2; marginal rows distinctly displaced inwards (Fig. 139a, b, g). Dorsal bristles about 3 µm long in life, widely spaced, and usually arranged in six rows. Caudal cirri lacking (Fig. 139h). Occurrence and ecology: Due to the neotypification, the new type locality of A. marioni is the site were the neotype population was collected, namely the outlet of a mariculturing water system in Qingda (Tsingtao; 36°08'N, 120°43'E), China (Li et al. 2007). The sample was collected at following conditions: 25°C water temperature; 31‰ salinity; pH 7.9. Li et al. (2007) established raw cultures at room temperature using Petri dishes filled with seawater from the sample site and some rice grains to support microbial growth. After several days, a low number of specimens occurred. Attempts to establish a pure culture failed (Li et al. 2007).
Trachelostyla pediculiformis (Cohn, 1866) Borror, 1972 (see p. 478; Fig. 140a, b) 2007 Trachelostyla pediculiformis (Cohn, 1866) Borror, 1972 – Shao, Song, Yi, Gong, Li & Lin, Europ. J. Protistol., 43: 255, 256, Fig. 1A–J, 2A–H, 3A, B, 4A–L, 5A–J, 6 (detailed description of cell division and estimation of phylogenetic position; voucher slides likely deposited in the Laboratory of Protozoology, Ocean University of China, Qingdao, China).
Remarks: Shao et al. (2007) described the cell division of the type species of Trachelostyla. Details see next chapter. Cell division: For a detailed description of this process, including a documentation with 18 illustrations and 22 micrographs, see Shao et al. (2007). I only provide the most important features: (i) The oral primordium for the proter originates, without contact to the parental oral apparatus, right of the proximal portion of the adoral zone. From the right side of this primordium the anlagen for the undulating membranes and the frontalventral-transverse cirri are formed. The oral primordium for the opisthe occurs si1
I suppose that the cirrus marked with an arrow in Fig. 139g is the pretransverse ventral cirrus VI/2 (cp. Fig. 2a, 3a).
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Fig. 140a, b Scheme of dorsal kinety pattern and origin of individual kineties in Trachelostyla and oxytrichids (original; a, reconstructed from data provided by Shao et al. 2007; b, reflects the situation in many oxytrichids). Both taxa form six dorsal kineties and three caudal cirri. However, the formation is rather different so that one can exclude a close relationship of Trachelostyla and the oxytrichids. A major difference is the lack of dorsomarginal kineties (interphasic kineties 5 and 6 in b) in Trachelostyla. RMR = right marginal row. Further details, see text. Page 687.
multaneously at about 66% of body length. The parental adoral zone is completely replaced, which is reminiscent of the pseudokeronopsids (for review, see Berger 2006). (ii) The 18 frontal-ventral-transverse cirri are formed, as in many other hypotrichs, from the plesiomorphic number of six (I–VI) anlagen (Fig. 4a). From the de-
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scription by Shao et al. (2007) it is not possible to estimate the exact allocation of the postoral ventral cirri (Fig. 4a, b). (iii) The marginal rows are formed in the plesiomorphic manner, that is, each two anlagen occur within the parental rows. (iv) The macronuclear nodules fuse to a single mass. The replication bands were observed in almost all non-diving specimens and in early dividers. (v) The dorsal kinety formation of T. pediculiformis is highly interesting (Fig. 140a). This species has almost invariable six dorsal kineties. In early dividers each two primordia (one for the proter, one for the opisthe) occur in kineties 1 and 6. In the next stage described by Shao et al. (2007), each two primordia are arranged in parallel in kinety 6, that is, two for the proter and two for the opisthe; both anlagen per filial product form a caudal cirrus! In middle or late dividers, the primordia in kinety 1 fragment and form four kineties (new kineties 1–4) per filial product; the rightmost fragment forms a caudal cirrus. The parental kineties 2–5 are obviously not involved in primordia formation and become resorbed in later stages. Consequently, each filial product has six new dorsal kineties, namely, four from parental kinety 1 and two from kinety 6 (Fig. 140a). It is not known at which level this curious dorsal kinety formation is an apomorphy because the cell division of no other trachelostylid is described. Trachelostyla caudata has, according to live observations by Kahl (1932), 8–10 kineties whereas Spirotrachelostyla tani has only two kineties (Table 31). This high variability in the number of kineties indicates that the formation pattern is not very uniform within the tracheostylids. The Trachelostyla pediculiformis pattern of dorsal kinety formation is rather different from the Oxytricha-pattern, which also creates six dorsal kineties (Berger & Foissner 1997; Fig. 24a in Berger 1999; Fig. 140a, b). In the Oxytricha-pattern, which is rather common within the oxytrichids, kineties 1 and 2 form each one anlage per filial product, kinety 3 fragments and forms the new kineties 3 and 4, and kineties 5 and 6 are dorsomarginal rows originating from/near the right marginal row (Fig. 140b). Caudal cirri are formed on kineties 1, 2, and 4. These distinct differences indicate that the two patterns evolved independently from each other. The phylogenetic position of Trachelostyla is not yet known in detail. The 18cirri pattern is without value for the estimation because it is (very likely) an apomorphy for the Hypotricha and therefore a plesiomorphy within this group (Fig. 6a, square 9). The (multiple) dorsal kinety fragmentation in Trachelostyla proceeds basically in the same way as in the oxytrichids (e.g., Oxytricha granulifera; Foissner & Adam 1983, for review see Berger 1999). However, in the oxytrichids the fragmentation occurs primarily in kinety 3 whereas in Trachelostyla kinety 1 is involved (Fig. 140a, b). In addition, Trachelostyla lacks dorsomarginal kineties (vs. present in oxytrichids), but forms kineties 5 and 6 from the parental kinety 6. Especially the lack of dorsomarginal kineties indicates that Trachelostyla does not belong to the Dorsomarginalia, that is, branched off rather basally in the Hypotricha tree (Fig. 9a).
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This “morphological” position agrees in the main with the locations inferred from molecular data (Schmidt et al. 2007, Shao et al. 2007, Gong et al. 2007). The estimation of the next relative of Trachelostyla is rather difficult. The molecular data propose different candidates. According to Schmidt et al. (2007; their Fig. 2), Trachelostyla pediculiformis forms the most basal branch of the Hypotricha. In the tree published by Shao et al. (2007), Trachelostyla groups with a clade composed of Gonostomum strenuum, G. namibiense, Orthoamphisiella breviseries, and Hemiurosoma terricola (p. 619; dorsomarginal kinety present!). By contrast, Gong et al. (2007) found Parabirojimia similis (for review see Berger 2006) as sister group; the next relatives are Amphisiella annulata (p. 100) and Hemigastrostyla enigmatica (for review, see Berger 1999).
Protogastrostyla Gong, Kim, Kim, Min, Roberts, Warren & Choi, 2007 2007 Protogastrostyla n. g.1 – Gong, Kim, Kim, Min, Roberts, Warren & Choi, J. Euk. Microbiol., 54: 477 (original description). Type species (by original designation on p. 477): Stilonichia pulchra Pereyaslawzewa, 1886. 2008 Maregastrostyla gen. nov. – Berger, present publication, p. 136 (original description). Type species (by original designation): Stilonichia pulchra Pereyaslawzewa, 1886 (junior objective synonym; for details, see nomenclature).
Nomenclature: As mentioned above, the data by Gong et al. (2007) became available too late to be included in the main text and to avoid the publication of this synonymy. However, the situation is rather simple: Maregastrostyla Berger, 2008 (p. 136) has the same type species as Protogastrostyla Gong, Kim, Kim, Min, Roberts, Warren & Choi, 2007 and therefore Maregastrostyla is the junior, objective synonym of Protogastrostyla. Protogastrostyla is a composite of the Greek prefix proto (first, primitive) and the generic name Gastrostyla, referring to the early branch-off of this taxon in the phylogenetic tree (Gong et al. 2007). For derivation of Gastrostyla, see Maregastrostyla. Like Gastrostyla of feminine gender (Aescht 2001, p. 283). Remarks: Gong et al. (2007) studied three Korean populations, two from Jawol Island and one from Ganghwa. The illustrations of the life specimens show distinct differences, for example, in the position of the transverse (distinctly displaced ante1 Gong et al. (2007) provided the following diagnosis: Adoral zone of membranelles formed like a question mark. Undulating membranes in Oxytricha pattern. More than seven frontoventral and post-oral cirri; frontoventral cirri in more or less continuous row. Five transverse cirri and two pre-transverse cirri. One right and one left row of marginal cirri. Caudal cirri present. During cell division the apical part of the old adoral zone of membranelles is retained combining with the newly built membranelles that develop from the proter’s oral primordium; the dorsal kinety’s primordia are of the primary type (a primordium that first appears within the middle part of each kinety, and then divides into two primordia for the proter and the opisthe, respectively); and marginal primordia commence de novo without definite contribution from the parental structure.
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Table 45 Morphometric data on Protogastrostyla pulchra (from Gong et al. 2007) a Characteristics b
mean
M
SD
SE
CV
Min
Max
n
Body, length Body, width Macronuclear nodule, length Macronuclear nodule, width Macronuclear nodules, number Micronuclei, number Transverse cirri e to rear cell end, distance Adoral zone of membranelles, length Adoral membranelles, number Paroral, length Frontal-ventral cirri, number c Transverse cirri, number Left marginal cirri, number d Right marginal cirri, number Dorsal kineties, number
119.1 48.4 25.3 15.9 2.1 2.7 21.6 46.3 35.9 33.2 16.0 5.0 29.8 26.2 5.0
– – – – – – – – – – – – – – –
18.8 9.5 3.4 3.0 0.4 0.8 5.4 6.2 4.7 4.5 0.0 0.0 3.6 3.4 0.2
3.6 1.8 0.7 0.6 0.1 0.2 1.2 1.2 0.9 0.9 0.0 0.0 0.8 0.8 0.0
15.8 19.7 13.4 19.0 18.6 30.0 24.9 13.4 9.6 13.6 0.0 0.0 12.0 12.9 4.2
80.0 160.0 35.0 68.0 19.0 32.0 11.0 24.0 2.0 4.0 2.0 4.0 14.0 37.0 35.0 58.0 30.0 42.0 25.0 41.0 16.0 16.0 5.0 5.0 21.0 39.0 20.0 36.0 4.0 5.0
27 27 24 24 26 20 22 28 27 27 27 27 25 18 23
a
Unfortunately, Gong et al. (2007) did not indicate to which population (Jawol Island or Ganghwa) the data refer.
b
All measurements in µm. CV = coefficient of variation in %, M = median, Max = maximum value, mean = arithmetic mean, Min = minimum value, n = number of individuals investigated, SD = standard deviation, SE = standard error of arithmetic mean. Data based on protargol-impregnated specimens. c
Comprises invariably three frontal cirri, one buccal cirrus, cirrus III/2, nine cirri forming the amphisiellid median cirral row, and two pretransverse ventral cirri.
d
Caudal cirri (usually three in number) included.
e
Likely the rearmost transverse cirrus (V/1) is meant.
riad and not protruding beyond rear cell end [Fig. 141a] vs. only slightly subterminal and therefore distinctly projecting beyond rear cell end [Fig. 141b]). In addition, the cirri of the amphisiellid median cirral row are linearly arranged (Fig. 141a, c) against irregularly (Fig. 141b, f). Interestingly, the population from Ganghwa has a large food vacuole in the posterior body portion (Fig. 141a, c); Gourret & Roeser (1888) and Jones (1974) obviously misinterpreted this organelle as contractile vacuole (Fig. 25a). The morphological differences indicate that subspecies exist. This is supported by the 18S rDNA data (Fig. 49 in Gong et al. 2007). Gong et al. (2007, p. 477) classified Protogastrostyla as incertae sedis in the “subclass Stichotrichia” (= Hypotricha in my terminology). They correctly stated that Foissner et al. (2002) overlooked the rather simple mode of dorsal kinety formation in P. pulchra described by Hu & Song (2000). According to the maximum likelihood tree based on the 18S rDNA, Protogastrostyla is the sistergroup to all other hypotrichs (Gonostomum to Pattersoniella in Fig. 49 of Gong et al. 2007). By contrast, in the tree based on 28S rDNA and in a combined tree (18S + 28S), Protogastrostyla branches off in a more central position (Holosticha + (Hemigastrostyla + (Protogastrostyla + (Uroleptus to Oxytricha))). This clearly shows that the exact po-
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Fig. 141a–g Protogastrostyla pulchra (from Gong et al. 2007. a–e, from life; f, g, protargol impregnation). a: Ventral view of representative specimen of Ganghwa population (brackish water), 122 µm. b: Ventral view of representative specimen of Jawol population (marine), 111 µm. Note that in this illustration the transverse cirri are only slightly subterminal which does not agree very well with the remaining descriptions. c–e: Arrangement of cortical granules on ventral (c) and dorsal side (d) and detail (e). Asterisk marks large food vacuole in posterior body portion. f, g: Infraciliature of ventral and dorsal side and nuclear apparatus of same(?) specimen, size not indicated. TC = transverse cirri, 1–5 = dorsal kineties. Page 693.
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sition of Protogastrostyla within the Hypotricha is not yet known, but the data basically confirms the idea that it is, like other amphisiellids, a non-dorsomarginalian hypotrich (Fig. 9a). However, in the molecular tree Protogastrostyla does not cluster with Amphisiella annulata (Fig. 49 in Gong et al. 2007).
Protogastrostyla pulchra (Pereyaslawzewa, 1886) Gong, Kim, Kim, Min, Roberts, Warren & Choi, 2007 (see p. 137; Fig. 141a–g, Table 45) 2007 Gastrostyla pulchra (Perejaslawzewa 1886) Kahl, 1932 – Gong, Kim, Kim, Min, Roberts, Warren & Choi, J. Euk. Microbiol., 54: 469, Fig. 1–27, Table 1 (Fig. 141a–g; redescription and estimation of phylogenetic position using molecular data; see nomenclature). 2007 Protogastrostyla pulchra (Perjaslawzewa 1886) n. comb. – Gong, Kim, Kim, Min, Roberts, Warren & Choi, J. Euk. Microbiol., 54: 477 (combination with Protogastrostyla). 2008 Maregastrostyla pulchra (Pereyaslawzewa, 1886) comb. nov. – Berger, present publication, p. 137 (combination with Maregastrostyla; junior, objective synonym of Protogastrostyla pulchra).
Nomenclature: Type species of Protogastrostyla. For explanation of synonymy of Protogastrostyla Gong et al., 2007 and Maregastrostyla Berger, 2008 (p. 136) see genus Protogastrostyla. Remarks: See genus section. Morphology: In the present chapter, the Korean populations studied in detail by Gong et al. (2007) are described. The description is obviously a combination of all three populations; unfortunately, the Korean authors did not indicate from which population the morphometric data are (Table 45). For a detailed documentation with further illustrations and 16 micrographs, see Gong et al. (2007). Further details about Protogastrostyla pulchra, see Berger (1999, p. 818) and present review (p. 137). Body size about 110–180 × 30–50 µm in life. Body non-contractile, but flexible, dorso-ventrally flattened 2:1. Body outline variable, usually slender and elongate in brackish populations (Fig. 141a, c, d), but stout and distinctly cephalised in marine populations (Fig. 141b); both ends usually widely rounded, but anterior portion slightly tapering, especially in estuarine specimens; margins slightly convex to more or less straight. Two (rarely four) ellipsoidal macronuclear nodules about in midbody slightly left of midline; individual nodules about 25 × 16 µm, chromatin bodies of ordinary size. 2–4 micronuclei, about 2.5 µm across, close to macronuclear nodules (Fig. 141a, b, g). No contractile vacuole observed. Cortical granules (extrusomes) colourless, elliptical (1.5 × 0.5 µm) in lateral view, perpendicularly arranged to pellicle in life and irregularly grouped between cirral rows and in nine rows on dorsal side (Fig. 141c–e). Cytoplasm colourless to slightly greyish. Commonly one large food vacuole in posterior body portion, contains flagellates and bacteria (Fig. 141a, c), but usually it feeds on diatoms. Movement moderately rapid, benthic.
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ADDENDA
Adoral zone about 25–33% of body length, composed of 30–42 membranelles; bases1 of membranelles 11–16 µm long in life. Distal end of adoral zone extends far posteriorly (DE-value of specimen shown in Fig. 141f is 0.56). Undulating membranes in Oxytricha-pattern, that is, more or less distinctly intersecting in top view (Fig. 141a, b, f); cilia 6–7 µm long. Invariably (n = 27) 16 frontal-ventral cirri, namely three frontal cirri, one buccal cirrus, one cirrus (= cirrus III/2) left of anterior portion of amphisiellid median cirral row which is composed of nine rather irregularly arranged cirri, and two pretransverse ventral cirri (Fig. 141f); specimens shown in Fig. 141a, c with a distinctly leftwards shifted postperistomial cirrus. Transverse cirri distinctly displaced anteriad2 and thus not projecting beyond rear cell end although 20–25 µm long (Fig. 141a, f). Right marginal row commences about at level of cirrus III/2, terminates at rear cell end. Left marginal row begins left of proximal end of adoral zone, extends to near rear end of right marginal row. Dorsal bristles only 1.5–2.0 µm long, arranged in five kineties; kineties 2 and 5 distinctly shortened anteriorly (Fig. 141f, g). Caudal cirri present, however, difficult to recognise in life and protargol preparations because in elongation with left marginal row. Cell division: The frontal-ventral-transverse cirri are formed from the plesiomorphic number of six (I–VI) cirral anlagen (Gong et al. 2007). In addition,Gong aet al. (2007) found, like Hu & Song (2000), that per filial product only three new dorsal kineties are produced. Thus, one or two parental kineties must be retained because 4–5 kineties are present in interphasic specimens (Fig. 141f, g, Table 45). The macronuclear nodules fuse to a single mass. Molecular data: The length of the 18S rDNA sequence (including primers) is 1767 bp (GenBank accession numbers: EF194081– brackish water population; Fig. 141a; EF194082 and EF194083 – marine populations; Fig. 141b; Gong et al. 2007). The similarity between the two marine populations is 99.6%, whereas the similarities between brackish water population and marine populations is 98.4% and 98.5%. The partial sequence of 28S rDNA was 1284 bp long (EF631977; Gong et al. 2007). Occurrence and ecology: Gong et al. (2007) studied three Korean populations, namely two from a “coastal beach” of Jawol Island (37°15'N 126°32'E; salinity 30‰), Yellow Sea, Incheon and one brackish population from a tidal flat at Ganghwa (37°35'N 126°32'E; salinity 20–25‰). Samples were gathered from the top layer of the sediments.
1
Gong et al. (2007) wrote “... (AZM) is composed of 30–42 membranelles, which measure 11–16 µm long in vivo.” Thus, it is uncertain whether they allude to the length of the membranellar bases or the length of the cilia. 2 According to the description by Gong et al. (2007), the transverse cirri are positioned in the posterior 1/4 to 1/5 of the body. This is basically correct, but inexact because in the specimen shown in Fig. 141a the rearmost transverse cirrus is at 24% of body length (from behind), whereas at only 8% in the specimen shown in Fig. 141b. Perhaps, the specimen/population shown in Fig. 141b is a subspecies.
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Systematic Index The index contains all names of hypotrichous ciliates mentioned in the book, including vernacular names for example, amphisiellids; food organisms and other ciliates, except for those mentioned in the general section, are not included. The index is two-sided, that is, species appear both with the genus-group name first (for example, Amphisiella annulata) and with the species-group name first (annulata, Amphisiella). Valid (in my judgement) species and genera are in boldface italics print. Valid taxa not revised in the present book, invalid taxa, junior homonyms, synonyms, outdated combinations, incorrect spellings, nomina nuda are given in italics. The scientific name of a subgenus, when used with a binomen or trinomen, must be interpolated in parentheses between the genus-group name and the species-group name (ICZN 1999, Article 6.1). In the following index, these parentheses are omitted to simplify electronic sorting. Thus, the name Holosticha (Amphisiella) annulata is listed as Holosticha Amphisiella annulata. Note that this name is also listed under “Amphisiella annulata, Holosticha” and “annulata, Holosticha Amphisiella”. Suprageneric taxa are represented in normal type, valid ones revised in the present review in boldface. Boldface page number indicates the location of a valid taxon in the systematic section. “T” indicates the location of the table with the morphometric characterisation; “K” marks the page where a taxon is mentioned in a key (note that some taxa are mentioned two times in one key or in two different keys).
18-cirri hypotrich(s) xii, 1, 10, 12, 13, 15–17, 21, 25–27, 34, 45, 47, 71, 80, 104, 140, 164, 289, 352, 390, 410, 471, 473, 482, 507, 530, 533, 578, 582, 589, 593, 603, 614 18-cirri oxytrichid(s) 44, 104, 120, 454, 469, 675, 681 abdita, Afroamphisiella 4, 165, 371, 372K, 373, 377, 380T abdita, Lamtostyla 165, 372, 377 acrobates, Kahlia 468 acuta, Amphisiella 89, 351, 352 acuta, Paramphisiella 5, 72, 89, 255, 322, 352K, 352, 356T adami, Anteholosticha 39, 639K affine, Gonostomum 38, 53, 471, 476, 478 affine, Trachelostyla 476 affinis, Oxytricha 512 Afroamphisiella 78K, 166, 323, 353, 359, 371, 564 Afroamphisiella abdita 4, 165, 371, 372K, 373, 377, 380T Afroamphisiella multinucleata 4, 8, 74, 247, 372K, 372, 380T agiliformis, Urosomoida 579, 581, 582 agilis, Uroleptus 513, 530 agilis, Urosomoida 587 Allotricha 36, 468, 675, 682 Allotricha antarctica 615 Allotricha mollis 675 alpestris, Anteholosticha 636K Amphisellidae 79 Amphisia 85, 91, 94 Amphisia marioni 91
Amphisia Oxytricha capitata 91 Amphisiela 85 Amphisiela milnei 118 Amphisiella 5, 10, 17, 21, 25, 49–52, 66, 75K, 81, 84, 129, 130, 133, 148, 151, 163, 164, 169, 171, 183, 187, 190, 192, 201, 244, 245, 147, 249, 255, 262, 264, 267, 273, 280, 285, 320, 351, 371, 464, 476, 539, 560, 599, 652, 655, 683 Amphisiella acuta 89, 351, 352 Amphisiella annulata 1, 4, 5, 7, 8, 15–17, 47, 72, 86T, 90K, 94, 96, 99, 100, 119, 120, 123, 130, 348, 472, 683, 690, 693 Amphisiella annulata, Holosticha 101 Amphisiella antarctica 89, 129 Amphisiella anulata 103 Amphisiella arenicola 89, 127 Amphisiella australis 4, 89, 165, 167K, 169, 173, 179, 226T, 246K, 263 Amphisiella binucleata 261, 262, 264, 267 Amphisiella binucleata binucleata 262, 264 Amphisiella binucleata multicirrata 89, 262, 263, 267 Amphisiella capitata 4, 81, 90K, 91, 103, 108, 109, 116, 117, 119, 123, 128, 187, 190, 201, 683, 686T Amphisiella capitata, Holosticha 93 Amphisiella dorsicirrata 89 Amphisiella edaphoni 324 Amphisiella elegans 89, 164, 165, 192 Amphisiella faurei 89 Amphisiella gellerti 424, 426 Amphisiella Holosticha annulata 100 Amphisiella kihni 247
723
724
SYSTEMATIC INDEX
Amphisiella lithophora 89 Amphisiella longiseries 90, 245, 249 Amphisiella magnigranulosa 47, 90, 245, 273 Amphisiella marioni 82, 84, 91, 93–95, 97, 100, 108, 244, 247, 683, 685, 686T Amphisiella marioni, Holosticha 93 Amphisiella milenei 119 Amphisiella milnei 4, 5, 8, 90K, 94, 108, 109, 112, 118, 123 Amphisiella milnei, Holosticha 89, 119 Amphisiella multinucleata 90, 245, 255 Amphisiella namibiensis 90, 533, 534, 538 Amphisiella oblonga 463 Amphisiella oblonga, Holosticha 464 Amphisiella oscensis 4, 90, 129, 129K, 130, 133, 654K, 655, 657T Amphisiella ovalis 4, 89, 91K, 125 Amphisiella Oxytricha capitata 91 Amphisiella perisincirra 200 Amphisiella polycirrata 90, 245, 285 Amphisiella procera 90, 165, 183 Amphisiella quadrinucleata 90, 165, 190 Amphisiella raptans 90, 245, 252, 253 Amphisiella terricola 90, 245, 279 Amphisiella Tetrastyla oblonga 463 Amphisiella thiophaga, Holosticha 130 Amphisiella turanica 4, 89, 91K, 109, 117, 123 Amphisiella vitiphila 187 Amphisiella, Holosticha 48, 85, 90 amphisiellid(s) 1, 3, 5, 7, 10, 12, 13, 15–19, 21, 22, 28, 38, 41, 45, 47, 48, 51–53, 71, 72, 74, 512, 534, 538, 560, 561, 571, 598, 599, 636, 652, 653, 656 Amphisiellidae 1, 12, 18, 24, 46, 48, 49, 71, 79, 476, 534, 560, 561, 563, 651–653, 659 Amphisiellides xii, 49–51, 78K, 83, 88, 130, 133, 466, 561, 569, 651 Amphisiellides atypicus 4, 77, 130, 132, 133, 245, 246K, 561, 567T, 569, 652, 653, 654K, 654, 657T, 662 Amphisiellides illuvialis 561, 562, 569, 652–655 amphisiellids, core 82 Amphysiella 85 Amphysiella milnei 119 Ancystropodium 139 annulata, Amphisiella 1, 4, 5, 7, 8, 15–17, 47, 72, 86T, 90K, 94, 96, 99, 100, 119, 120, 123, 130, 348, 472, 683, 690, 693 annulata, Amphisiella Holosticha 100 annulata, Holosticha 89, 100 annulata, Holosticha Amphisiella 101 antarctica, Allotricha 615 antarctica, Amphisiella 89, 129
antarctica, Caudiamphisiella 4, 8, 72, 89, 129K, 129, 133, 135T, 151, 437, 655 antarctica, Urosomoida 578 antecirrata, Anteholosticha 638K Anteholosticha 77K, 389, 391, 392, 424, 636, 640, 643, 645, 647, 648 Anteholosticha adami 39, 639K Anteholosticha alpestris 636K Anteholosticha antecirrata 638K Anteholosticha arenicola 637K Anteholosticha australis 637K Anteholosticha azerbaijanica 638K, 639K Anteholosticha bergeri 638K, 647–649 Anteholosticha brachysticha 638K, 639K Anteholosticha brevis 636K Anteholosticha camerounensis 637K Anteholosticha distyla 638K Anteholosticha estuarii 638K Anteholosticha extensa 637K Anteholosticha fasciola 639K Anteholosticha gracilis 638K, 639K Anteholosticha grisea 639K Anteholosticha heterocirrata 4, 77, 339K, 640K, 640 Anteholosticha intermedia 639K Anteholosticha longissima 639K Anteholosticha macrostoma 637K Anteholosticha manca 639K Anteholosticha mancoidea 638K Anteholosticha monilata 637K Anteholosticha multistilata 638K Anteholosticha muscicola 637K Anteholosticha oculata 119, 122, 637K Anteholosticha plurinucleata 639K Anteholosticha pulchra 28, 638K Anteholosticha randani 638K, 639K Anteholosticha scutellum 639K Anteholosticha sigmoidea 637K Anteholosticha sphagni 638K Anteholosticha thononensis 639K Anteholosticha verrucosa 4, 393K, 396T, 402, 424, 426, 427, 636, 640K, 647 Anteholosticha violacea 637K, 639K Anteholosticha vuxgracilis 637K Anteholosticha warreni 639K Anteholosticha xanthichroma 637K, 638K anulata, Amphisiella 103 Apoamphisiella 46, 151, 539 Apokeronopsis 47 Apourosomoida 5, 48, 52, 71, 75K, 78K, 514, 582, 589, 615 Apourosomoida halophila 4, 10, 18, 21, 75, 515, 516K, 516, 527T, 530, 582
SYSTEMATIC INDEX Apourosomoida natronophila 4, 52, 515, 516K, 530 Architricha 36, 675 Architricha indica 675 arenicola, Amphisiella 89, 127 arenicola, Anteholosticha 637K arenicola, Circinella 391, 392, 467 arenicola, Erimophrya 4, 578, 579K, 579, 583T, 586, 590, 593 arenicola, Vermioxytricha 5, 9, 77, 391, 428, 596, 597T, 599, 600, 607 Aspidisca 25, 28, 30, 31, 33, 43 Aspidiscidae 24 aspidiscids 475 atypica, Uroleptoides 466, 653–655 atypicus, Amphisiellides 4, 77, 130, 132, 133, 245, 246K, 561, 567T, 569, 652, 653, 654K, 654, 657T, 662 atypicus, Uroleptoides 245, 651, 652, 654, 655 auricularis, Epiclintes 493 auricularis, Oxytricha 467 australis, Amphisiella 4, 89, 165, 167K, 169, 173, 179, 226T, 246K, 263 australis, Anteholosticha 637K australis, Lamtostyla 3, 4, 20, 89, 164, 167K, 167, 169, 179, 180, 182, 183, 187, 201, 212, 226T, 329, 331, 341 Australocirrus 46 azerbaijanica, Anteholosticha 638K, 639K Balladyna 49, 466 Balladyna fusiformis 466 Balladyna parvula 466 Balladynella 49, 466 Banylsellidae 467 Banyulsella 49, 466 Banyulsella viridis 466 Banyulsellidae 467 bavariensis, Gastrostyla 138 bavariensis, Gastrostyla Kleinstyla 140 bavariensis, Kleinstyla 140 bergeri, Anteholosticha 638K, 647–649 bicirratum, Tachysoma 200 Biholosticha discocephalus 472 binucleata binucleata, Amphisiella 262, 264 binucleata multicirrata, Amphisiella 89, 262, 263, 267 binucleata, Amphisiella 261, 262, 264, 267 binucleata, Amphisiella binucleata 262, 264 binucleata, Uroleptoides 261, 262, 264 binucleatus binucleatus, Uroleptoides 4, 9, 226T, 263K, 264, 267 binucleatus multicirratus, Uroleptoides 4, 9, 89, 226T, 251, 264K, 264, 267
725
binucleatus, Uroleptoides 72, 180, 183, 246K, 249, 253, 261, 273 binucleatus, Uroleptoides binucleatus 4, 9, 226T, 263K, 264, 267 Bistichella 13, 77K, 289, 514, 532, 660 Bistichella buitkampi 4, 75, 193, 533, 534K, 535, 548, 550T, 554, 661 Bistichella humicola 5, 533, 534K, 548, 550T, 556, 661 Bistichella namibiensis 4, 90, 285, 533, 534K, 538, 552T, 552, 544 Bistichella procera 4, 533, 534K, 535, 547, 550T, 554, 555, 661 Bistichella terrestris 4, 533, 534K, 548, 550T, 552, 554, 557, 661 Bistichella terrestris-group 548 borrori, Prodiscocephalus 44 brachysticha, Anteholosticha 638K, 639K brachytona, Eschaneustyla 467 brandti, Cyttarocylis 41 brevis, Anteholosticha 636K breviseries, Orthoamphisiella 39, 373, 469, 690 bryonicolum, Trachelostyla 475 bryophila, Trachelochaeta 470 buitkampi, Bistichella 4, 75, 193, 533, 534K, 535, 548, 550T, 554, 661 buitkampi, Hemisincirra 5, 74, 388, 390, 393K, 393, 396T, 407, 408, 428, 437, 599, 640 buitkampi, Paraurostyla 533–535, 661 buitkampi, Perisincirra 393, 394 buitkampi, Pseudouroleptus 535 camerounensis, Anteholosticha 637K canadiensis, Trachelostyla 478 capitata, Amphisia Oxytricha 91 capitata, Amphisiella 4, 81, 90K, 91, 103, 108, 109, 116, 117, 119, 123, 128, 187, 190, 201, 683, 686T capitata, Amphisiella Oxytricha 91 capitata, Holosticha Amphisiella 93 capitata, Oxytricha 89, 91, 93–95 caudata, Paramphisiella 5, 18, 245, 351, 352K, 353, 356T, 358 caudata, Trachelostyla 5, 383, 475, 476, 478, 478K, 494, 501, 502, 511, 689 caudata, Uroleptoides 245, 352, 358, 359 caudata, Urosoma 631 caudatus caudatus, Pseudouroleptus 662, 662K, 663, 667T, 669, 671 caudatus namibiensis, Pseudouroleptus 312, 662, 663K, 667T, 668 caudatus, Pseudouroleptus 4, 77, 289, 293, 308, 469, 533, 554, 653, 658, 660, 661
726
SYSTEMATIC INDEX
caudatus, Pseudouroleptus caudatus 662, 662K, 663, 667T, 669, 671 caudatus, Uroleptoides 359 Caudiamphisiella 75K, 84, 90, 129, 437, 655, 657 Caudiamphisiella antarctica 4, 8, 72, 89, 129K, 129, 133, 135T, 151, 437, 655 Caudiholosticha 389, 392, 636 Caudiholosticha gracilis 392 Caudiholosticha navicularum 636K Caudiholosticha setifera 103, 104 Caudiholosticha sylvatica 120 Certesia 25, 28, 31 Chlamydodontidae 3 Choreotrichia 24, 38 Choreotrichida 24 cienkowskii, Urosoma 513, 631 ciliophorum, Trachelostyla 475 Circinella 50, 82, 370, 389, 391, 392, 432, 467, 599 Circinella arenicola 391, 392, 467 Circinella filiformis 428, 432 Circinella vettersi 428 citrina, Uroleptopsis 118, 225 Cladotricha 49, 467, 517 Cladotricha koltzowii 467 Cladotricha variabilis 516, 517, 529 Cladotrichidae 163, 244 Codonella 25 colpodid(s) 30, 39, 40 Coniculostomum 36, 46, 49, 467, 673, 675 core amphisiellids 82 corlissi, Meseres 28 cornuta, Stichochaeta 477 coronata, Gastrostyla Holosticha 137 coronata, Holosticha 137, 138, 141, 148 coronata, Holosticha Keronopsis 138 coronata, Keronopsis Holosticha 137 corsica, Stichochaeta 479, 482, 483, 485, 492 Cossothigma 72K, 382, 472, 478, 481, 510 Cossothigma dubium 4, 74, 153, 380K, 383, 386, 387, 506 crassa, Oxytricha 47 crassum, Strongylidium 297 cristata, Pseudourostyla 39 Cyrtohymena 44, 46, 206, 259, 261, 536 Cyrtohymena tetracirrata 464 Cyrtostrombidium 38 Cytharoides 28, 31, 43 Cyttarocylis brandti 41 decorata, Lamtostyla 4, 8, 164, 165, 166K, 205, 206, 212, 218, 226T deserticola, Urosomoida 578 diademata, Holosticha 90, 130
Diophrys 22, 25, 26, 28–31, 33–44 Diophrys hystrix 41 Diophrys scutum 33, 35, 40 discocephalid(s) 13, 22, 25, 26, 28–31, 33–44 Discocephalidae 25 Discocephalus 25, 31 discocephalus, Biholosticha 472 distichum, Metastrongylidium 148–151, 153, 160T, 161 distyla, Anteholosticha 638K dorsicirrata, Amphisiella 89 dorsicirrata, Gastrostyla 140 dorsicirrata, Kleinstyla 140 Dorsomarginalia xi, xii, 18, 21, 26, 28, 37, 39–42, 44–47, 80, 83, 163, 206, 321, 390, 472, 515, 534, 560, 561, 563, 566, 577, 578, 596, 598, 599, 614, 615, 617, 618, 651, 653, 658, 689 Dorsomarginalia, non-oxytrichid 46, 77, 472, 513, 560, 578 dubia, Gastrostyla 383 dubia, Trachelostyla 382, 383, 478 dubium, Cossothigma 4, 74, 153, 380K, 383, 386, 387, 506 dubium, Holosticha Parurosoma 469 Dysteriidae 3 edaphoni, Amphisiella 324 edaphoni, Lamtostyla 163, 165, 201, 323–325 edaphoni, Lamtostylides 3, 4, 72, 163, 165, 201, 226T, 324K, 324, 334, 336 elegans, Amphisiella 89, 164, 165, 192 elegans, Lamtostyla 4, 89, 164, 166K, 192, 226T elongata, Oxytricha 477 elongata, Oxytricha Opisthotricha 477 Engelmanniella 618 Engelmanniella mobilis 42, 607, 618 enigmatica, Hemigastrostyla 47, 140, 472, 473, 690 enigmatica, Mixotricha 673 enigmatica, Paraurostyla 672, 673 enigmatica, Ponturostyla 4, 77, 651, 673, 681T Entodinium 33, 34 Epiclintes 49, 467 Epiclintes auricularis 493 Erimophrya 71, 78K, 515, 560, 577, 599, 615, 618, 629 Erimophrya arenicola 4, 578, 579K, 579, 583T, 586, 590, 593 Erimophrya glatzeli 4, 77, 577, 578, 579K, 579, 583T, 589 Erimophrya quadrinucleata 4, 516K, 517, 577, 578, 579K, 583T, 592, 593, 595, 614 Erimophrya sylvatica 4, 578, 579K, 583T, 590, 595 Eschaneustyla 40, 49
SYSTEMATIC INDEX Eschaneustyla brachytona 467 espeletiae, Fragmocirrus 193 estuarii, Anteholosticha 638K Euhypotrichina 79 Euplota 24, 25 Euplotes 25, 28, 30, 31, 33, 34, 36–40, 43 Euplotia 24 euplotid(s) 17, 20, 22–26, 28–31, 33–35, 37, 39–44, 66, 88, 117, 118, 468, 475 Euplotida 24 Euplotidae 24 Euplotidium 31 extensa, Anteholosticha 637K fasciola, Anteholosticha 639K faurei, Amphisiella 89 faurei, Marginotricha 30, 33, 34, 89 faurei, Psammocephalus 33, 89 filiformis, Circinella 428, 432 filiformis, Perisincirra 392 flexilis, Onychodromopsis 468, 615 fossicola, Paraurostyla 539 Fragmocirrus espeletiae 193 furcata, Tachysoma 200 fusiformis, Balladyna 466 Gastrocirrhidae 24 Gastrocirrhus 31, 138 Gastrostyla 45, 46, 50, 82, 84, 136–141, 151, 383, 384, 467, 476, 665, 690 Gastrostyla bavariensis 138 Gastrostyla dorsicirrata 140 Gastrostyla dubia 383 Gastrostyla Gastrostyla 138, 468 Gastrostyla Gastrostyla pulchra 138 Gastrostyla Holosticha coronata 137 Gastrostyla Kleinstyla 138 Gastrostyla Kleinstyla bavariensis 140 Gastrostyla minima 140 Gastrostyla muscorum 139 Gastrostyla mystacea 138 Gastrostyla opisthoclada 140 Gastrostyla pulchra 84, 137, 140, 693 Gastrostyla pulchra, Gastrostyla 138 Gastrostyla setifera 139 Gastrostyla Spetastyla 138, 468 Gastrostyla Spetastyla mystacea 468 Gastrostyla steinii 82, 84, 138–140, 151, 390 Gastrostyla stenocephala 151, 383 Gastrostyla Stylonychia pulchra 137 Gastrostyla, Gastrostyla 138, 468 gastrostylid(s) 139, 390 Gastrostylidae 139, 140, 390 geleii, Trachelostyla 475
727
gellerti gellerti, Hemisincirra 424, 647 gellerti verrucosa, Hemisincirra 391, 424, 647, 648 gellerti, Amphisiella 424, 426 gellerti, Hemisincira 388 gellerti, Hemisincirra 4, 5, 8, 393K, 396T, 418, 424, 437, 647, 648 gellerti, Hemisincirra gellerti 424, 647 gellerti, Perisincira 424 gellerti, Perisincirra 388, 391, 424, 647 gibba, Holosticha 42 glatzeli, Erimophrya 4, 77, 577, 578, 579K, 579, 583T, 589 Glaucoma 38 Glaucoma scintillans 53 goertzi, Hemiurosoma 4, 614, 616T, 618, 619K, 628 Gonostomatidae 471, 472 Gonostomum xii, 16, 40, 45–47, 51, 80, 81, 222, 307, 390, 407, 415, 448, 469, 471–473, 475–477, 479, 481, 482, 486, 503, 505, 512, 515, 533, 614, 615, 617, 618, 623, 625, 631, 691 Gonostomum affine 38, 53, 471, 476, 478 Gonostomum namibiense 690 Gonostomum pediculiforme 479 Gonostomum pediculiformis 479 Gonostomum strenuum 690 gracilis, Anteholosticha 638K, 639K gracilis, Caudiholosticha 392 gracilis, Perisincirra 388, 392 grandis, Onychodromus 468 grandis, Pleurotricha 675 grandis, Urostyla 41, 42 granulifera, Hemiamphisiella 4, 8, 289, 290T, 294K, 311, 315, 662 granulifera, Lamtostyla 4, 8, 10, 16, 48, 72, 164, 165, 166K, 205, 206, 212, 218, 220, 222, 223, 226T, 336, 473, 482, 486 granulifera, Oxytricha 482, 615, 689 granulifera-group, Lamtostyla 71, 78K, 164, 165, 205 granuliferum, Strongylidium 294, 315 grelli, Orthoamphisiella 373 grisea, Anteholosticha 639K group, Bistichella terrestris- 548 group, Lamtostyla granulifera- 71, 78K, 164, 165, 205 group, Lamtostyla lamottei- 164, 165, 167 group, Lamtostyla longa- 71, 78K, 164, 165, 218 group, Spirotrachelostyla simplex- 506 halophila, Apourosomoida 4, 10, 18, 21, 75, 515, 516K, 516, 527T, 530, 582 halophila, Lamtostyla 165, 323, 336 halophila, Parakahliella 193
728
SYSTEMATIC INDEX
halophilus, Lamtostylides 4, 8, 165, 226T, 323, 324K, 336, 344, 345, 347, 449, 449K Halteria 24, 33, 37 halterid(s) 28, 33, 38 Halteriida 38 haptorid(s) 30, 39 hembergeri, Spiroamphisiella 4, 17, 72, 140, 148, 149T, 150, 160T, 384, 506 Hemiamphisiella 21, 49–51, 77K, 80, 83, 88, 161, 244, 288, 352, 442, 443, 446, 650, 662, 665, 671 Hemiamphisiella granulifera 4, 8, 289, 290T, 294K, 311, 315, 662 Hemiamphisiella qingdaoensis 310 Hemiamphisiella quadrinucleata 4, 8, 10, 187, 190, 245, 289, 290T, 294K, 318, 562, 562K, 563, 564, 567T Hemiamphisiella terricola 72, 288, 289, 294K, 294, 311, 312, 320–322, 352, 442, 562, 660, 690 Hemiamphisiella terricola qingdaoensis 5, 245, 290T, 295, 296K, 310 Hemiamphisiella terricola terricola 5, 8, 289, 290T, 293–295, 296K, 296, 311 Hemiamphisiella wilberti 4, 289, 290T, 294K, 311, 315, 660, 662, 671 Hemicinsirra sp. 388 Hemigastrostyla 141, 151, 384, 472, 473, 691 Hemigastrostyla enigmatica 47, 140, 472, 473, 690 Hemigastrostyla stenocephala 140, 383, 472 Hemigastrostyla szaboi 472 Hemisincira gellerti 388 Hemisincirra 51, 71, 78K, 83, 139, 200, 201, 323, 387, 448–450, 458, 512, 578, 579, 598, 599, 607, 614, 618, 633, 634, 636, 643, 645 Hemisincirra buitkampi 5, 74, 388, 390, 393K, 393, 396T, 407, 408, 428, 437, 599, 643 Hemisincirra gellerti 4, 5, 8, 393K, 396T, 418, 424, 437, 647, 648 Hemisincirra gellerti gellerti 424, 647 Hemisincirra gellerti verrucosa 391, 424, 647, 648 Hemisincirra heterocirrata 391, 640, 643 Hemisincirra inquieta 5, 8, 390, 391, 393K, 394, 396T, 403, 437, 473 Hemisincirra interrupta 5, 389, 393K, 396T, 428, 432, 600, 607 Hemisincirra livida 391, 449, 458 Hemisincirra muelleri 391, 599, 607 Hemisincirra namibiensis 4, 193, 391, 392K, 396T, 418 Hemisincirra octonucleata 4, 391, 392K, 396T, 421, 450 Hemisincirra polynucleata 391, 618, 634 Hemisincirra pori 323, 344
Hemisincirra quadrinucleata 4, 193, 245, 391, 392K, 396T, 418, 419, 449K, 454 Hemisincirra rariseta 5, 389, 391, 393K, 396T, 428, 599, 600, 607 Hemisincirra similis 633 Hemisincirra vermiculare 435 Hemisincirra vermicularis 5, 389, 391, 392K, 396T, 428, 435, 599 Hemisincirra verrucosa 647 Hemisincirra vettersi 391 Hemisincirra viridis 450 Hemisincirra wenzeli 5, 8, 391, 393K, 394, 396T, 408, 424, 437 Hemiurosoma 13, 46, 71, 78K, 389, 391, 392, 437, 515, 560, 578, 579, 599, 614 Hemiurosoma goertzi 4, 614, 616T, 618, 619K, 628 Hemiurosoma polynucleatum 4, 619K, 628, 634 Hemiurosoma similis 4, 587, 614, 619K, 622, 631, 633 Hemiurosoma terricola 4, 77, 614, 616T, 618, 619K, 619 heterocirrata, Anteholosticha 4, 77, 339K, 640K, 640 heterocirrata, Hemisincirra 391, 640, 643 heterocirrata, Perisincirra 640 heterofoissneri, Holosticha 481 heterotrichid(s) 30 Histriculus 46 holomilnei, Holosticha 119 Holosticha 25, 40, 42, 66, 81, 85, 91, 93, 94, 101, 119, 128, 138, 141, 464, 691 Holosticha Amphisiella 48, 85, 90 Holosticha Amphisiella annulata 101 Holosticha Amphisiella capitata 93 Holosticha Amphisiella marioni 93 Holosticha Amphisiella milnei 89, 119 Holosticha Amphisiella oblonga 464 Holosticha Amphisiella thiophaga 130 Holosticha annulata 89, 100 Holosticha annulata, Amphisiella 100 Holosticha coronata 137, 138, 141, 148 Holosticha coronata, Gastrostyla 137 Holosticha coronata, Keronopsis 137 Holosticha diademata 90, 130 Holosticha gibba 42 Holosticha heterofoissneri 481 Holosticha holomilnei 119 Holosticha Holosticha milnei 119, 122 Holosticha Keronopsis coronata 138 Holosticha kessleri 42 Holosticha milnei 122 Holosticha milnei, Holosticha 119, 122 Holosticha multistilata 279
SYSTEMATIC INDEX Holosticha obliqua 103, 104 Holosticha Parurosoma dubium 469 Holosticha pullaster 28 Holosticha setifera 103 Holosticha thiophaga 90 holostichid(s) 81, 163, 225 Holostichidae 24, 88, 163, 476 Holostichides 469 hospes, Mucotrichdium 4, 74, 75, 153, 441 hospes, Uroleptus 440, 441 humicola, Bistichella 5, 533, 534K, 548, 550T, 556, 661 humicola, Pseudouroleptus 557 humicola, Uroleptus 534, 556, 557, 661 hyalina, Lamtostyla 165, 323, 347 hyalina, Tachysoma 324, 347 hyalinum, Tachysoma 348 hyalinus, Lamtostylides 4, 164, 165, 226T, 324K, 347 hypotrich(s) 1, 3, 5, 7, 10, 12, 16–23, 25, 26, 28, 30, 33–36, 38, 41–44, 51, 52, 65, 66, 117, 294, 390, 472, 515, 517, 560, 562, 563, 688 hypotrich(s), 18-cirri xii, 1, 10, 12, 13, 15–17, 21, 25–27, 34, 45, 47, 71, 80, 104, 140, 164, 289, 352, 390, 410, 471, 473, 482, 507, 530, 533, 578, 582, 589, 593, 603, 614 Hypotricha 1, 12, 17, 21–26, 28–31, 34–38, 40, 41, 43–48, 66, 75, 79, 80, 81, 220, 294, 472, 477, 503, 514, 533, 534, 560, 596, 615, 617, 660, 689, 691 Hypotricha, non-dorsomarginalian 46 Hypotrichia 24, 28, 44 Hypotrichida 25, 79, 384 Hypotrichidium 38 hystrix, Diophrys 41 illuvialis, Amphisiellides 561, 562, 569, 652–655 illuvialis, Nudiamphisiella 4, 562K, 563, 565, 566, 567T, 569, 576T, 654K, 654–656 indica, Architricha 675 inquieta, Hemisincirra 5, 8, 390, 391, 393K, 394, 396T, 403, 437, 473 inquieta, Perisincirra 403 intermedia, Anteholosticha 639K interrupta, Hemisincirra 5, 389, 393K, 396T, 428, 432, 600, 607 interrupta, Nudiamphisiella 4, 9, 12, 18, 77, 289, 560, 561, 562K, 562, 567T, 571, 654K interrupta, Paruroleptus (?) 432 interrupta, Perisincira 432 interrupta, Perisincirra 388, 391, 432 islandica, Lamtostyla 4, 164, 165, 166K, 196, 201, 220, 226T, 325, 334,
729
kahli, Perisincirra 393 kahli, Uroleptus 389, 391, 393, 394, 450, 458, 512, 530 Kahlia acrobates 468 Kahliella 49, 468, 472 Kahliellidae xii, 88, 244, 660 Keronella 38 Keronopsidae 660 Keronopsis 53, 138 Keronopsis coronata, Holosticha 138 Keronopsis Holosticha coronata 137 kessleri, Holosticha 42 kihni, Amphisiella 247 kihni, Uroleptoides 5, 88, 224, 244, 245, 246K, 246, 294, 352, 372–374 Kiitricha 31 kirkeniensis, Lamtostyla 165, 323, 333 kirkeniensis, Lamtostylides 4, 165, 226T, 324K, 325, 333, 336 kirkensis, Lamtostyla 334 Kleinstyla 140 Kleinstyla bavariensis 140 Kleinstyla bavariensis, Gastrostyla 140 Kleinstyla dorsicirrata 140 Kleinstyla, Gastrostyla 138 koltzowii, Cladotricha 467 Laboea 38 Lacazea 49, 468 Lacazea ovalis 468 lacazei, Pseudoamphisiella 117 lamottei, Lamtostyla 4, 72, 88, 161, 163–165, 166K, 167, 171, 187, 190, 201, 220, 245, 512 lamottei-group, Lamtostyla 164, 165, 167 Lamtostyla 10, 16, 19, 38, 50, 51, 78K, 80, 81, 88, 161, 245, 246, 273, 322–324, 334, 336, 348, 353, 377, 390, 422, 448, 449, 512 Lamtostyla abdita 165, 372, 377 Lamtostyla australis 3, 4, 20, 89, 164, 167K, 167, 169, 179, 180, 182, 183, 187, 201, 212, 226T, 329, 331, 341 Lamtostyla decorata 4, 8, 164, 165, 166K, 205, 206, 212, 218, 226T Lamtostyla edaphoni 163, 165, 201, 323–325 Lamtostyla elegans 4, 89, 164, 166K, 192, 226T Lamtostyla granulifera 4, 8, 10, 16, 48, 72, 164, 165, 166K, 205, 206, 212, 218, 220, 222, 223, 226T, 336, 473, 482, 486 Lamtostyla granulifera-group 71, 78K, 164, 165, 205 Lamtostyla halophila 165, 323, 336 Lamtostyla hyalina 165, 323, 347
730
SYSTEMATIC INDEX
Lamtostyla islandica 4, 164, 165, 166K, 196, 201, 220, 226T, 325, 334 Lamtostyla kirkeniensis 165, 323, 333 Lamtostyla kirkensis 334 Lamtostyla lamottei 4, 72, 88, 161, 163–165, 166K, 167, 171, 187, 190, 201, 220, 245, 512 Lamtostyla lamottei-group 164, 165, 167 Lamtostyla longa 4, 72, 164, 165, 166K, 206, 212, 218, 222, 223, 226T Lamtostyla longa-group 71, 78K, 164, 165, 218 Lamtostyla perisincirra 3, 4, 164, 166K, 197, 199, 200, 220, 226T, 329, 341 Lamtostyla procera 1, 4, 90, 164, 167K, 183, 226T, 273 Lamtostyla quadrinucleata 4, 90, 164, 166K, 187, 190, 193, 226T Lamtostyla raptans 4, 164, 166K, 206, 212, 218, 220, 221, 226T Lamtostyla vitiphila 4, 164, 166K, 187, 190, 193, 226T, 245, 320 Lamtostylides 78K, 164, 166, 246, 322, 391, 392, 448, 449, 454 Lamtostylides edaphoni 3, 4, 72, 163, 165, 201, 226T, 324K, 324, 334, 336 Lamtostylides halophilus 4, 8, 165, 226T, 323, 324K, 336, 344, 345, 347, 449, 449K Lamtostylides hyalinus 4, 164, 165, 226T, 324K, 347 Lamtostylides kirkeniensis 4, 165, 226T, 324K, 325, 333, 336 Lamtostylides pori 4, 226T, 324K, 344 lanceolata, Oxytricha 617 lanceolata, Paragastrostyla 469, 564 lanceolata, Periholosticha 469 Laurentia monilata 467 Laurentiella 46, 653 lemnae, Stylonychia 42 levis, Pseudorurostyla 39 Limnostrombidium 25 lithophora, Amphisiella 89 lithophora, Marginotricha 89 lithophora, Psammocephalus 89 livida, Hemisincirra 391, 449, 458 livida, Terricirra 5, 9, 448, 449K, 458, 460T Lohmanniella 25 longa, Lamtostyla 4, 72, 164, 165, 166K, 206, 212, 218, 222, 223, 226T longa, Tachysoma 165, 218 longa-group, Lamtostyla 71, 78K, 164, 165, 218 longiseries, Amphisiella 90, 245, 249 longiseries, Uroleptoides 4, 9, 90, 193, 226T, 246K, 249
longissima, Anteholosticha 639K macrostoma, Anteholosticha 637K macrostoma, Trachelostyla 475 magnigranulosa, Amphisiella 47, 90, 245, 273 magnigranulosus, Uroleptoides 4, 9, 47, 90, 182, 183, 226T, 246K, 249, 255, 263, 267, 272, 273, 390 manca, Anteholosticha 639K mancoidea, Anteholosticha 638K Maregastrostyla 21, 36, 40, 72K, 84, 90, 130, 136, 151, 468, 690, 693 Maregastrostyla pulchra 4, 9, 36, 72, 137, 147T, 153, 693 Marginotricha 25 Marginotricha faurei 30, 33, 34, 89 Marginotricha lithophora 89 marioni, Amphisia 91 marioni, Amphisiella 82, 84, 91, 93–95, 97, 100, 108, 244, 247, 683, 685, 686T marioni, Holosticha Amphisiella 93 matsusakai, Terricirra 4, 9, 193, 420, 448, 449, 449K, 450, 454, 458, 460T Meseres 24, 33, 37, 39, 40 Meseres corlissi 28 Metastrongylidium 148–151 Metastrongylidium distichum 148–151, 153, 160T, 161 milenei, Amphisiella 119 milnei, Amphisiela 118 milnei, Amphisiella 4, 5, 8, 90K, 94, 108, 109, 112, 118, 123 milnei, Amphysiella 119 milnei, Holosticha 122 milnei, Holosticha Amphisiella 89, 119 milnei, Holosticha Holosticha 119, 122 minima, Gastrostyla 140 minima, Spetastyla mystacea 140 minima, Urosomoida 515, 516K, 517 Mitra radiosa 469, 512 Mixotricha 672, 673, 675 Mixotricha enigmatica 673 mobilis, Engelmanniella 42, 607, 618 mollis, Allotricha 675 monilata, Anteholosticha 637K monilata, Laurentia 467 monostyla, Urosomoida 578 mucicola, Strongylidium 445 Mucotrichdium hospes 4, 74, 75, 153, 441 Mucotrichidium 50, 77K, 440 muelleri, Hemisincirra 391, 599, 607 muelleri, Vermioxytricha 5, 428, 597T, 598–600, 600K, 605, 607
SYSTEMATIC INDEX multicirrata, Amphisiella binucleata 89, 262, 263, 267 multicirratus, Uroleptoides binucleatus 4, 9, 89, 226T, 251, 264K, 264, 267 multinucleata, Afroamphisiella 4, 8, 74, 247, 372K, 372, 380T multinucleata, Amphisiella 90, 245, 255 multinucleatus, Uroleptoides 5, 9, 77, 90, 226T, 246K, 251, 253, 255, 272 multistilata, Anteholosticha 638K multistilata, Holosticha 279 muscicola, Anteholosticha 637K muscorum, Gastrostyla 139 muscorum, Strongylidium 289, 294, 296, 297, 311 musculus, Trichoda 443 mutsusakai, Tericirra 448 mystacea minima, Spetastyla 140 mystacea mystacea, Spetastyla 140 mystacea, Gastrostyla 138 mystacea, Gastrostyla Spetastyla 468 mystacea, Oxytricha 140 mystacea, Spetastyla 140 mystacea, Spetastyla mystacea 140 mytilus, Stylonychia 482 namibiense, Gonostomum 690 namibiensis, Amphisiella 90, 533, 534, 538 namibiensis, Bistichella 4, 90, 285, 533, 534K, 538, 552T, 552, 554 namibiensis, Hemisincirra 4, 193, 391, 392K, 396T, 418 namibiensis, Pseudouroleptus caudatus 312, 662, 663K, 667T, 668 natronophila, Apourosomoida 4, 52, 515, 516K, 530 natronophilus, Uroleptus 515, 530, 531 navicularum, Caudiholosticha 636K Neokeronopsis xi, 25, 44, 46, 47, 65, 82, 359, 560, 661 Neokeronopsis spectabilis 18 non-dorsomarginalian Hypotricha 46 non-oxytrichid Dorsomarginalia 46, 77, 472, 513, 560, 578 non-stylonychine Oxytrichidae 46 Notohymena 46 nova, Pseudourostyla 636 Nudiamphisiella 21, 46, 78K, 83, 289, 320, 321, 560, 561, 653, 655 Nudiamphisiella illuvialis 4, 562K, 563, 565, 566, 567T, 569, 576T, 654K, 654–656 Nudiamphisiella interrupta 4, 9, 12, 18, 77, 289, 560, 561, 562K, 562, 567T, 571, 654K obliqua, Holosticha 103, 104
731
oblonga, Amphisiella 463 oblonga, Amphisiella Tetrastyla 463 oblonga, Holosticha Amphisiella 464 oblonga, Tetrastyla 4, 74, 75, 463 octonucleata, Hemisincirra 4, 391, 392K, 396T, 421, 450 octonucleata, Perisincirra 421 octonucleata, Urosoma 614 oculata, Anteholosticha 119, 122, 637K oligotrich(s) 23–26, 28–31, 33, 34, 36–44, 117 Oligotricha 24, 25, 37 Oligotrichea 24, 38, 41 Oligotrichia 24, 37, 38 Oligotrichida 24, 41 Onychodromopsis 49, 468 Onychodromopsis flexilis 468, 615 Onychodromus 23, 46, 49, 468, 653 Onychodromus grandis 468 Onychodromus quadricornutus 468 Ophrydium 447 opisthoclada, Gastrostyla 140 Opisthotricha elongata, Oxytricha 477 Orthoamphisiella 50, 51, 82, 190, 244, 371, 377, 468 Orthoamphisiella breviseries 39, 373, 469, 690 Orthoamphisiella grelli 373 Orthoamphisiella stramenticola 373, 468 Orthoamphisiellidae 467, 469 oscensis, Amphisiella 4, 90, 129, 129K, 130, 133, 654K, 655, 657T ovalis, Amphisiella 4, 89, 91K, 125 ovalis, Lacazea 468 Oxytricha 23, 25, 33, 34, 38, 42, 43, 46, 81, 104, 112, 220, 473, 577, 596, 615, 659, 689–691 Oxytricha affinis 512 Oxytricha auricularis 467 Oxytricha capitata 89, 91, 93–95 Oxytricha capitata, Amphisia 91 Oxytricha capitata, Amphisiella 91 Oxytricha crassa 47 Oxytricha elongata 477 Oxytricha granulifera 482, 615, 689 Oxytricha lanceolata 617 Oxytricha mystacea 140 Oxytricha Opisthotricha elongata 477 Oxytricha setigera 466 Oxytricha stenocephala 141 Oxytrichia 24 oxytrichid(s) xii, 12, 18–21, 24, 26, 28, 34, 37, 39, 40, 42, 45, 51, 65, 71, 81–83, 88, 138, 140, 141, 151, 163, 206, 289, 293, 390, 466, 468–470, 472, 473, 475, 477, 513, 515, 533, 539, 560, 561, 578,
732
SYSTEMATIC INDEX
582, 596, 615, 651–653, 659–661, 665, 673, 675, 682, 688, 689 oxytrichid(s), 18-cirri 44, 104, 120, 454, 469, 675, 681 Oxytrichidae xi, 12, 18, 24–26, 29, 42, 44–47, 77, 81, 82, 88, 140, 162, 163, 293, 384, 447, 448, 467, 470, 473, 475, 514, 515, 533, 560, 577, 578, 596, 598, 615, 651, 653, 660, 672 Oxytrichidae, non-stylonychine 46 Oxytrichidea 24 Oxytrichinae 44, 47 oxytrichines 45 Parabirojimia similis 690 Paragastrostyla 50, 51, 392, 469, 564 Paragastrostyla lanceolata 469, 564 Paragonostomum 473 Paraholosticha 53 Parakahliella 36, 46 Parakahliella halophila 193 Parakahliellidae 653 Paramecium 33, 34, 38 Paramphisiella 49–51, 78K, 88, 244, 294, 322, 351, 352 Paramphisiella acuta 5, 72, 89, 255, 322, 352K, 352, 356T Paramphisiella caudata 5, 18, 245, 351, 352K, 353, 356T, 358 Parastylonychia 139 Paraurostyla 33, 34, 43, 46, 49, 469, 535, 560, 673 Paraurostyla buitkampi 533–535, 661 Paraurostyla enigmatica 672, 673 Paraurostyla fossicola 539 Paraurostyla viridis 618 Paraurostyla weissei 42, 535, 673 Parentocirrus 539 Paruroleptus (?) interrupta 432 Paruroleptus 225 Parurosoma 49, 469 Parurosoma dubium, Holosticha 469 parvula, Balladyna 466 Pattersoniella xi, 45–47, 653, 691 paucicirrata, Periholosticha 407 pediculariformis, Trachelostyla 480 pediculiforme, Gonostomum 479 pediculiforme, Stichochaeta 493 pediculiforme, Trachelostyla 480 pediculiformis, Gonostomum 479 pediculiformis, Stichochaeta 4, 470, 474, 475, 477, 478, 500 pediculiformis, Trachelostyla 5, 12, 18, 26, 75, 383, 384, 386, 387, 390, 476, 478K, 478, 494–498, 500, 501T, 502, 505, 683, 687
pediculiformis, Trachelostyla Stichochaeta 479, 480 pediculiformis, Traxhelostyla 475 Pelagohalteria 24 Pelagotrichidium 38 pellionellum, Tachysoma 5, 120 pendiculiformis, Trachelostyla 480 Periholosticha 51, 391, 392, 407, 469, 643 Periholosticha lanceolata 469 Periholosticha paucicirrata 407 Perilemmaphora 25, 26, 37, 38, 40, 42 Perisincira gellerti 424 Perisincira interrupta 432 Perisincira viridis 450 Perisincirra 200, 387–391, 394, 424, 432, 450, 634 Perisincirra buitkampi 393, 394 Perisincirra filiformis 392 Perisincirra gellerti 388, 391, 424, 647 Perisincirra gracilis 388, 392 Perisincirra heterocirrata 640 Perisincirra inquieta 403 Perisincirra interrupta 388, 391, 432 Perisincirra kahli 393 Perisincirra octonucleata 421 Perisincirra pori 324, 344 Perisincirra quadrinucleata 419 Perisincirra similis 392, 618, 631 Perisincirra vermiculare 435 Perisincirra viridis 392, 447–450, 512 perisincirra, Amphisiella 200 perisincirra, Lamtostyla 3, 4, 164, 166K, 197, 199, 200, 220, 226T, 329, 341 perisincirra, Tachysoma 165, 196, 200 peritrich(s) 28 Petalotricha 41 Phacodinium 29, 31, 34, 36, 40 piscis, Uroleptus 447, 617 Pleurotricha 46, 675 Pleurotricha grandis 675 Pleurotricha setifera 139 plurinucleata, Anteholosticha 639K polycirrata, Amphisiella 90, 245, 285 polycirratus, Uroleptoides 4, 90, 226T, 246K, 285, 539 Polyhymenophora 24 polynucleata, Hemisincirra 391, 618, 634 polynucleata, Urosoma 634 polynucleatum, Hemiurosoma 4, 619K, 628, 634 Ponturostyla 21, 36, 78K, 651, 672 Ponturostyla enigmatica 4, 77, 651, 673, 681T pori, Hemisincirra 323, 344 pori, Lamtostylides 4, 226T, 324K, 344 pori, Perisincirra 324, 344 procera, Amphisiella 90, 165, 183
SYSTEMATIC INDEX procera, Bistichella 4, 533, 534K, 535, 547, 550T, 554, 555, 661 procera, Lamtostyla 1, 4, 90, 164, 167K, 183, 226T, 273 procerus, Pseudouroleptus 534, 535, 547, 661 Prodiscocephalus 25, 31 Prodiscocephalus borrori 44 Protocruzia 23 Protogastrostyla 137, 138, 690 Protogastrostyla pulchra 683, 691T, 693 Psammocephalus faurei 33, 89 Psammocephalus lithophora 89 Psammomitra 49–51, 469, 471, 473, 512 Psammomitra retractilis 469 Pseudoamphisiella 118 Pseudoamphisiella lacazei 117 pseudocrassum, Strongylidium 83 pseudokeronopsid(s) 40 Pseudokeronopsidae 477 Pseudokeronopsinae 30, 65 Pseudokeronopsis 47 Pseudokeronopsis qingdaoensis 47 Pseudokeronopsis rubra 279 Pseudorurostyla levis 39 Pseudostrombidium 38 Pseudouroleptus xii, 18, 21, 46, 49, 50, 77K, 79, 83, 151, 289, 293, 307, 312, 469, 470, 533, 535, 548, 554, 557, 651, 653, 658 Pseudouroleptus buitkampi 535 Pseudouroleptus caudatus 4, 77, 289, 293, 308, 469, 533, 554, 653, 658, 660, 661 Pseudouroleptus caudatus caudatus 662, 662K, 663, 667T, 669, 671 Pseudouroleptus caudatus namibiensis 312, 662, 663K, 667T, 668 Pseudouroleptus humicola 557 Pseudouroleptus procerus 534, 535, 547, 661 Pseudouroleptus terrestris 534, 554, 557, 661 Pseudourostyla 36, 675, 682 Pseudourostyla cristata 39 Pseudourostyla nova 636 Psilotricha succisa 117 pulchra, Anteholosticha 28, 638K pulchra, Gastrostyla 84, 137, 140, 693 pulchra, Gastrostyla Gastrostyla 138 pulchra, Gastrostyla Stylonychia 137 pulchra, Maregastrostyla 4, 9, 36, 72, 137, 147T, 153, 693 pulchra, Protogastrostyla 683, 691T, 693 pulchra, Stilonichia 137, 690 pullaster, Holosticha 28 pyriformis, Tetrahymena 53
733
qingdaoensis, Hemiamphisiella 310 qingdaoensis, Hemiamphisiella terricola 5, 245, 290T, 295, 296K, 310 qingdaoensis, Pseudokeronopsis 47 qingdaoensis, Uroleptoides 245, 294, 295, 308 quadricornutus, Onychodromus 468 quadrinucleata, Amphisiella 90, 165, 190 quadrinucleata, Erimophrya 4, 516K, 517, 577, 578, 579K, 583T, 592, 593, 595, 614 quadrinucleata, Hemiamphisiella 4, 8, 10, 187, 190, 245, 289, 290T, 294K, 318, 562, 562K, 563, 564, 567T quadrinucleata, Hemisincirra 4, 193, 245, 391, 392K, 396T, 418, 419, 449K, 454 quadrinucleata, Lamtostyla 4, 90, 164, 166K, 187, 190, 193, 226T quadrinucleata, Perisincirra 419 quadrinucleata, Uroleptoides 245, 318 quadrinucleatus, Uroleptoides 294, 318, 320 radiosa, Mitra 469, 512 randani, Anteholosticha 638K, 639K raptans, Amphisiella 90, 245, 252, 253 raptans, Lamtostyla 4, 164, 166K, 206, 212, 218, 220, 221, 226T raptans, Tachysoma 165, 221 raptans, Uroleptoides 4, 90, 246K, 249, 252, 255 rariseta, Hemisincirra 5, 389, 391, 393K, 396T, 428, 599, 600, 607 retractilis, Psammomitra 469 Retroextendia 40 Rigidocortex 46 Rigidothrix 1, 29, 38, 65 Rimostrombidium 39 Rootletphorida 89 rostrata, Trachelostyla 4, 385, 387, 476, 478, 478K, 481, 483, 498 rubra, Pseudokeronopsis 279 Rubrioxytricha 46 schiffmanni, Wallackia 470 scintillans, Glaucoma 53 scutellum, Anteholosticha 639K scutum, Diophrys 33, 35, 40 setifera, Caudiholosticha 103, 104 setifera, Gastrostyla 139 setifera, Holosticha 103 setifera, Pleurotricha 139 setigera, Oxytricha 466 sigmoidea, Anteholosticha 637K similis, Hemisincirra 633 similis, Hemiurosoma 4, 587, 614, 619K, 622, 631, 633 similis, Parabirojimia 690
734
SYSTEMATIC INDEX
similis, Perisincirra 392, 618, 631 similis, Urosoma 587, 631 simplex Trachelostyla 509 simplex, Spirotrachelostyla 4, 384, 503K, 506, 509 simplex, Stichotricha 503, 509 simplex, Strichotricha 510 simplex, Stychotricha 510 simplex-group, Spirotrachelostyla 506 sp., Hemicinsirra 388 sp./spec., Trachelostyla 4, 382, 383K, 384, 386, 475, 477, 480, 481, 485, 489, 501T spathidiids 36 Spathidium 33, 34 spec./sp., Uroleptus 42, 394, 395, 661 spectabilis, Neokeronopsis 18 Spetastyla 140 Spetastyla mystacea 140 Spetastyla mystacea minima 140 Spetastyla mystacea mystacea 140 Spetastyla mystacea, Gastrostyla 468 Spetastyla, Gastrostyla 138, 468 sphagni, Anteholosticha 638K spiralis, Spirotrachelostyla 5, 75, 503K, 503 spiralis, Trachelostyla 478, 502–504 Spiretella 38 Spiroamphisiella 21, 72K, 78K, 84, 90, 148, 149T, 445, 472, 473, 478, 503, 510 Spiroamphisiella hembergeri 4, 17, 72, 140, 148, 149T, 150, 160T, 384, 506 Spirofilidae 148, 440, 442 Spirotrachelostyla 384, 473, 473K, 478, 502 Spirotrachelostyla simplex 4, 384, 503K, 506, 509 Spirotrachelostyla simplex-group 506 Spirotrachelostyla spiralis 5, 75, 503K, 503 Spirotrachelostyla tani 4, 384, 502, 503K, 504, 506, 507T, 510, 689 Spirotricha 23–25, 37, 42 Spirotrichea 24 spirotrichoides, Trachelostyla 475 spirotrichs xi, 3, 24, 30, 33, 35, 38, 39, 42 Sporadotrichina 473 Steinia 46, 603 steinii, Gastrostyla 82, 84, 138–140, 151, 390 stenocephala, Gastrostyla 151, 383 stenocephala, Hemigastrostyla 140, 383, 472 stenocephala, Oxytricha 141 Sterkiella 25, 46 Stichochaeta 476, 477, 480, 481 Stichochaeta cornuta 477 Stichochaeta corsica 479, 482, 483, 485, 492 Stichochaeta pediculiforme 493 Stichochaeta pediculiformis 4, 470, 474, 475, 477, 478, 500
Stichochaeta pediculiformis, Trachelostyla 479, 480 Stichotricha 384, 472, 476, 477, 481, 501, 503, 509, 510 Stichotricha simplex 503, 509 Stichotrichia 24, 38, 391, 691 Stichotrichida 24 stichotrichines 619 Stilonichia pulchra 137, 690 stramenticola, Orthoamphisiella 373, 468 stramenticola, Territricha 470 strenuum, Gonostomum 690 Strichotricha simplex 510 strobilidiids 117 Strobilidium 25, 30 Strobilidium gyrans 30 Strobilidium velox 41 Strombidiida 38 strombidiids 117 Strombidium 25, 38 strongylidids 83 2 Strongylidium 83, 149, 297, 311, 315, 445, 503 3 Strongylidium crassum 297 Strongylidium granuliferum 294, 315 Strongylidium mucicola 445 Strongylidium muscorum 289, 294, 296, 297, 311 Strongylidium pseudocrassum 83 Strongylidium wilberti 294, 311 Stychotricha simplex 510 Stylonychia 17, 23, 25, 33, 34, 38, 43, 46, 137, 489 Stylonychia lemnae 42 Stylonychia mytilus 482 Stylonychia pulchra, Gastrostyla 137 Stylonychinae 1, 29–31, 44, 46, 47, 65, 82, 138–140, 467, 468, 653 stylonychine(s) 7, 30, 38, 44, 45, 65, 138, 139, 390, 653, 675 Styxophrya 46, 468 succisa, Psilotricha 117 suctorian(s) 51 sylvatica, Caudiholosticha 120 sylvatica, Erimophrya 4, 578, 579K, 583T, 590, 595 szaboi, Hemigastrostyla 472 Tachysoma 201, 220, 222, 348 Tachysoma bicirratum 200 Tachysoma furcata 200 Tachysoma hyalina 324, 347 Tachysoma hyalinum 348 Tachysoma longa 165, 218 Tachysoma pellionellum 5, 120 Tachysoma perisincirra 165, 196, 200 Tachysoma raptans 165, 221
SYSTEMATIC INDEX tani, Spirotrachelostyla 4, 384, 502, 503K, 504, 506, 507T, 510, 689 tani, Trachelostyla 478, 503, 506 Tericirra mutsusakai 448 terrestris, Pseudouroleptus 534, 554, 557, 661 terrestris, Bistichella 4, 533, 534K, 548, 550T, 552, 554, 557, 661 terrestris-group, Bistichella 548 Terricirra 51, 77K, 164, 166, 323, 339, 389, 391, 392, 418, 420, 421, 447, 512 Terricirra livida 5, 9, 448, 449K, 458, 460T Terricirra matsusakai 4, 9, 193, 420, 448, 449, 449K, 450, 454, 458, 460T Terricirra viridis 4, 9, 74, 420, 422, 448, 449K, 450, 458, 460T terricola qingdaoensis, Hemiamphisiella 5, 245, 290T, 295, 296K, 310 terricola terricola, Hemiamphisiella 5, 8, 289, 290T, 293–295, 296K, 296, 311 terricola, Amphisiella 90, 245, 279 terricola, Hemiamphisiella 72, 288, 289, 294K, 294, 311, 312, 320–322, 352, 442, 562, 660, 690 terricola, Hemiamphisiella terricola 5, 8, 289, 290T, 293–295, 296K, 296, 311 terricola, Hemiurosoma 4, 77, 614, 616T, 618, 619K, 619 terricola, Uroleptoides 4, 90, 225, 226T, 246K, 279, 285 Territricha xi, 26, 46, 50, 470 Territricha stramenticola 470 Tetmemena 46 tetracirrata, Cyrtohymena 464 Tetrahymena 33, 34, 38 Tetrahymena pyriformis 53 Tetrastyla 77K, 463 Tetrastyla oblonga 4, 74, 75, 463 Tetrastyla oblonga, Amphisiella 463 Thigmokeronopsis 36, 47 thiophaga, Holosticha 90 thiophaga, Holosticha Amphisiella 130 thononensis, Anteholosticha 639K tintinnid(s) 28, 41 Trachelochaeta 49, 50, 470, 477 Trachelochaeta bryophila 470 Trachelostya 475 Trachelostyla xii, 12, 40, 47, 50, 51, 141, 383, 384, 390, 448, 470–473, 473K, 474, 502, 503, 506, 510, 685, 688 Trachelostyla affine 476 Trachelostyla bryonicolum 475 Trachelostyla canadiensis 478
735
Trachelostyla caudata 5, 383, 475, 476, 478, 478K, 494, 501, 502, 511, 689 Trachelostyla ciliophorum 475 Trachelostyla dubia 382, 383, 478 Trachelostyla geleii 475 Trachelostyla macrostoma 475 Trachelostyla pediculariformis 480 Trachelostyla pediculiforme 480 Trachelostyla pediculiformis 5, 12, 18, 26, 75, 383, 384, 386, 387, 390, 476, 478K, 478, 494–498, 500, 501T, 502, 505, 683, 687 Trachelostyla pendiculiformis 480 Trachelostyla rostrata 4, 385, 387, 476, 478, 478K, 481, 483, 498 Trachelostyla simplex 509 Trachelostyla sp./spec. 4, 382, 383K, 384, 386, 475, 477, 480, 481, 485, 489, 501T Trachelostyla spiralis 478, 502–504 Trachelostyla spirotrichoides 475 Trachelostyla Stichochaeta pediculiformis 479, 480 Trachelostyla tani 478, 503, 506 trachelostylid(s) 4, 10, 16, 24, 51, 52, 75, 448, 471 Trachelostylidae xii, 1, 12, 24, 46, 48, 51, 71, 72K, 163, 390, 448, 470, 471 Traxhelostyla pediculiformis 475 Trichoda musculus 443 turanica, Amphisiella 4, 89, 91K, 109, 117, 123 uroleptid(s) 26 Uroleptoides 19, 49–51, 77K, 81, 88, 161, 165, 166, 180, 183, 187, 224, 294, 310, 318, 320, 352–354, 372, 376, 377, 655 Uroleptoides atypica 466, 653–655 Uroleptoides atypicus 245, 651, 652, 654, 655 Uroleptoides binucleata 261, 262, 264 Uroleptoides binucleatus 72, 180, 183, 246K, 249, 253, 261, 273 Uroleptoides binucleatus binucleatus 4, 9, 226T, 263K, 264, 267 Uroleptoides binucleatus multicirratus 4, 9, 89, 226T, 251, 264K, 264, 267 Uroleptoides caudata 245, 352, 358, 359 Uroleptoides caudatus 359 Uroleptoides kihni 5, 88, 224, 244, 245, 246K, 246, 294, 352, 372–374 Uroleptoides longiseries 4, 9, 90, 193, 226T, 246K, 249 Uroleptoides magnigranulosus 4, 9, 47, 90, 182, 183, 226T, 246K, 249, 255, 263, 267, 272, 273, 390 Uroleptoides multinucleatus 5, 9, 77, 90, 226T, 246K, 251, 253, 255, 272
736
SYSTEMATIC INDEX
Uroleptoides polycirratus 4, 90, 226T, 246K, 285, 539 Uroleptoides qingdaoensis 245, 294, 295, 308 Uroleptoides quadrinucleata 245, 318 Uroleptoides quadrinucleatus 294, 318, 320 Uroleptoides raptans 4, 90, 246K, 249, 252, 255 Uroleptoides terricola 4, 90, 225, 226T, 246K, 279, 285 Uroleptoides vitiphila 165, 187, 245 Uroleptoides vitiphilus 187 Uroleptopsis 225 Uroleptopsis citrina 118, 225 Uroleptus xi, 25, 42, 44, 46, 47, 65, 82, 224, 394, 531, 659, 691 Uroleptus agilis 513, 530 Uroleptus hospes 440, 441 Uroleptus humicola 534, 556, 557, 661 Uroleptus kahli 389, 391, 393, 394, 450, 458, 512, 530 Uroleptus natronophilus 515, 530, 531 Uroleptus piscis 447, 617 Uroleptus spec./sp. 42, 394, 395, 661 Uronychia 25, 28, 31, 33, 36, 41, 43 Urosoma 13, 41, 45, 46, 51, 78K, 389, 437, 439, 471, 473, 513–515, 560, 564, 599, 614, 615, 618, 618K, 619, 623, 625, 630, 633, 634 Urosoma caudata 631 Urosoma cienkowskii 513, 631 Urosoma octonucleata 614 Urosoma polynucleata 634 Urosoma similis 587, 631 Urosomoida 45, 46, 178, 471, 473, 513–515, 530, 531, 566, 577, 589, 596, 599, 603, 615, 619, 634 Urosomoida agiliformis 579, 581, 582 Urosomoida agilis 587 Urosomoida antarctica 578 Urosomoida deserticola 578 Urosomoida minima 515, 516K, 517 Urosomoida monostyla 578 Urosomoides 51, 513 Urospinula 1, 29 Urostyla 25, 618, 672 Urostyla grandis 41, 42 Urostyla viridis 618 Urostyla weissei 469 Urostylida 24 Urostylidae 24, 81 Urostylina 24 urostyloid(s) xii, 3, 12, 13, 15, 17–20, 24–26, 28, 29, 35, 38, 39, 42, 42, 44, 45, 47, 51, 65, 66, 71,
80–82, 88, 94, 119, 163, 390, 391, 445, 469, 470, 476, 477, 560, 564, 569, 615, 636, 648, 661, 675 Urostyloidea xi, 12, 24, 29, 39, 40, 46, 77, 88, 512, 636, 643 variabilis, Cladotricha 516, 517, 529 velox, Strobilidium 41 vermiculare, Hemisincirra 435 vermiculare, Perisincirra 435 vermicularis, Hemisincirra 5, 389, 391, 392K, 396T, 428, 435, 599 Vermioxytricha 46, 71, 78K, 389, 391, 392, 428, 462, 515, 560, 578, 579, 596, 615, 618 Vermioxytricha arenicola 5, 9, 77, 391, 428, 596, 597T, 599, 600, 607 Vermioxytricha muelleri 5, 428, 597T, 598–600, 600K, 605, 607 verrucosa, Anteholosticha 4, 393K, 396T, 402, 424, 426, 427, 636, 640K, 647 verrucosa, Hemisincirra 647 verrucosa, Hemisincirra gellerti 391, 424, 647, 648 vettersi, Circinella 428 vettersi, Hemisincirra 391 violacea, Anteholosticha 637K, 639K viridis, Banyulsella 466 viridis, Hemisincirra 450 viridis, Paraurostyla 618 viridis, Perisincira 450 viridis, Perisincirra 392, 447–450, 512 viridis, Terricirra 4, 9, 74, 420, 422, 448, 449K, 450, 458, 460T viridis, Urostyla 618 vitiphila, Amphisiella 187 vitiphila, Lamtostyla 4, 164, 166K, 187, 190, 193, 226T, 245, 320 vitiphila, Uroleptoides 165, 187, 245 vitiphilus, Uroleptoides 187 vuxgracilis, Anteholosticha 637K Wallackia 46, 50, 470, 472, 473 Wallackia schiffmanni 470 warreni, Anteholosticha 639K weissei, Paraurostyla 42, 535, 673 weissei, Urostyla 469 wenzeli, Hemisincirra 5, 8, 391, 393K, 394, 396T, 408, 424, 437 wilberti, Hemiamphisiella 4, 289, 290T, 294K, 311, 315, 660, 662, 671 wilberti, Strongylidium 294, 311 xanthichroma, Anteholosticha 637K, 638K
Table Index Table 1. . . . . . . . . . . . . . Table 2. . . . . . . . . . . . . . Table 3. . . . . . . . . . . . . . Table 3a. . . . . . . . . . . . . Table 4. . . . . . . . . . . . . . Table 5. . . . . . . . . . . . . . Table 6. . . . . . . . . . . . . . Table 7. . . . . . . . . . . . . . Table 8. . . . . . . . . . . . . . Table 9. . . . . . . . . . . . . . Table 10. . . . . . . . . . . . . Table 11. . . . . . . . . . . . . Table 12. . . . . . . . . . . . . Table 13. . . . . . . . . . . . . Table 14. . . . . . . . . . . . . Table 15. . . . . . . . . . . . . Table 16. . . . . . . . . . . . . Table 17. . . . . . . . . . . . . Table 18. . . . . . . . . . . . . Table 19. . . . . . . . . . . . . Table 20. . . . . . . . . . . . . Table 21. . . . . . . . . . . . . Table 22. . . . . . . . . . . . . Table 23. . . . . . . . . . . . . Table 24. . . . . . . . . . . . . Table 25. . . . . . . . . . . . . Table 26. . . . . . . . . . . . . Table 27. . . . . . . . . . . . . Table 28. . . . . . . . . . . . . Table 29. . . . . . . . . . . . . Table 30. . . . . . . . . . . . . Table 31. . . . . . . . . . . . . Table 32. . . . . . . . . . . . . Table 33. . . . . . . . . . . . . Table 34. . . . . . . . . . . . . Table 35. . . . . . . . . . . . . Table 36. . . . . . . . . . . . . Table 37. . . . . . . . . . . . . Table 38. . . . . . . . . . . . . Table 39. . . . . . . . . . . . . Table 40. . . . . . . . . . . . . Table 41. . . . . . . . . . . . . Table 42. . . . . . . . . . . . . Table 43. . . . . . . . . . . . . Table 44. . . . . . . . . . . . . Table 45. . . . . . . . . . . . .
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