Süßwasserflora von Mitteleuropa, Bd. 21 1 Freshwater Flora of Central Europe, Vol. 21 1: Fungi: 1. Teil 1st Part: Lichens (German and English Edition)

Holger Thüs / Matthias Schultz Fungi · 1. Teil / 1st Part: Lichens

6‰ZDVVHUÀRUD von Mitteleuropa Freshwater Flora of Central Europe Begründet von / Founded by A. Pascher Herausgegeben von / Edited by Burkhard Büdel · Georg Gärtner Lothar Krienitz · Hans-Rudolf Preisig Michael Schagerl

Band / Volume 21/1

Holger Thüs Matthias Schultz Fungi 1. Teil / 1stt Part: Lichens

Fungi 1. Teil / 1stt Part: Lichens Holger Thüs Matthias Schultz

171 Abbildungen / Figures

Authors’ Adresses Dr. Holger Thüs, Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom of Great Britain Dr. Matthias Schultz %LR]HQWUXP .OHLQ )ORWWEHN 6\VWHPDWLN GHU 3ÀDQ]HQ Ohnhorststr. 18, 22609 Hamburg, Germany Notice for the reader The Authors, the Editors, and the Publisher of this work have made every effort to ensure that the information herein is accurate and in accord with the standards accepted at the time of publication. To the extent permissible under applicable laws, no responsibility is assumed by Spektrum Akademischer Verlag GmbH for any injury and/or damage to persons or property as a result of any actual or alleged libellous statements, infringement of intellectual property or privacy rights, or products liability, whether resulting from negligence or otherwise, or from any use or operation of any ideas, instructions, procedures, products or methods contained in the material therein. The Publishers have made an extensive effort to trace original copyright holders for permission to use borrowed material. If any has been overlooked, it will be FRUUHFWHGDWWKH¿UVWUHSULQW Bibliographic information published by Die Deutsche Nationalbibliothek Die Deutsche Nationalbibliothek lists this publication in the Deutsche NationalELEOLRJUD¿H Detailed bibliographic data is available in the internet at http://dnb.d-nb.de. Springer is a part of Springer Science+Business Media springer.de © Spektrum Akademischer Verlag Heidelberg 2009 Spektrum Akademischer Verlag is an Imprint of Springer 09 10 11 12 13

5 4 3 2 1

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Acknowledgements &KULVWLQH.HOOHU%LUPHQVGRUI has PXFKLQÀXHQFHGWKHFRQFHSWLRQRIWKLVERRN DQGVKHIXUWKHUVWLPOXDWHGLWVSURJUHVVE\MRLQW¿HOGZRUNDQGGLVFXVVLRQVRIYDULRXV parts of the manuscript. We are grateful to the curators of B, BM, FR, G, J, H, HBG, KR, M, POL, PRM, S, STU, TU, UPS and W for quickly handling loan UHTXHVWV :H DUH IXUWKHU LQGHSWHG WR %XUNKDUG %GHO .DLVHUVODXWHUQ  )UDQ] %HUJHU.RS¿QJ 3LHWHUYDQGHQ%RRP6RQ %ULDQ&RSSLQV(GLQEXUJK  Rainer &H]DQQH'DUPVWDGW +HOHQH *HUKDUG&]HLND9LHQQD 5KLQDL[D 'XTXH7KV /RQGRQ 2OLYHU'UKDPPHU5HJHQVEXUJ 0DULRQ(LFKOHU'DUPVWDGW &HFLOH *XHLGDQ8WUHFKW 0DUWLQ*UXEH*UD] $QQD*XWWRYi%UDWLVODYD 3HU0DJQXV -¡UJHQVHQ%HUJHQ %HDWD.U]HZLF]ND.UDNyZ +HOPXW0D\UKRIHU *UD] -XUL 1DVFLPEHQH7ULHVWH $ODQ2UDQJH&DUGLII =GHQHN3DOLFH3UĤKRQLFH $NLUD 3HWHUV5RVFRII 8OI6FKLHIHOEHLQ8HFNHUPQGH 5RPDQ7UN6DO]EXUJ DQG 9RONPDU:LUWK 0XUU  IRU VXSSO\LQJ XV ZLWK YDOXDEOH ¿HOG REVHUYDWLRQV DGGLtional study material, additional photographs and critical remarks on earlier verVLRQVRIWKHPDQXVFULSW5KLQDL[D'XTXH7KV/RQGRQ IXUWKHUFRQWULEXWHGE\ tracing literature as well as by digitizing and improving the line drawings. Studies of H. Thüs for this book were supported by a grant from the German Research )RXQGDWLRQ7+ DQGE\WKH6<17+(6<S Project http://www.synthesys. LQIR ZKLFK LV ¿QDQFHG E\ WKH (XURSHDQ &RPPXQLW\ 5HVHDUFK ,QIDVWUXFWXUH Action under the FP6 “Structuring the European Area” Programme. We would also like to thank our families for their patience and lasting support.

Contents Acknowledgement . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

V

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

1

Glossary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

14

General Key . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

25

Keys to the Species and Species PrR¿OHVLQDOSKDEHWLFRUGHU . . . . . . . . . . . Anisomeridium 0OOHU$UJRYLHQVLV 0&KRLV\ . . . . . . . . . . . . . . . . Aspicilia A. Massalongo 1852 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bacidina 9Č]GD . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caloplaca Th. Fries 1860 nom. cons. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Collema Weber ex F.H. Wiggers 1780 nom. cons .. . . . . . . . . . . . . . . . . . . Dermatocarpon Eschweiler 1824 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Endocarpon Hedwig 1789 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ephebe Fries 1825. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gonohymenia J. Steiner 1902 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gyalidea /HWWDXH[9Č]GD . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydropunctaria *XHLGDQ.HOOHU 7KV . . . . . . . . . . . . . . . . . . . . . Hymenelia Krempelhuber 1852 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ionaspis Th. Fries 1871. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Koerberiella Stein 1879 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lecidea Acharius 1803 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lempholemma Körber 1855 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leptogium $FKDULXV *UD\ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lobothallia &ODX]DGH &O5RX[ +DIHOOQHU . . . . . . . . . . . . . . . . . . Phaeophyscia Moberg 1977 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Placynthium $FKDULXV *UD\ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polyblastia A. Massalongo 1852 nom. cons. . . . . . . . . . . . . . . . . . . . . . . . Porina Müller Argoviensis 1883 nom. cons. . . . . . . . . . . . . . . . . . . . . . . . Porocyphus Körber 1855 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Porpidia Körber 1855 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudarthopyrenia Keissler 1935 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterygiopsis Vainio 1890 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pyrenocarpon Trevisan 1855 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pyrenocollema Reinke 1895 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pyrenopsis Nylander 1858 nom. cons. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhizocarpon de Candolle 1805. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rinodina $FKDULXV *UD\ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sporodictyon A. Massalongo 1852 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Staurothele Norman 1853 nom. cons. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thelidium A. Massalongo 1855 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thelignya A. Massalongo 1855 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Verrucaria Schrader 1794 nom. cons.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Literature . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Index of Lichen Names . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Index to Figures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

35 35 36 40 42 43 49 54 56 59 61 63 68 68 71 72 73 75 82 83 84 92 95 100 104 107 109 111 113 116 118 122 124 125 135 155 156 187 195 205

Colour Plates 1–6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

211

Introduction Amphibious lichens are widely distributed but little recognized components of freshwater vegetation. In most assessments of freshwater organisms however, lichens are still neglected which is partly due to the fact that corresponding literature is widely scattered and a profound treatment of these lichens in Central Europe is still missing. This book is intended to provide a comprehensive tool for the determination of freshwater lichens in Central Europe for both, the non-specialized limnologist and the interested lichenologist. It further summarizes basic information on ecology and distribution. The book roughly covers an area ranging from the Alps to the south shores of the North and Baltic Sea and from the Western Carpathians in the east to the Vosgues-Mountains in the west. Amphibious lichens are found in springs, streams and rivers wherever hard and stable VXEVWUDWDDVZHOODVVXI¿FLHQWOLJKWDUHJUDQWHG,QFHUWDLQVWUHDPVRIFRROHUXSland areas, lichens can represent the most diverse group of organisms covering virtually any rock surface (e.g. Frey 1922). On the other hand, they are very rarely found in lakes. Rock surfaces incrusted by silt, surface erosion by sediment particles or water with a pH below 5 can exclude any growth of lichens. Resulting IURP WKDW IUHVKZDWHU OLFKHQV DUH YDOXDEOH ORQJWHUP LQGLFDWRUV RI WKH VSHFL¿F structure of a given watercourse, as well as they indicate key factors of water chemistry (Nascimbene et al. 2007). Water pollution was – and certainly still is – the main reason for the dramatic decline of lichen growth especially along larger watercourses during the past decades. However, increasing water quality and reduced acidic impacts have enabled some species to recover even along the banks of large Central European rivers as for example the Rhine (Thüs 2006). Consequently, the potential use of lichens for bioindication purposes is regarded very high (Aptroot & Seaward 2003, Nascimbene et al. 2007, Thüs 2002). Unfortunately, the knowledge on ecology and indicator values of many freshwater liFKHQVDQGEDVLFÀRULVWLFLQIRUPDWLRQWKDWZRXOGDOORZWRHYDOXDWHZKHWKHUDFHUtain species is endangered or not are still sketchy. Therefore, corresponding notes KDYHEHHQPDGHRQO\LQFDVHVZKHUHVXI¿FLHQWGDWDZHUHDYDLODEOH,QVSLWHRI this, we hope that the present book will stimulate the recognition of freshwater lichens and promote the use of these organisms in matters of nature conservation. Lichens do not form a natural group of organisms (Liu & Hall 2004, Lutzoni et al. 2001) but represent an exceedingly delightful lifestyle in higher fungi, characterized by the symbiotic association with one or more photosynthetic partner(s) which supply the lichenized fungus with organic metabolites (Honegger1998). More than 2000 species of lichen forming fungi occur in Central Europe, but only a much smaller number is found in and along watercourses. The lichen symbiosis is a nutrition strategy with an excellent performance under conditions characterized by frequent changes in water availability, the total lack of liquid water or extended periods of desiccation (Lange 2003a, b, Lange & Green 1997, Ried 1960a). On the other hand, ecosystems with permanent supply of liquid water are rather poor in lichen species (Ried 1960b). In Central Europe, roughly a dozen taxa of lichens are able to survive under permanent submersion. In the splash water zone of watercourses and in temporarily inundated micro-sites the species number is much higher, reaching more than 100 taxa in upland areas.

2

Introduction

,QWKH¿HOGLWLVRIWHQGLI¿FXOWWRDVVHVVZLWKDVLQJOHYLVLW ZKHWKHUDVWUHDPRU small river is permanent or ephemeral, and in large rivers the water level can FKDQJHVHYHUDOPHWHUV'XHWRWKHVSHFL¿FHFRORJLFDOQHHGVRILQGLYLGXDOOLFKHQ species, the water regime of such water bodies can be assessed by analysing the composition of the lichen communities at a given site. The inundation-dependent zonation of the lichen vegetation can provide important information for the prediction of regularly occuring high water levels as well as for the assessment of H[WUHPH ÀRRGZDWHUV LQ WKH SDVW %HVFKHO  +DOH  0DUVK  7LPRQH\ 2005, Rosentreter 1984, Timoney & Marsh 2004, 2005). Furthermore, the analysis of amphibious lichen vegetation provides valuable information on physicochemical properties of both, the water (Pentecost 1977, Nascimbene et al. 2007) DQGWKHVXUURXQGLQJDLUERXQGLQÀXHQFHV7KV 7KHLQFOXVLRQRIOLFKHQVLQ the assessment of benthic communities is thus highly recommended for applied issues in nature conservation and ecology – but has received little interest among limnologists so far. The common practice of neglecting amphibious lichens in HFRORJLFDOVWXGLHVLVPDLQO\GXHWRWKHODFNRIDFRPSUHKHQVLYHÀRULVWLFWUHDWPHQW of these organisms and the fact that amphibious lichen communities are often composed by representatives of taxonomically highly problematic genera. 7KLV ERRN LV LQWHQGHG DV D WRRO IRU WKH LGHQWL¿FDWLRQ RI DOO OLFKHQL]HG IXQJL LQ Central Europe which can be called freshwater lichens, either because they are adapted to freshwater habitats or because they are frequently observed in or at streams and rivers due to their exceptionally wide ecological amplitude. While WKH¿UVWJURXSLVUHDVRQDEO\GH¿QHGWKHVHFRQGJURXSLVGLI¿FXOWWRGHOLPLWDQG may also be described as facultative amphibious species. Many lichens can be found occasionally at the banks of watercourses at temporarily inundated microsites. Especially on carbonaceous rocks it may prove impossible to distinguish between hydrophilic communities (amphibious, regularly submersed or strongly LQÀXHQFHGE\VSODVKZDWHU RQWKHRQHKDQGDQGK\JURSKLORXVFRPPXQLWLHVSUHferring humid sites, with either high air humidity or frequent access to liquid water) on the other. Local characteristics can strongly affect the composition of hygrophilous lichen communities, especially in the less frequently inundated zones. The composition of the freshwater lichen vegetation at a certain site furthermore largely depends on macroclimatic factors. Some species that are frequently found inundated in Northern or Western Europe, expose a less pronounced hydrophytic performance in Central Europe (compare species lists and lichen communities described by Gilbert 1996, Gilbert & Giavarini 1997 for the British Isles; Santesson 1939, Nordin 2002a for Sweden; Beschel 1954, Frey 1921, Keller 2000, Motyka 1926, Ried 1960b, Scheidegger & Keller 1994 and Thüs 2002 for Central Europe). In the Mediterranean region, high summer temperaWXUHVDQGSHULRGLFGURXJKWSHULRGVUHVXOWLQDVSHFL¿FFRPSRVLWLRQRIWKHOLFKHQ ÀRUDLQVWUHDPVHVSHFLDOO\DWORZDOWLWXGHV&RQVHTXHQWO\WKH OLFKHQYHJHWDWLRQ along watercourses in the Mediterranean differs considerably from that in Central Europe (Pereira 1992). In Central Europe, freshwater lichens are best developed LQ UDSLGO\ ÀRZLQJ ZDWHU ZKHUHDV LQ VWDJQDQW ZDWHU VXFK DV DORQJ URFN\ ODNH shores the species number remains low (e.g. Erichsen 1957). In Nordic countries and the British Isles instead, lake shores are important habitats for freshwater lichens (Gilbert & Giavarini 2000, Santesson 1939).

Introduction

3

The present book comprises all specialized aquatic lichens occurring in Central Europe. However, it proved impossible to key out every species that potentially occurs among the facultative “guests” in freshwater ecosystems. Therefore, only WKHPRVWIUHTXHQWRQHVDUHLQFOXGHGLQWKHNH\V,IWKHXVHU¿QGVPRUHIDFXOWDWLYH freshwater lichens than more or less specialized amphibious species, it can be concluded that the watercourse under study is either non-permanent or the colOHFWLQJVLWHLVQRWPXFK LQÀXHQFHGE\VSODVKZDWHURU periodic inundation. The socalled “riparian” lichens are typically found in the vicinity of large water bodies, EXWRXWVLGHWKHGLUHFWLQÀXHQFHRILQXQGDWLRQRUZDWHUVSUD\7KHVHOLFKHQVEHQH¿WIURPPRUHIUHTXHQWIRJHYHQWVDVZHOODVIURPLQFUHDVHGGHZIDOODURXQGODUJH ZDWHUERGLHVEXWWKH\DUHQRWVSHFL¿FDOO\DGDSWHGWRDQDPSKLELRXV OLIH VW\OH and therefore the present book offers only limited information on these lichens. )RUWKHLGHQWL¿FDWLRQRIULSDULDQDVZHOODVIRUIDFXOWDWLYHO\ DPSKLELRXVOLFKHQV the user is refered to comprehensive determination books such as Wirth (1995), Clauzade & Roux (1985) or Purvis et al. (1992).

Ecology of Freshwater Lichens Inundation tolerance versus inundation demand Most freshwater lichens are inundation tolerant to a certain degree, but only a small number is really inundation demanding and appears to be highly sensitive to complete desiccation (Ried 1960a )LHOGREVHUYDWLRQVKRZHYHUFRQ¿UPWKDW even lichens typically found in permanently inundated communities (Verrucaria aquatilis, V. funckii, Hydropunctaria rheitrophila) are able to survive limited periods of desiccation up to several weeks (!) at shaded micro-sites with high air humidity. Obviously, the circumstances under which desiccation occurs decide about the survival of the lichens from usually permanently submersed sites. Aptroot & Seaward (2003) followed Santesson (1939) in his suggestion that possibly no freshwater lichen is completely restricted to submersed micro-sites and added the aspect that every freshwater lichen relies on a limited desiccation tolerance at least for the dispersal of the ascospores via the air. They concluded that the term “aquatic” should be avoided and instead proposed the term “amphibious“. Our own observations of populations of Verrucaria aquatilis, V. funckii and Hydropunctaria (Verrucaria) rheitrophila from Central Europe over almost ten years however, leave no doubt that at least for these species the complete life cycle from the establishment of young thalli to the production of perithecia and ascospores takes place at micro-sites that remain inundated even during extended heat and drought periods. Similar observations were made by Hawksworth (2000) for Collema dichotomum and Hydropunctaria (Verrucaria) rheitrophila. Therefore, we see no reason to avoid the term “aquatic” for these organisms, even if they have the principal ability to survive short periods of time without liquid water irrespective of their ability to disperse ascospores via the air. In this book we use the term “aquatic” synonymously to “growing submersed” for lichens able to grow for more than a year under permanent submersion in the whole distribution area. In contrast, the term “amphibious” is used for species that are best developed in the splash water zone or at micro-sites that are regularly subject to

4

Introduction

desiccation for at least some weeks during the year and for species that occur permanently submersed only in rare cases and under special conditions (usually only in parts of their distribution range, e.g. in cooler upland streams or Nordic regions). A third group of lichens does not have any physiological adaptation to LQXQGDWLRQEXWEHQH¿WVIURPKLJKHUDLUKXPLGLW\LQWKHYLFLQLW\RIFHUWDLQW\SHV of waterbodies. These species are referred to as “riparian” lichens. Widespread and common species in Central Europe such as Lecanora muralis fall under this category, but also rare and highly specialized taxa adapted to shaded situations with high air humidity such as Lecidea ahlesii. We have included a selection of VSHFLHVIURPWKHULSDULDQÀRUDLQWKHNH\VDVIDUDVWKHUHLVHYLGHQFHIRURFFDsional records from temporarily inundated sites. A few of the species which are rare and often neglected are presented in more detail in order to stimulate their consideration in future studies. In general, a high frequency of riparian lichens at a certain site indicates WKDWWKH\DUHQRWUHJXODUO\LQXQGDWHGDQGUHJLRQDOÀRULVWLF surveys or determination books should be consulted. Finally, “terrestrial” species are those with a generally low submergence tolerance and which are usually not found in the direct vicinity of water bodies. This group includes all lichens growing in any kind of non-aquatic ecosystem (including rocks, living or dead plants or soil) and it should not mixed up with the term “terricolous” which refers to lichens growing on soil.

Substratum stability Unlike many free living algae and mosses, freshwater lichens are rather slow growing organisms although large differences exist in growth rates among aquatic lichen species (Keller 2005). While some freshwater lichens can even be conVLGHUHGSLRQHHUVSHFLHV.U]HZLFND *DODV RWKHUVVSHFL¿FDOO\RFFXURQ immobile substrata that offer a hard and stable surface (Thüs 2002). The knowledge on the individual preferences of particular species regarding the substrate VWDELOLW\LVYHU\VNHWFK\DQGRQO\LQFDVHVZKHUHVXI¿FLHQWREVHUYDWLRQVH[LVW a corresponding note was added in the species description part of this book.

Silting tolerance Most freshwater lichens are very sensitive against deposition of silt-like sediments of either organic or inorganic origin on the thallus surface (e.g. Erichsen 1957, Gilbert 1996, Gilbert & Giavarini 1997, Thüs 2002, 2006). Possibly, the establishment of ascospores is hindered by silted substrate surfaces, but no study addressed this aspect so far. A small number of species however, appears to be tolerant to moderate silting (e.g.: Bacidina inundata, Placynthium tantaleum, PyUHQRFDUSRQÀRWRZLDQXP, Leptogium subtorulosum, Verrucaria praetermissa). In general, species with thin subgelatinous thalli (consistency like “solid jelly”, becoming ± transparent when wet) are most sensitive to silting while thick areolate VSHFLHVDUHRIWHQOHVVVHQVLWLYH7KV 6LOWLQJLVGLI¿FXOWWRTXDQWLI\DQG JHQHUDOQRWHVRQWKHVLOWWROHUDQFHRIDVSHFLHVZHUHPDGHRQO\ ZKHUHVXI¿FLHQW ¿HOGREVHUYDWLRQVH[LVW

Introduction

5

Nutrient supply Experimental studies on nutrient demands and tolerance limits to high nutrient supply are very scant (Davis et al. 2000) and therefore, generalisations are premature. Field observations suggest a higher species diversity in oligotrophic environments. In eutrophic watercourses most lichen species are easily out-competed by cyanobacteria, algae and mosses. Only a few taxa such as Verrucaria praetermissa are able to survive even in polluted streams when the current is rapid and silting does not exceed a critical level. Possibly, lichen growth is often not limited simply by eutrophication (nutrient content) but by the silting effect of detritus which is related to high degree of saproby and directly affects the lichen vegetation. Detailed studies on direct or indirect effects of eutrophication versus saproby related silting of organic matter are still missing.

Light demands The highest species diversity is usually found at well lit streams and rivers, but shade demanding species do exist (Pentecost 1977, Gilbert 1996, Gilbert & Giavarini 1997, Thüs 2002). Therefore, more or less shaded places should also be examined for a complete assessment of species richness. Shade demanding speFLHVFDQEHIRXQGIRUH[DPSOHRQWKHORZHUVLGHRUDWWKHÀDQNV RIODUJHUERXOGHUV or on small pebbles in the shade of weedy vegetation at the banks of a watercourse. Deep shade, however, is generally avoided by all freshwater lichens. Therefore, streams in dense thickenings of e.g. young spruce forests are usually completely devoid of freshwater lichens.

Preferences for acidity )UHVKZDWHUOLFKHQVKDYHYHU\VSHFL¿FUHVSRQVHVWRWKHS+YDOXHRIWKHLUKDELWDW sometimes not only of the surrounding water but also of the substratum and in case of amphibious species also of the precipitation. Permanently submersed lichens are only affected by the pH value of the water body. In contrast, amphibious lichens in the splash water zone or at micro-sites with frequent changes of the ZDWHU OHYHO DUH FRQVLGHUDEO\ LQÀXHQFHG E\ DLUERUQH DFLGLF HPLVVLRQV 7KV 2002). The underlying processes for the different ranges of pH tolerance are still poorly XQGHUVWRRG 6SHFL¿F DFLGLW\ OHYHOV PD\ KDYH GLIIHUHQW HIIHFWV RQ WKH IXQJDO RU algal partner of a lichen and certain developmental stages may also have different tolerance limits. Some general patterns however are obviously related to the kind of associated photobionts: freshwater lichens with cyanobacterial photobionts are best developed in neutral or slightly (?) alkaline waters while they are usually absent in waters below pH 5. The same pattern is observed for lichens with obligatory symbioses with 'LODEL¿OXP-like algae (Verrucaria aquatilis) or Diplosphaera chodatii and other Stichococcus-like green algae (most species of Dermatocarpon, Polyblastia, Staurothele, Thelidium, and Verrucaria). Freshwater lichens with “trebouxioid” photobionts (genera Asterochloris, Myrmecia, Trebouxia), Trentepohlia or Heterococcus tend to prefer neutral to acidic water. Species with a wider pH-tolerance (Verrucaria hydrela, Hydropunctaria rheitrophila) are able

6

Introduction

to associate with different photobionts (Hydropunctaria rheitrophila: 'LODEL¿lum, Heterococcus; Verrucaria hydrela:'LODEL¿OXP, Diplosphaera, Elliptochloris, Heterococcus  VXLWDEOH IRU WKH UHVSHFWLYH S+UDQJH LQ D VSHFL¿F HQYLURQment. Freshwater lichens are usually slow growing and – compared to other benthic cryptogams – rather long living organisms. The knowledge of these preference SUR¿OHVPDNHVLQYHQWRULHVRIIUHVKZDWHUOLFKHQYHJHWDWLRQYDOXDEOHIRUDUDSLG in situ assessment of pH-extremes occurring on the large time scale. They allow for a more integrative assessment compared to the short term indication by common indicators such as diatoms or other free living algae with a short life cycle. Furthermore, the separate analysis of the composition of lichens from submersed and amphibious sites gives a detailed impression of the acidic load transported by the water body and the air.

Interactions with other organisms and eutrophication Fungi Saprophytic or commensalistic ¿ODPHQWXRVIXQJLDUHIUHTXHQWO\JURZLQJDPRQJ lichen thalli, colonising the cavities of over-aged perithecia or the cracks between the areoles of crustose species. Only few lichenicolous fungi are known to rely on freshwater lichens as their hosts (for details see Clauzade et al. 1989, Rambold et al. 1990, Molitor & Diederich 1997, Orange 2002). The external appearance of lichens may change considerably when being infected by lichenicolous fungi, RIWHQPDNLQJDSURSHULGHQWL¿FDWLRQLPSRVVLEOH,QIHFWHGOLFKHQWKDOOLDUHW\SLcally much more cracked and have brownish to blackish coloured areas on the thallus with a brittle consistency.

Algae Epiphytic algae are often present in certain periods of the year, e.g. during mass developments of diatoms in spring. Long-living, non motile algae are much rarer on completely submersed lichens. On the other hand, the brownish cyanobacterium Oncobrysa rivularis can regularly be observed as an epiphyte on submersed Verrucaria thalli without causing any visible damage to the lichen. In alpine areas mostly reddish coloured cyanobacteria are sometimes covering crustose species of Verrucaria and Staurothele with a continuous layer. Free-living cyanobacteria with reddish sheaths are often found on moist rock and may resemble accompanying, juvenile thalli of Pyrenopsis which contain Gloeocapsa-like cyanobacteria as photobionts (= cyanobiont). Likewise, free-living Rivulariaceae can be observed growing abundantly on slack stones in the freshwater tidal zone along the river Elbe in and around Hamburg, Germany. These cyanobacteria form a blackish, irregular crust and may resemble young (or dying?) thalli of species of Porocyphus which contain Rivulariaceae as photobionts and are known to occur in the same habitat. The red alga Hildenbrandia rivularis and the lichen Hydropunctaria rheitrophila are frequently observed side by side at shady places in cooler and neutral to alkaline streams and lakes. Only rarely one of the two species is able to overgrow the other causing considerable changes in the colour of the lichen. Chlorophycean taxa such as Stichococcus spp., or Trebouxia

Introduction

7

spp. have also been observed as epiphytes on amphibious lichens. They are more frequent at low altitudes and in more or less nutrient rich waters. In the splash water zone of moderately eutrophic watercourses an epiphytic cover of lichens with a huge diversity of cyanobacteria and eukaryotic algae is rather frequent DQG FDQ FRQVLGHUDEO\ LQÀXHQFH WKH FRORXUDWLRQ DQG external appearance of the lichens.

Bryophytes Bryophytes (Mosses and Liverworts) are generally strong competitors in shaded habitats. When they start overgrowing lichens the latter will soon die, probably due to shading or allelopathic effects. Mats of pleurocarpous mosses however, are favoured substrata for some foliose lichens (Collema spp., Leptogium spp.) and the membranaceous species Lempholemma polyanthes. Shade tolerant crustose lichens can be found among the shoots of turf forming acrocarpous species.

Vascular Plants 9DVFXODU SODQWV DUH XVXDOO\ QRW FRPSHWLQJ ZLWK OLFKHQV IRU VSDFH EXW LQÀXHQFH their occurrence by shading. Roots of Black Elder (Alnus ( glutinosa) and Willow (Salix spp.) are occasionally colonized by crustose freshwater lichens. In areas where larger boulders are rare, this substratum can be the most important habitat IRU DPSKLELRXV OLFKHQV 0RWLMnjQDLWơ   (SLSK\WLF JURZWK RI IUHVKZDWHU OLchens on the roots of vascular plants is more often observed in the Alps and in 1RUWK(DVWHUQ(XURSH0RWLHMnjQDLWơ3\NNlOl EXWLVD UDUHSKHQRPenon in most areas of Central Europe at low or moderate elevations (Thüs 2002).

Animals Herbivores attack amphibious lichens preferably in the splash water zone where snails, springtails and other insects feed on algae from the thallus and the hymenium of the lichen ascomata. Determination of such thalli is often impossible and regeneration of grazed lichen thalli may result in manifold aberrations of the normal thallus appearance (“doubled” or threefold basal layer, changes in colour due to the absence or paucity of cortex, uneven surface etc.). The cushions of large, foliose lichens such as Dermatocarpon spp. may also play a role as resting sites for aquatic invertebrates in streams with strong current but detailed studies on the associated fauna of these lichens are still missing.

Phytosociology Freshwater lichen vegetation can be arranged in characteristic and more or less colourful belts at the shores and banks of lakes and watercourses. This phenomenon has attracted the interest of phytosociologists for a long time and many relevées from various European regions exist. More frequent however, is a patchy, small-scale distribution of species with different hydration needs and the FLUFXPVFULSWLRQRIKRPRJHQRXVUHOHYpHVLQWKH¿HOGLVHVSHFLDOO\GLI¿FXOWLQWKH splash water zone. Comprehensive overviews of the currently described phytosociological unities from Central Europe are described and discussed by Wirth

8

Introduction

(1972) and Thüs (2002). For communities occurring on siliceous substrata detailed systems of phytosociological entities already exist, but for those from carbonaceous rocks (especially for the amphibious and rarely inundated communities) more relevées and systematic arrangements are still needed before a reasonable system can be proposed. Basic data for freshwater limestone communities have been collected by Pereira (1992) with a strong focus on the Mediterranean region and thus cannot be directly adopted for Central European rivers.

Collection and Storage of Herbarium Specimens Aptroot & Seaward (2003) provide a good introduction of the methodology for collecting freshwater lichens, and their description is followed here. Usually, it is LQHYLWDEOHWRFROOHFWVSHFLPHQVIURPWKH¿HOGIRUWKHGHWHUPLQDWLRQRIIUHVKZDWHU OLFKHQV(YHQIRULGHQWL¿FDWLRQVWRJHQXVOHYHOPLFURVFRSLFDO FKDUDFWHUVKDYHWR be consulted by means of a compound microscope. In general, large thalli with reproductive structures (apothecia, perithecia, pycnidia, soralia or isida) should be selected for collection, because material without fruiting bodies or other dispersal structures is often impossible to determine. In the case of foliose species, it is possible to remove the thalli from the substratum by using a knife, but care has to be taken that the lower side is not damaged and possibly present rhizines are preserved during the collection process. Most freshwater lichens however, are crustose and tightly connected to their substratum. Some species are fast growing and colonize even small and moving pebbles. They are easily collected with their substratum, but the collection of loose pebbles will usually cover only a small fraction of the total species diversity. For a complete assessment of species, staEOHURFNVÀXVKHGRUVSODVKHGE\ZDWHUHJLQUDSLGV KDYHWREHVWXGLHGDVZHOO as small pebbles, old wood or roots of Elder (Alnus ( nigra) and Widdow (Salix spp.). For the collection of the species from rocky substrates, the use of hammer and chisel is inevitable. Working with the chisel below water level in a current is VRPHWLPHVGLI¿FXOWDQGSHULRGVZLWKORZZDWHUDSSHDUDVWKHSUHIHUUHGWLPHIRU inventories of freshwater lichens. A few species however (especially Verrucaria aquatilis, V. funckii and Hydropunctaria rheitrophila) live at sites that are permanently submerged and it may be necessary to examine rocks from at least half a PHWHUEHORZWKHFXUUHQWZDWHUOHYHOLQRUGHUWR¿QGWKHVHGHVLFFDWLRQVHQVLWLYH species in larger streams even during low water periods. Large boulders in rapidly running streams usually have smooth surfaces, thus PDNLQJLWGLI¿FXOWWR SODFHWKHFKLVHOLQWKHULJKWSRVLWLRQ,QVXFKFDVHVLWLVXVHIXOWR¿UVWPDNHDIXUURZDIHZFHQWLPHWUHVEHVLGHWKHOLFKHQWKDOOXVXVLQJWKH FKLVHOLQYHUWLFDOGLUHFWLRQ,QDVHFRQGVWHSWKHFKLVHOLVSRVLWLRQHGDWWKHÀDQNRI WKHIXUURZPRUHRUOHVVSDUDOOHOWRWKHURFNVXUIDFHDQGDWKHQ DÀDWSLHFHZLWKWKH lichen can be chipped off the rock. After collection, lichen samples should be dried as soon as possible in order to avoid moulding. For photobiont isolation, subsequent cultivation and detailed LGHQWL¿FDWLRQWKHVSHFLPHQVPXVWQRWGU\RXWDVVRPHRIWKHSKRWRELRQWVDUHYHU\ sensitive to desiccation. If it is impossible to dry the lichen sample directly in the ¿HOG (e.g. during rainy weather), transportation of lichen samples should be done

Introduction

9

LQSODVWLFEDJV)RU¿QDOVWRUDJHLQWKHKHUEDULXPWKHOLFKHQWKDOOLPXVWEHGULHG completely. It should be noted however, that rapid drying may cause the formation of wide cracks in the thalli of submersed lichens which may be problematic for determination, especially when the thallus surface is usually continuous. The paper capsules which are commonly used by lichenologists for epiphytic samples become rapidly soaken and instable when wet specimens of rock are transported. Small freezer bags with a volume of 1 litre are usually best suited since the plastic is more robust. An additional smaller set of 3-litre freezer bags is useful for transportation of large samples which is sometimes necessary when the rock is very hard and smaller samples are impossible to obtain. Each bag should contain only a single piece of rock and the bags have to be carried carefully in order to avoid damage of the thallus surfaces. Sample numbers and some basic collection data should be noted with a water resistant permanent pen directly on the freezer bag FRPSOHWHGE\GHWDLOHGGHVFULSWLRQVRIWKHFROOHFWLRQVLWHLQWKH¿HOGERRNORFDOity, distance from current water level, turbulence of the streaming, accompanying organisms, vegetation type in the surrounding etc.).

Preparation and Storage of Sections Thallus dimensions, inner and outer thallus structure as well as fruit body morSKRORJ\RIWKHIXQJDOV\PELRQWDUHLPSRUWDQWFKDUDFWHULVWLFVIRUVSHFLHVLGHQWL¿cation. For the study of internal morphology sections of thallus and fruit bodies are necessary. Best results are obtained using a freezing microtome, but in most cases sections by hand with a razor blade will have to suit for determination purposes. Recommendable is the use of smaller blades which are used to remove KRUQ\VNLQ7KHVHEODGHVDUHFKHDSHUXVXDOO\VKDUSHUDQGOHVVÀH[LEOHFRPSDUHG to many common razor blades. Care should be taken to use only new, sharp blades and it is advisable to have a set of new blades at hand. Depending on the consistency of the lichen thallus, sections should be done with slightly moistened material. Usually it is recommendable to put a drop of distilled water on the lichen thallus, wait until it is absorbed and afterwards test several times for optimal stability of the lichen tissue during preparation. If the thallus is too dry, sections will be brittle and the surface of the sections becomes uneven. If the thallus is too wet, the thallus will be squeezed instead of being cut by the blade. Finer structures of the fungal tissues and ascospores are sensitive to shrinking or swelling in different media. Therefore, all measurements should be taken in distilled water. If chemical tests are required, the application of e.g. caustic potash solution, Lugol’s solution or Nitric acid should always be done after the measurePHQWVKDYHEHHQ¿QLVKHG8QVWDLQHGVHFWLRQVFDQHDVLO\EHVWRUHGGU\RQDVOLGH FRYHUHGE\DFRYHUVOLS7KHFRYHUVOLSFDQEH¿[HGZLWKVPDOOGURSVRIQDLOYDUnish which are placed at the corners of the coverslip, while the section is embedGHGLQDGURSRIZDWHU7KHZDWHUSUHYHQWVWKHYDUQLVKWRÀRZXQGHUWKHFRYHUVOLS and additionally serves as a distance keeper. After study, the water will evaporate IURP WKH RSHQ ÀDQNV EHWZHHQ VOLGH DQG FRYHU VOLS DQG WKH GU\ VHFWLRQV FDQ EH stored without further treatment in a dry and tightly closed slide box. For re-exDPLQDWLRQDGURSRIGLVWLOOHGZDWHULVSODFHGDWWKHÀDQNRIWKHFRYHUVOLSDQGWKH water is soaked under the coverslip via capillary forces.

10

Introduction

Morphological Characteristics of Freshwater Lichens 0RUSKRORJLFDO VWUXFWXUHV UHOHYDQW IRU LGHQWL¿FDWLRQ RI IUHVKZDWHU OLFKHQV DUH shortly described and illustrated in the glossary. For more comprehensive descriptions of thallus anatomy and function textbooks like Nash (2008) and the introductions of the books written by Wirth (1995, in German) or Purvis et al. (1992) and Brodo et al. (2001) are recommended. 5XQQLQJZDWHULVDVWURQJVHOHFWLYHIRUFHZKLFKLQÀXHQFHVWKHHFRSK\VLRORJ\DV well as the mechanical constraints of the evolution of amphibious lichens. Consequently, lichens adapted to amphibious habitats are often very similar in morphology. Structures are optimised for minimum resistance to the current, e.g. most freshwater lichens are crustose, a much smaller number is foliose and only a few taxa with fruticose thalli occur in or at watercourses (Santesson 1939, Nordin 2002a). The reproductive organs are either perithecia which are often protected by a solid, carbonised cap, the involucrellum, or apothecia which are often immersed in the thallus or situated in thallus warts. Prominent and unprotected reproductive structures like soredia or sessile apothecia are rare exceptions in freshwater lichens. Lichen thalli are often differentiated in functional layers: a cortex (with cells of distinctly different shape compared to those in the algal layer) or pseudocortex (lacking photobionts, often with a special pigmentation but with fungal cells of similar size and shape as in the algal layer), a photobiont layer containing the algae, and a medullary layer which is usually composed of loosely interwoven hyphae. In terrestrial lichens, the medullary hyphae are often coated by water repellent substances, thus facilitating the mobility of gaseous components within the thallus (Honegger 1998). In the water saturated environPHQWRIIUHVKZDWHUOLFKHQVWKHDLU¿OOHGPHGXOODU\OD\HULVJHQHUDOO\UHGXFHGRU composed of a carbonised and tightly glued paraplectenchyma referred to as “black basal layer”.

Photobionts The algal component of the lichen symbiosis was long regarded as a fundamental basis for both, large-scale systematics and for species separation, especially in freshwater lichens (Zahlbruckner 1921, Zschacke 1934, Servít 1954). However, WKHLGHQWL¿FDWLRQRIWKHSKRWRELRQWVZDVRIWHQEDVHGRQWKHVWXG\RIWKHOLFKHQL]HG phenotypes alone. In some cases it is possible to determine lichen photobionts to JHQXV OHYHO +RZHYHU LQ PRVW FDVHV WKH LGHQWL¿FDWLRQ RI WKH DOJDH IURP OLFKHQ WKDOOXVVHFWLRQVZLOOSURYHWREHYHU\GLI¿FXOWRUYLUWXDOO\LPSRVVLEOH±HYHQWR family level. The use of photobiont characters for species determination in lichens LVWKXVPXFKUHVWULFWHGE\SUDFWLFDOOLPLWDWLRQV5HFHQWVWXGLHVRQWKHVSHFL¿FLW\RI the fungal-algal relationship by means of molecular markers revealed a limited ÀH[LELOLW\LQVSHFL¿FJURXSVRIOLFKHQIXQJLZKLFKLVIXUWKHUGHSHQGLQJRQDYDULHW\ RIKDELWDWVSHFL¿FIDFWRUV%HFN%HFNHWDO 3LHUFH\1RUPRUH  DePriest 2001, Guzow-Krzemínska 2006). Still, there are some general patterns, e.g. the obligatory association of all members of the Collemataceae, Lichinaceae

Introduction

11

and Placynthiaceae with prokaryotic, cyanobacterial photobionts or the limited submersion tolerance of lichens with Trebouxia s.str., Asterochloris or Trentepohlia photobionts in contrast to the generally hydrophilic and desiccation sensitive lichens with 'LODEL¿OXP or Heterococcus as photosynthetic partners. For many VSHFLHVKRZHYHUPRUHGDWDDUHQHHGHGWRDFFXUDWHO\GHWHUPLQHWKHVSHFL¿FLW\RI the fungal-algal relationship and to achieve a better understanding of the reasons IRUVLWHVSHFL¿FGLIIHUHQFHV7KHHFRSK\VLRORJ\RIVXLWDEOHOLFKHQSKRWRELRQWVFHUtainly plays an important role in the understanding of the ecological preferences of OLFKHQVEXWVWLOOPXFK¿HOGREVHUYDWLRQVDQGODERUDWRU\H[SHULPHQWVDUHQHHGHGLQ order to verify preliminary conclusions as outlined above, e.g. for the preferences of amphibious lichens for acidic water. We have provided information about the identity of lichen-associated photobionts only when these informations have been FRQ¿UPHG E\ RXU RZQ REVHUYDWLRQV LQ WKH VWXG\ DUHD RU KDYH EHHQ UHSURGXFHG from reliable literature sources clearly referring to Central Europe and including detailed drawings and descriptions of the isolation process. In the keys we have attempted to make only little use of characters of the photobionts. In order to facilitate the distinction of relevant colour characters, examples of several photobionts in the thalli of amphibious lichens are given in colour plate 6. $SURSHULGHQWL¿FDWLRQRIDSKRWRELRQWWRVSHFLHVOHYHOXVXDOO\ UHTXLUHVWKHLVRODtion of photobiont cells from the lichen thallus and the cultivation of the algae or cyanobacteria in culture media. For isolation of amphibious lichen photobionts only freshly collected material should be used which has to be kept in moist condiWLRQXQWLOWKHLVRODWLRQSURFHGXUH)RULVRODWLRQDODPLQDUÀRZEHQFKDQGVWHULOH FRQGLWLRQVDUHUHTXLUHG,QWKHODPLQDUÀRZEHQFKDVWHUHRPLFURVFRSHLVVWHULOLVHG with ethanol. The surface of many amphibious lichens can be covered by epiphytic algae which are easily mistaken for photobiont cells when cultivation experiments are performed with entire thallus fragments. Therefore, it is recomPHQGDEOHWR¿UVWGHFRUWLFDWHDSSUR[LPDWHO\PP2 of the lichen by a horizontal section with a razor blade. In the decorticated area a small drop of sterile water is placed and the algae containing layer of the thallus is disrupted by means of a sterile preparation needle. The resulting dispersion of algal and fungal cells is VRDNHGZLWKWKHWLSRID¿QHJODVVFDSLOODUHJWKHWLSRID 3DVWHXUSLSHWWHDIWHU careful elongation over a gas burner). The suspension is then transferred onto a Petri-dish containing a suitable culture medium. For green algae and Xanthophytes good results are received with Bold’s Basal Medium with an additional drop of soil extract per litre. Axenic (bacteria free) cultures can be prepared either by repeated streaking of small cell groups to fresh media or by picking single cells from thallus fragments or pre-cultured thallus suspensions XVLQJDPRGL¿HGPLFURPDQLSXODWRU%HFN .RRS 6WLOODJRRGVWDUWLQJSRLQW IRULGHQWL¿FDWLRQ is the comprehensive “Syllabus der Boden-, Luft- und Flechtenalgen“ (Ettl & *lUWQHU which also includes instructions for isolation and cultivation of lichen photobionts.

12

Introduction

Taxonomic Concept The relatively uniform structure of the vegetative thallus makes quantitative characters of the reproductive structures a principal source of information for morphological species delimitation in many freshwater lichens. When using TXDQWLWDWLYHFKDUDFWHUVLQWKH NH\VFDUHPXVWEH WDNHQWRREWDLQDVXI¿FLHQWVDPple size. For example, the variation in ascospore size should be assessed by measuring at least 10 ascospores, if necessary from different asci and ascomata. This book is intended for use in ecological studies and we attempted to follow a VWULFWO\PRUSKRORJLFDOFRQFHSWIRUVSHFLHVLGHQWL¿FDWLRQ5HFHQWUHVXOWVRIPRlecular studies are commented where appropriate, but changes in species delimitation were only proposed if a morphology based distinction of the taxa is possible. In general, only those taxa were accepted which are corrobated by at least two independent characters from otherwise similar taxa. Ecological characters are KHOSIXOIRULGHQWL¿FDWLRQSXUSRVHVLQVRPHFDVHVEXWIRUSURSHUVSHFLHVGHOLPLWDtion morphological, chemical – or if available – molecular characters are mandatory. In some cases such as the crustose genera of the Verrucariales we followed traditional genus concepts based on single characters which have been demonstrated to have evolved convergently in different lineages of the Verrucariales (Gueidan et al. 2007, 2008). Although the polyphyletic origin of genera such as Verrucaria, Polyblastia and Thelidium has become evident, new monophyletic HQWLWLHVDUHRIWHQGLI¿FXOWWRGLVWLQJXLVKE\PRUSKRORJLFDOFKDUDFWHUVDORQHDQG for several lineages a morphological circumscription has not been achieved so far. We have followed Gueidan et al. (2008) in the separation of the black dotted species of Verrucaria with small-sized ascospores as the new genus Hydropunctaria and the treatment of Sporodictyon at genus rank separated from the remaining species of Polyblastia 6DYLüHWDO . However, further changes in the delineation of the genera Verrucaria, Polyblastia and Thelidium are to be expected in the near future. Similarly, the cyanobacterial lichen genera Porocyphus, Pterygiopsis, Pyrenocarpon, Pyrenopsis and Thelignya are currently based on (sometimes still rather vague) morphological concepts (Henssen ‘1979’1980, Henssen & Jørgensen 1990, Schultz & Büdel 2002) and thus, changes are to be H[SHFWHGRQFHVXI¿FLHQWPROHFXODUGDWDKDYHEHFRPHDYDLODEOH

Photography Only few of the taxa treated in this book have been illustrated elsewhere. Since SKRWRJUDSKVRIWKHRYHUDOOWKDOOXVKDELWXVFDQFRQWULEXWHPXFKWRZDUGVLGHQWL¿cation, we provide images for almost all of the species treated here. However, we are aware that images can illustrate only a small proportion of the species variation. Furthermore, species of genera such as Verrucaria and Thelidium may look very much alike and can be distinguished only using microscopic characters. Therefore, the photographic illustrations are intended to serve as a supplement to the information provided in the species descriptions and keys.

Introduction

13

Most of the lichens are illustrated by black and white photographs directly accompanying the descriptions in the text. Some colour images have been assembled at the end of the book to give an impression of the general appearance and colour for a selection of the species of Ionaspis, Staurothele, Verrucaria as well as some others. With a few exceptions, we did not attempt to take photographs at the natural habitat of the species. Instead, we selected recently collected material for illustration whenever available. Many aquatic species exhibit a vivid green colour when studied in nature. Unfortunately, this fresh green colour fades rapidly once the lichen is dried, mounted and kept under herbarium conditions. On the other hand, the dark colour of many cyanobacterial lichens changes little with time so that old herbarium material is still suitable for photography. A number of species occurs in terrestrial as well as in amphibious or riparian habitats. Sometimes, the external appearance of these lichens varies depending on the habitat. We therefore attempted to provide images based on material from habitats that the book is focused on (i.e. margins of rivers, lakes etc.). Most images were produced using the following digital equipment: a Canon EOS 'GLJLWDOFDPHUDPRXQWHGRQD1RYRÀH[*HUPDQ\ EHOORZVZLWK&DQRQ)' mount using an adapter (Kood, Japan Canon AF/FD, central correction lens rePRYHG DQG¿[HGLQD1RYRÀH[*HUPDQ\ PDFURVWDQG$VHWRIPDFUROHQVHV ZDVXVHGIRUDPDJQL¿FDWLRQUDQJHRI±WLPHV0RVWLPDJHVZHUHWDNHQXVLQJ a Zeiss (Germany) Luminar 40 mm/1:4 lens. Additionally, Zeiss (Germany) Luminar 63 mm/1:4.5 and 25 mm/1:3.5 lenses and two Canon Macro lenses 35 mm/1:2.8 and 20 mm/1:3.5 were used. Images 15 & 160 (Collema dichotomum, Dermatocarpon arnoldianum) were taken with a Canon Macro Lens EF-S 60 mm/1:2.8 USM. For illumination a Schott (Germany) cold-light source was used. An image of white photo paper taken with the camera set to automatic white balance was used as reference for correction of light temperature to obtain natural light conditions.

14

Glossary

Glossary A accessory lobule, small ± erect, simple or branched thallus lobe formed by outgrowing marginal isidia or originating from the margin or surface of larger, main lobes; term mostly used in Collema (DEGELIUS 1954, 1974). acicular, needle-shaped. amphibious, temporarily inundated or ± frequently wetted by splash water. amyloid, staining blue, blue-violet, orange or red in iodine (I). anastomosing, repeatedly branched and reconnected. apical dome, thickened inner part of the ascus wall in the ascus apex, in some species amyloid (staining I+ blue with iodine). apothecial disc, the surface of the hymenium surrounded by the exciple. apothecium, disciform to cup shaped ascoma (fruit body) with hymenium exposed at maturity. aquatic, (in the sense used here) completely submersed, or very rarely subject to desiccation. areolate, crustose thallus with deep cracks separating the thallus into mosaic-like fragments. areole, one of the fragments in an areolate thallus. ascoma, fruit body in which the asci are formed; in freshwater lichens either perithecia or apothecia. ascospore, diaspore type, meiotically formed in an ascus. ascus, cylindrical to sack-like, swollen hyphal cell in which ascospore formation takes place.

B bacilliform, rod-shaped. (black) basal layer, algal free tissue of compact fungal cells with brownishblack cell walls in the basal part of a crustose thallus.

Glossary

15

biatorine, type of apothecial margin formed by a pale, not carbonised true exciple free of algae. blastidium, rounded lichen propagule produced by the budding (yeast-like propagation) of a thallus.

C C, saturated aquaceous solution of calcium hypochlorite, Ca(OCl)2. Commercially available for disinfection purposes and principal compound of common house-hold bleach products. Instable, proper reactivity should be checked regularly on lichens with known reaction (e.g. the common epiphyte Lecanora expallens, which should turn C+ yellow to reddish if the reagent is still in good condition). Caustic! canaliculate, channelled. carbonised, optically dense, brown-black appearance of fungal tissue with usually small cell lumina and thickened cell walls, containing large amounts of dark coloured phenolic substances. cilia, hair-like outgrowths on the thallus surface or margin. clavate, club-shaped. conidium (conidiospore), mitotically produced spores, in freshwater lichens exclusively found in pycnidia. cortex, outermost layer in heteromerous thalli composed of differentiated fungal hyphae, often with compact arrangement and special cortical pigments. crenate, with toothed edges, teeth rounded. crenulate, delicately crenate. crustose, lichen growth form with thalli tightly attached to the substratum and lacking a lower cortex.

D decumbent, resting on the substratum with the ends turned up.

16

Glossary

E HI¿JXUDWH, thallus margin consisting of ± radially arranged, elongated areoles. epihymeniumRIWHQFRORXUHGOD\HUDERYHWKHK\PHQLXP¿J  epinecral layer, layer of dead fungal hyphae with indistinct, often collapsed lumina. epithecium, upper part of hymenium. eumuriform, spores having numerous horizontal and longitudinal septa. euparaplectenchymatous, composed of ± isodiametric, strongly conglutinate hyphae; term refers to a type of proper exciple mostly in Collema (DEGELIUS 1954, 1974). euthyplectenchymatous, composed of parallel, elongated, not or weakly conglutinate hyphae; term refers to a type of proper exciple mostly in Collema (DEGELIUS 1954, 1974). exciple, tissue forming the wall of an ascoma (apothecium or perithecium). excipulum proprium (excip. prop., proper or true exciple), margin of an apothecium composed of a fungal plectenchyme, pale or variously coloured, carbonised or not, texture various from loosely interwoven to strongly agglutinated, from reticulate to euthyplectenchymatous or paraplectenchymatous.

F foliose, leaf-like growth form of lichens with the thallus attached to the substratum by rhizines, hapters or punctually by the lower surface, but never with the entire lower surface. Often with differentiated upper or lower cortex. fruticose, shrub-like growth form of lichens with ± cylindrical, often furcate and erect thallus branches. fusiform, spindle-like, with acute apices.

G globose, with rounded outline.

H halonate, ascospores surrounded by a thick transparent coat or perispore (often distinctly swelling in K).

Glossary

17

hamathecium, sterile hyphae between asci and ascoma walls. hapterVWUXFWXUHZKLFKLV¿[LQJWKHWKDOOXVWRWKHVXEVWUDWXP heterocyte, (often incorrectly referred to as “heterocyst”) special cells in some ¿ODPHQWRXV F\DQREDFWHULD IUHHOLYLQJ RU OLFKHQL]HG FKDUDFWHUL]HG E\ D WKLFN PXOWLSO\ OD\HUHG ZDOO DQG HQ]\PHV DEOH WR ¿[ DWPRVSKHULF QLWURJHQ SRVLWLRQ basal (e.g. Rivulariaceae) or intercalary (Nostoc, Stigonema, Scytonema). heteromerous, photobiont cells arranged in a distinct layer in the lichen thallus. homoiomerous, photobiont cells not forming a distinct layer in the lichen thallus. holdfast, structure for punctual attachment of the lichen thallus to the substratum. hydrophytic, demanding frequent or constant supply of liquid water. hygrophytic, demanding high air humidity or supply of liquid water. hymenial algae, photobiont algae in the cavity of a perithecium. hymenium,IXQJDOWLVVXHFRPSRVHGRILQWHUDVFDO¿ODPHQWHJSDUDSK\VHV DQG asci. hypothecium, layer of fungal tissue below the hymenium.

I I, chemical reagent, 0.3% aquaceous solution of 1 part iodine, I and 2 parts poWDVVLXPLRGLQH.,7KH¿QDOVROXWLRQLVLQVWDEOHDQGWKHSUHSDUDWLRQRIDVWRFN solution (1 g I and 2 g KI diluted in 30 ml distilled water) is recommended for storage. The working solution is a freshly prepared 10% dilution of the stock solution. involucrellumFRQLFDOWLVVXHRIFRPSDFWFHOOVVXUURXQGLQJWKHH[FLSOH¿J   Sometimes fused with the exciple and only recognizable by a abrupt thickening of the perithecial wall. isidium, lichen propagule covered by a distinct cortex and containing fungal and algal cells. isodiametric, more or less equally sized in all directions.

18

Glossary

K K, chemical reagent, 10% aquaceous solution of potassium hydroxide, KOH. Caustic!

L laminal, situated on the surface of a thallus. lecanorine, apothecium with thalline exciple, i.e. exciple composed of thallus WLVVXHFRQWDLQLQJDOJDH¿J  lecideine, apothecium with blackish, carbonised exciple not containing algae ¿JV–3). lichenicolous, growing on lichens. lignicolous, growing on dead wood. lobateOHDIOLNHXVXDOO\HDVLO\OLIWHGRIIWKHVXEVWUDWXP¿[HGE\ UKL]LQHVKDSWHUV or with one or several attachment points, but not with the entire lower surface.

M medulla, fungal tissue lacking algae in the central or lower part of a lichen thallus. Either composed of loosely intervowen hyphae (typical for terrestrial lichens) or of compact and sometimes deep brown-coloured tissue (“black basal layer”, typical for some amphibious lichens). Meltzer’s Reagent, chemical reagent, composed of iodine, I solved in 50 % aquaceous chloral hydrate. Instable, iodine and chloral hydrate solutions should be stored separately and mixed before application. muriform, ascospores having transverse as well as horizontal septa.

N N, chemical reagent, concentrated nitric acid, HNO3, very aggressive, also against microscope lenses etc. Caustic!

O ostiole, the apical opening of a perithecium or pycnidium.

Glossary

19

P P, chemical reagent, alcoholic solution of para-phenylenediamine. The solution is instable and it is recommended to be prepared freshly before application by solving some small crystals in a drop of 96 % ethanol. Para-phenylenediamine is also a principal component of most commercially available developing solutions for colour photographs. Cancerogenic and strongly dying clothes! papilla, small wart-like protuberance. paraplectenchymatous, compact fungal hyphae arrangements with ± isodiametric cells. In a loose sense also used for any compact fungal tissue without intercellular spaces. Characteristic feature of most obligately aquatic lichens. prosoplectenchymatous, fungal hyphae type with elongated lumina. Often with intercellular spaces and typical for terrestrial lichens. paraphysisVOHQGHUIXQJDO¿ODPHQWVJURZLQJXSZDUGVEHWZHHQWKHDVFLVRPHWLPHVEUDQFKHGDQGDQDVWRPRVLQJDQGZLWKVZROOHQWLSV¿J  peltate, shield-like with central attachment. periclinal, arranged ± parallel to the surface or the circumference. periphysis, hair-like fungal hyphae growing on the inner surface of the upper SDUWRIDSHULWKHFLXPWRZDUGVWKHRVWLROH¿J  perispore, the outer layer of an ascospore. peritheciumW\SHRIDVFRPDZLWKVXEJORERVHWRÀDVNOLNHVKDSH¿J LQ ZKLFK the hymenium is enclosed and the ascospores are released via a small apical opening, the ostiole. photobiont, photosynthetically active partner of the lichen symbiosis. Either a cyanobacterium or an eucaryotic alga. Form and inner architecture of photobiont cells in the lichen symbiosis are often much different from those of free living or cultivated cells. placodioid, lichen thallus with radiating marginal lobes. polarilocular, type of ascospores with two lumina separated by a ± thick septum which is perforated by a thin central plasmatic channel. polysporous, with more than eight spores per ascus. prothallusWKLQDQGRIWHQ“¿PEULDWHDOJDOIUHH]RQHRIIXQJDOK\SKDHERUGHULQJ the algae containing part of the thallus. Usually of different colour in comparison to the algal containing lichen thallus.

20

Glossary

pustulate, blistered. pycnidium JORERVH WR ÀDVNOLNH VWUXFWXUH LQ ZKLFK FRQLGLD S\FRQVSRUHV  DUH produced. pycnospore, mitotically produced spore which is formed in a pycnidium (see also conidium). pycnoascocarp, type of fruit body in the Lichinaceae with ascogones and trichogynes formed beneath pycnidia and conidiophores often becoming elongated and serving as primary paraphyses; these primary paraphyses often becoming replaced by common (secondary) paraphyses once the pycnidium has been transformed into an apothecium. pyrenocarpous, with perithecia. pyriform, pear-shaped.

R rhizine, root-like compact bundle of hyphae serving to attach the lichen thallus to the substratum. rhizohypha (pl. rhizohyphae)IXQJDO¿ODPHQWDULVLQJIURPWKHORZHUVXUIDFHRI a thallus serving to attach the thallus to the substratum. rimose, irregular cracked thallus surface. riparian RFFXUULQJ UHJXODUO\ LQ WKH YLFLQLW\ RI ZDWHU ERGLHV EHQH¿Wting from increased humidity or fog formation, but not adapted to submersion and usually avoiding temporarily inundated sites and frequent wetting by splash water. rugulose, rough from small wrinkles.

S soralium, a structure or area on a lichen thallus pruducing soredia. soredium, lichen propagule containing fungal hyphae and algal cells but lacking a cortex (compare isidium). squamulose, growth form of lichens with the thallus composed of scale-like units, sometimes scales arranged like roof tiles and forming ± dense mats. subfruticose, lichen growth form intermediate between crustose and fruticose. subgelatinous, thallus consistency like solid jelly when wetted, usually thallus or WKDOOXVPDUJLQVEHFRPLQJ“WUDQVSDUHQW¿Js. 157–158).

Glossary

21

subglobose, almost rounded, slightly longer than wide. subsquamulose, lichen growth form intermediate between crustose and squamulose. submerged, (in sense used here) being inundated for most of the time. submuriform, ascospores with several transversal but only one or very few longitudinal septa. substipitate, almost stalked.

T teretiform, circular in section but narrowing to one end. terrestrial, (in the sense used here) living in micro-sites without inundation or VSODVKZDWHULQÀXHQFHV thalline exciple, apothecial margin containing algae, usually concolorous with WKHVXUURXQGLQJWKDOOXV¿J  thallinocarp, type of fruit body in the Lichinaceae with continuous or discontinuous, rather irregular hymenium covered by sterile thallus plectenchyma. thallus, the vegetative lichen body. thallus mantle, perithecia with a ± distinct cover of thallus tissue. t.l.c. WKLQOD\HUFKURPDWRJUDSK\ WHFKQLTXHWRVFUHHQIRUVSHFL¿FRUJDQLFFRPpounds in lichen thalli. Recommended is the use of HPTLC (High Performance Thin Layer Chromatographie, see Arup et. al. 1993 for details and further references). trichomesFHOOURZVRI¿ODPHQWRXVF\DQREDFWHULDZLWKRXWJHODWLQRXVVKHDWK true exciple, see excipulum propium.

U umbilicate, growth form of lichens with a foliose thallus attached by a central, basal holdfast. The thallus often has a bowl-like appearance.

22

Glossary

Fig. 1

Fig. 2

Fig. 3

Fig. 4

Plate 1. Ascocarps. Fig. 1. Lecanorine apothecium with thalline (algae containing) exciple, hypothecium (grey), asci, paraphyses and epihymenium. Fig. 2. Lecidein apothecium, exciple of Lecidea-type. Fig. 3. Lecidein apothecium, exciple of Porpidia-type. Fig. 4. Structure of perithecium with asci, paraphyses, periphyses, exciple and involucrellum.

Glossary

Fig. 7

Fig. 5

Fig. 6

Fig. 8

Fig. 9

Plate 2. Asci and Ascospores. Fig. 5. Lecanora-type ascus. Fig. 6. Porpidia-type ascus. Fig. 7. 7 Ascospore septation in Caloplaca: polarilocular ascospore. Fig. 8. Ascospore wall thickening in Rinodina: Mischoblastia-type. Fig. 9. Ascospore wall thickening in Rinodina: Pachysporaria-type.

23

General Key (aquatic or amphibious lichens in bold, terrestrial species which are not treated in further detail in italics and with author citation). 1a Photobiont is a cyanobacterium, chloroplast absent, with yellow, brown, EOXHJUHHQRUUHGGLVKFRORXUV¿JV. 170–171) ......................................... 62 1b Photobiont is an eucaryotic alga, chloroplast present, with green, yellow or RUDQJHFRORXUV¿JV. 166–169) .................................................................. 2 2a Thallus crustose or squamulose ................................................................ 3 2b Thallus foliose ........................................................................................ 55 3a Thallus fertile, perithecia or apothecia present ......................................... 4 3b Thallus sterile, perithecia or apothecia lacking (but pycnidia may be present) ................................................................................................................. 47 4a Thallus with perithecia .............................................................................. 5 4b Thallus with apothecia ............................................................................ 14 5a Hamathecial hyphae (i.e. paraphyses) soon gelatinizing, usually absent in ripe perithecia, never richly branched, phycobiont green, rarely yellowish but never orange, ascospores simple or septate ............................................... 6 5b Hamathecial hyphae persistent, branched or anastomosing, phycobiont green or yellowish to orange (Trentepohlia), mature ascospores always septate .......................................................................................................... 12 6a Perithecia without hymenial algae ............................................................ 7 6b Perithecia with hymenial algae ............................................................... 11 7a Ascospores simple (Verrucaria) ................................................................ 8 7b Ascospores septate .................................................................................... 9 8a Thallus with black dots RU FROXPQV ¿JV. 30–33) and ascospores mostly < 18 m long ..................................................................... Hydropunctaria 8b Not with the combination of black dots or columns in the thallus and ascospores < 18 m long .................................................................. Verrucaria 9a Ascospores with transverse septa only ....................................... Thelidium

26

General Key

9b Ascospores with transverse and longitudal septa (Polyblastia) ................. 9 10a Perithecia naked, ascospores colourless .................................... Polyblastia 10b Perithecia covered by a thin thalline mantle, mature ascospores brownish ............................................................................... Sporodictyon cruentum 11a Thallus squamulose ................................................................. Endocarpon 11b Thallus crustose ....................................................................... Staurothele 12a Ascospores with more than one septum, perithecia black or orange .... Porina 12b Ascospores uniseptate, perithecia black .................................................. 13 13a Involucrellum present, only eucaryotic algae associated (Trentepohlia, sometimes also other green algae incorporated in the thallus) ... Anisomeridium 13b Involucrellum absent, with Trentepohlia and usually with additional procaryotic photobionts (small yellowish cells without plastids) ....... Pyrenocollema 14a Ascospores simple .................................................................................. 15 14b Ascospores septate .................................................................................. 37 15a Apothecia yellow to orange-brown, not greenish (predominantly terrestrial lichens, only a selection of species is keyed out which are found ± frequently in or at watercourses) .............................................................................. 16 15b Apothecia with different colours, never yellow to orange-brown ........... 18 16a Apothecial disc orange to brown-orange, soon becoming convex, thallus pale grey to grey-brown to grey-green ......................................................... ....................................... Protoblastenia rupestris (Scopoli) J. Steiner 1930 16b Apothecial disc yellow, thallus not grey ................................................. 17 17a Asci (12) 16–32-spored, thallus distinct, never with black prothallus, never on calcareous substrata ................................................................................ .............................. Candelariella vitellina (Hoffmann) Müller-Argau 1894 17b Asci 8-spored, thallus indistinct but black prothallus often present, on limestone or base-rich substrata .......................................................................... ................................ Candelariella aurella (Hoffmann) Zahlbruckner 1928 18a Apothecia with biatorine exciple or exciple indistinct to absent, never immersed in the thallus ............................................................................... 19 18b Apothecia with thalline exciple or apothecia immersed in the thallus with weakly developed exciple ....................................................................... 25

General Key

27

19a Apothecial disc brown to pink, asci without or weakly blue amyloid apical structures ................................................................................................. 20 19b Apothecial disc black to black-brown, asci with amyloid apical structures ................................................................................................................. 21 20a 7KDOOXVVOLJKWO\HI¿JXUDWHRUVXEVTXDPXORVHDUHROHVRIWHQRYHUODSSLQJ ..... .......................................................... Trapelia involuta (Taylor) Hertel 1973 20b 7KDOOXV FUXVWRVH QHYHU HI¿JXUDWH RU VXEVTXDPXORVH DUHROHV QRW overlapping ........................................ Trapelia coarctata (Smith) M. Choisy 1932 21a Exciple absent or soon becoming inconspicuous ......................................... .............................................. (Micarea spp., terrestrial species, not treated) 21b Exciple present and distinct .................................................................... 22 22a Exciple dark throughout (PorpidiaW\SH¿J ............................ Porpidia 22b Exciple with dark pigment restricted to peripheral parts ((LecideaW\SH¿J. 2, Lecidea/Lecidella, a selection of species is keyed out, but more species likely to be found; especially at the margins of upland creeks) ......................... 23 23a Paraphyses lax, easily separating in K when gently pressed, asci of Lecanora-type (amyloid cap surrounding a thin unstained channel at the tip of the DVFXV¿J5), hymenium and hypothecium with blue-violet to blackish, K+, green granules ..................................................................... Lecidea ahlesii 23b Paraphyses not lax, asci of Porpidia-type (amyloid tube at the tip of the DVFXV¿J6), hymenium and hypothecium without K+ green granules .... 24 24a Hypothecium colourless to pale yellow-brown, exciple edge blue-black, indigo to green-black, colourless within, thallus K-/K+, yellow, KC-, thallus contains zeorin, r atranorin, ± lichexanthone (detectable by thin layer chromatography) ......... Lecidella stigmatea (Acharius) Hertel & Leuckert 1969 24b Hypothecium yellow-brown, exciple edge bluegreen-black, reddish-brown within, thallus KC+, yellow, thallus contains diploicin, r thuringion (detectable by thin layer chromatography) ...... Lecidella carpathica Körber 1861 25a $SRWKHFLDVHVVLOHWKDOOXVJUHHQLVKRUZKLWHXVXDOO\LQWKHÀXYLDO[HULFRUterrestrial zone (Lecanora ( , predominantly terrestrial species, a selection of species which often occur on riverbanks and lake margins is keyed out) ......... 26 25b Apothecia r immersed, thallus grey, brown, orange or whitish, usually in the ÀXYLDOPHVLFRUVXEPHUJHG]RQH ............................................................. 30 26a Thallus distinct, old thalli up to several centimetres in diameter, greenish (occasionally whitish-green) ................................................................... 27

28

General Key

26b Thallus inconspicuous, of dispersed granules; if white, then thalli small and less than 0.5 cm in diameter .................................................................... 28 27a Thallus placodioid, with lobe-like margins (terrestrial, but characteristic component of the riparian vegetation) .......................................................... ............................................ Lecanora muralis (Schreber) Rabenhorst 1845 27b Thallus crustose, margins continuous to crenulate or only minutely lobed .. .............................................. Lecanora intricata (Acharius) Acharius 1810 28a Apothecial margin green or faintly yellowish green .................................... ....................................... Lecanora polytropa (Hoffmann) Rabenhorst 1854 28b Apothecial margin grey or white ............................................................. 29 29a Thallus inconspicuous or of scattered granules, apothecia dispersed ........... ................................................. Lecanora dispersa (Persoon) Röhling 1813 29b Thallus white, of aggregated areoles, apothecia concentrated in the centre ... ............................ Lecanora albescens (Hoffmann) Branth & Rostrup 1869 30a Apothecia (greenish-) whitish to pink, photobiont green (Trebouxia/Asterochloris sp.), never yellowish to orange, thallus smooth .... Ionaspis lacustris 30b Apothecia black or (reddish-) brown, photobiont green or (yellowish-) orange (Trentepohlia VS¿JV. 168–169), thallus smooth to warty or papillate .... 31 31a Apical dome I+ blue, thallus yellowish-brown ............................................ ......................................................... (LJOHUDÀDYLGD (Hepp) Hafellner 1984 31b Apical dome I-, thallus yellowish, brown, grey or pinkish ..................... 32 32a Thallus margins HI¿JXUDWH .................................... Lobothallia melanaspis 32b 7KDOOXVPDUJLQVQRWHI¿JXUDWH ................................................................ 33 33a Epihymenium colourless or blue-green and N+ red to mauve, photobiont green to orange (Trentepohlia), thallus yellowish, olive, brown or pinkishgrey ......................................................................................................... 36 34a on calcareous rock ........................................................................ Ionaspis 34b on siliceous rock ..................................................................................... 35 35a Apothecial margin distinctly projecting over the thallus, disc blue-black with N+ red upper hymenium, on siliceous substrata .................................. ................................................................................ Hymenelia cyanocarpa 35b Apothecial margin ± completely immersed in the thallus, apothecial disc either brownish to black and N-, or blue-black and N- ................. Ionaspis

General Key

29

36a Thallus papillate-isidiate, apothecia rare, immersed in the tips of isidiate papillae .............................................................. Koerberiella wimmeriana 36b Thallus not papillate-isidiate, apothecia never immersed in papillae ........... ....................................................................................................... Aspicilia 37a Apothecial disc yellow, orange, orange-brown, K+ red; ascospores colourless, polarilocular ........................................................................ Caloplaca 37b Apothecial disc differently coloured, K-; ascospores colourless or brown, not polarilocular, but in some cases with unequal wall thickenings ........ 38 38a Ascospores colourless ............................................................................. 39 38b Ascospores black or brown (when ripe) .................................................. 46 39a Ascospores with gelatinous perispore ..................................................... 40 39b Ascospores without gelatinous perispore ................................................ 41 40a Apothecia ± immersed in the thallus, epithecium K+ red ............................ ............................................................................. Rhizocarpon amphibium 40b Apothecia sessile, epithecium K- .................................................. Gyalidea 41a Apothecia with thalline exciple .................................................................... ............................ Lecania inundata (Hepp ex Körber) M. Mayrhofer 1987 41b Apothecia with true exciple .................................................................... 42 42a Ascospores with one septum, not fusiform to acicular ............................ 43 42b Ascospores with 2 or more septa, of various shape ................................. 45 43a Apothecial disc concave, yellowish to pink, hymenium I- or I+ yellow ...... .......................... Absconditella delutula (Nylander) Coppins & Kilias 1980 43b Apothecia ÀDWWRFRQYH[EURZQWREODFNK\PHQLXP,EOXH ................. 45 44a Exciple dark coloured throughout, hypothecium dark brown ...................... ...................................... Catillaria chalybeia (Borrer) A. Massalongo 1852 44b Exciple edge dark coloured, brown to colourless inside, hypothecium colourless or pale straw ..... Catillaria lenticularis (Acharius) Th. Fries 1874 45a Thallus with orange-yellow photobiont (Trentepohlia), ascospores muriform ................................... Gyalecta jenensis (Batsch) Zahlbruckner 1924 45b Thallus with green photobiont (never with Trentepohlia), ascospores with transversal septa only .................................................................... Bacidina

30

General Key

46a Ascospore walls unequally thickened, with one septum, without gelatinous perispore ...................................................................................... Rinodina 46b Ascospores walls equally thickened, with one or more septa, with gelatinous perispore ................................................................................ Rhizocarpon 47a 7KDOOXVZLWKPLQXWHÀDWOREHVDWOHDVWDWWKHPDUJLQVZLWKUKL]LQHVRQWKH lower surface ......................................................................... Phaeophyscia 47b Thallus never with lobes, never with rhizines ......................................... 48 48a Thallus with isidia ................................................................................... 49 48b Thallus lacking isidia .............................................................................. 50 49a Isidia large and easily recognized by the naked eye, one or only a few per areole, thallus greyish, K-, in and at clean upland watercourses .................. ............................................................................ Koerberiella wimmeriana 49b Isidia minute and easily mistaken for soralia, densely covering the thallus, often with a bluish tinge, thallus and iVLGLD.SDOHYLROHWPD\EHGLI¿FXOW to observe), at eutrophic margins of lowland and low mountain creeks and rivers ........................................................................... Caloplaca isidiigera 50a Thallus with soralia ................................................................................. 51 50b Thallus lacking soralia ............................................................................ 52 51a Thallus brownish, orange, grey to white, C-, KC-, without gyrophoric acid ....................................................................................................... Porpidia 51b Thallus grey or whitish, often with pinkish tinge, C+/KC+ red, with gyrophoric acid .......................... Trapelia placodioides Coppins & James 1984 52a Thallus with pycnidia .............................................................................. 53 52b Thallus lacking pycnidia, soralia, blastidia, isidia or marginal lobuli .......... .................................................................................................... not treated 53a Phycobiont greenish yellow to orange (Trentepohlia), thallus thin, without prothallus, conical pycnidia with bacilliform pycnospores, 3.5–4.5 × ±ȝP, extruded in cirrus like gelatinous matrix or immersed globose with pycnoVSRUHV±î±ȝP ................................................................................. ........................ Anisomeridium polypori (Ellis & Everhart) M.E. Barr 1996 53b Phycobiont green (chlorococcoid), white prothallus present or absent, pycnidia never with conical shape, pycnospores of various form and size ±ȝPORQJ WKDOOXVWKLQWRWKLFN ....................................................... 54 54a With white or crème-coloured prothallus ...................................... Bacidina

General Key

31

54b Without prothallus ....................................................................................... not treated (mostly Micarea spp. or morphs of some species of Bacidina/ Bacidia) 55a Thallus ascending, lobes usually much wider than 2 mm, umbilicate (one FHQWUDOKROGIDVW RU¿[HGZLWKPDQ\DWWDFKment points, never with rhizines ................................................................................................................. 56 55b 7KDOOXVIRUPLQJÀDWURVHWWHVr appressed to the substratum or, if lobes ascending, then < 2 mm wide or with rhizines but thallus not umbilicate ....... 58 56a Thallus with perithecia (ostioli are visible as black dots at least when wet), never with rhizines, amphibious or terrestrial lichens ........ Dermatocarpon 56b Thallus usually with isidia or apothecia, never with perithecia, with or without rhizines, predominantly terrestrial species, occasionally part of the riparian vegetation and extending inWRWKHÀXYLDO[HULF]RQHLQXSODQGDUHDV ......... 57 57a Thallus with isidia, brown-black (occasionally greenish), usually without apothecia ........................ Umbilicaria deusta (Linnaeus) Baumgarten 1790 57b Thallus grey, usually with apothecia ............................................................ Umbilicaria spp. (frequently at the banks of upland creeks: U. cylindrica (Linnaeus) Delise ex Duby 1830, with abundant marginal cilia). 58a Lobes < 2.0 mm wide, with numerous short rhizines .............................. 59 58b Lobes > 2.5 mm wide, without or with scattered rhizines, usually terrestrial lichens ........................................................................................ not treated 59a Lobes with marginal cilia, ± ascending ................................................... 60 59b Lobes without marginal cilia (but often with protruding rhizines), ± adpressed .................................................................................................... 61 60a Soralia lip-shaped ................... Physcia tenella (Scopoli) de Candolle 1805 60b Soralia helmet-shaped ............ Physcia adscendens (Fries) H. Olivier 1882 61a Medulla K+ yellow, thallus light grey, not changing to green when wet, with bluish-grey laminal soralia ....... Physcia caesia (Hoffmann) Fürnrohr 1839 61b Medulla K- (or rarely K+ red) thallus, grey, brown or greenish, usually turning green when wet ............................................................. Phaeophyscia 62a 7KDOOXV HQWLUHO\ FUXVWRVH PDUJLQDOO\ HI¿JXUDWH VTXDPXORVH RU URVHWWH shaped ..................................................................................................... 66 62b Thallus distinctly lobate, small fruticose or hair-like .............................. 63

32

General Key

63a 7KDOOXV KDLUOLNH ¿ODPHQWRXV ORRVHO\ DWWDFKHG WR WKH VXEVWUDWH RIWHQ DSpearing combed in the direction of running water, photobiont Stigonema .... .............................................................................................. Ephebe lanata 63b Thallus not hair-like ................................................................................ 64 64a 7KDOOXVXPELOLFDWHIRUPLQJVPDOOFXVKLRQVRIÀDWWHQHGVRPHZKDWEUDQFKHG blackish lobes, photobiont a single-celled cyanobacterium ......................... ................................................................................... Gonohymenia heppii 64b 7KDOOXVQRWXPELOLFDWHSKRWRELRQW¿ODPHQWRXVF\DQREDFWHULXPNostoc ... 65 65a Thallus corticate, usually greyish ............................................... Leptogium 65b Thallus not corticate, usually dark olive green or dark brownish olive to blackish .......................................................................................... Collema 66a Thallus squamulose-peltate, margins free, initially crenate, later distinctly lobate, surface tessellate, apothecia opening with pore-like disc, asci polyVSRURXV FRQLGLD ¿OLIRUP SDUDSK\VHV ODFNLQJ QRW DPSKLELRXV EXW SRVVLbly present on steep, ± exposed siliceous rocks along the upper margins of mountain streams and rivers ........................................................................ Phylliscum demangeonii (Mougeot & Montagne in Montagne) Nylander 1855 66b Thallus not squamulose-peltate and with free, lobate margins ................ 67 67a 7KDOOXVURVHWWHVKDSHGRUGLVWLQFWO\HI¿JXUDWHDWWKHPDUJLQ .... Placynthium 67b Thallus crustose, granulose to distinctly areolate, subsquamulose to squamuORVHRUPLQXWHO\FRUDOORLGQRWGLVWLQFWO\URVHWWHVKDSHGRUHI¿JXUDWH I ...... 68 68a Fruit bodies perithecia, perithecial wall dark, ascospores uniseptate, thallus uniformly crustose .................................................................................. 69 68b Fruit bodies apothecia, if apothecia appearing perithecioid with pore-like disc, then with pale proper exciple, ascospores septate or not, thallus crustose to small squamulose ........................................................................ 70 69a Paraphyses sparingly branched, never anastomosing ... Pseudarthopyrenia 69b Paraphyses richly branched and anastomosing ................... Pyrenocollema 70a Thallus usually with a distinct dark hypothallus, sometimes also with a dark bluish prothallus, apothecia lecideine, ascopsores septate, hypothecium dark reddish brown ........................................................................ Placynthium 70b Thallus having neither a dark hypothallus nor a dark blue prothallus, apothecia usually lecanorine, rarely biatorine, proper margin absent or very indistinct, rarely thickened and usually pale, rarely darkly pigmented but never carbonized, hypothecium pale, ascospores simple or septate .................. 71

General Key

33

71a Apothecial discs black, umbonate to gyrose, proper exciple at least apically distinctly coloured .................................................................................. 72 71b Apothecial discs brownish, not or rarely umbonate, never gyrose, proper exciple – if at all present – pale brownish ............................................... 73 72a 3KRWRELRQW ¿ODPHQWRXV 5LYXODULDFHDH  WKDOOXV VPDOO VTXDPXORVH WR VXEVTXDPXORVHLUUHJXODUO\VKDSHGDSRWKHFLDVPDOOLPPHUVHGGLVF ¿QDOO\ZLWK distinct central umbo, hymenium I-; submerged in cold mountain streams ....................................................................................... Thelignya lignyota 72b Photobiont single-celled, thallus crustose, composed of thick, angulose areoles dying from the base, apothecia rather conspicuous, immersed to adnate, disc umbonate to gyrose, hymenium I+ blue; on rock moist from seeping water, in the British Isles also on boulders in streams .................................. ........................................................ Metamelanea umbonata Henssen 1989 73a Ascospores septate, apothecia sessile, thallus thin, crustose, usually growing inundated ....................................................................... Leptogium aquale 73b Ascospores simple, apothecia sessile or ± immersed, thallus thin or thick, growing frequently or rarely inundated ................................................... 74 74a *URZLQJRYHUPRVVHVDORQJURFN\ULYHUPDUJLQVSKRWRELRQWWKH¿ODPHQWRXV cyanobacterium Nostoc, thallus occasionally submerged ............................. ........................................................................... Lempholemma polyanthes 74b $OZD\VJURZLQJGLUHFWO\RQURFNSKRWRELRQWDVLQJOHFHOOHGRU ¿ODPHQWRXV cyanobacterium (Rivulariaceae or Stigonema) but never Nostoc, thallus frequently or occasionally submerged ......................................................... 75 75a 3KRWRELRQW WKH ¿ODPHQWRXV F\DQREDFWHULXP Stigonema, thallus appearing crustose, composed of short, erect to ascending threads forming rather compact areoles, asci polysporous, apothecia globose, very small ....................... ......................................................................................... Ephebe multispora 75b 3KRWRELRQW D VLQJOHFHOOHG F\DQREDFWHULXP RU ¿ODPHQWRXV 5LYXODULDFHDH  with tapered, much contorted and split chains, thallus areoles compact, sometimes with vertical outgrowths but never composed of erect threads, asci 8-spored (in the species treated here) ............................................... 76 76a Thallus dark reddish to dark reddish-brown or almost black, photobiont cells ZLWKGDUNUHGGLVKWRSXUSOLVKJHODWLQRXVVKHDWK¿J ........ Pyrenopsis 76b Thallus blackish or dark olivaceous to brownish, photobiont cells with yelORZLVKEURZQJHODWLQRXVVKHDWK¿J ............................................... 77

34

General Key

77a Apothecia sessile, conical to eventually roundish and disc-like, apothecial GLVFDW¿UVWSXQFWLIRUPODWHURSHQDQGHYHQWXDOO\XPERQDWHSURSHUH[FLSOH distinct from the beginning, pale, thalline margin persisting, thallus dark brownish to olivaceous-brown, areoles very thin, small and smooth ........... ......................................................................... 3\UHQRFDUSRQÀRWRZLDQXP 77b Apothecia immersed to semi-sessile, sometimes at the tips of minute erect thallus outgrowths, discs punctiform to opened, proper exciple distinct or not, thalline margin receding or persisting, thallus blackish or dark olivaceous, usually not brownish, areoles thin or thick, usually not very small and smooth .................................................................................................... 78 78a Thalline and proper margin of the apothecia roughly equally thick, thalline margin eventually receding and proper margin becoming prominent, apically pale or brownish, paraphyses usually thin, indistinctly septate and apically hardly thickened, apothecia semi-immersed to sometimes at the tips of erect vertical thallus outgrowths, apothecial discs pale to dark reddish EURZQSKRWRELRQW¿ODPHQWRXV5LYXODULDFHDH EXWFKDLQVRIWHQKHDYLO\VSOLted and appearing single-celled ................................................ Porocyphus 78b Thalline margin of the apothecia persisting, always distinctly thicker than the proper exciple, proper margin remaining thin or virtually absent, never prominent, paraphyses usually robust, distinctly septate with thickened apical cells, apothecia immersed to semi-immersed, rarely sessile, apothecial discs pale to dark reddish brown or black, photobiont a single-celled cyanobacterium ................................................................................................ 79 79a $SRWKHFLDOGLVFVEODFNLVKWKDOOXVUHJXODUO\DUHRODWHDUHROHVÀDWZLWKVKDUS HGJHV VXUIDFH YHU\ ¿QHO\ ZDUW\ IXUIXUDFHRXV  EXW QRW JUDQXORVH WKDOOXV anatomy paraplectenchymatous with the symbionts arranged in vertical rows; amphibious or on rock moist from seeping water ............................... ........................................................................... Pterygiopsis concordatula 79b Apothecial discs yellowish to dark reddish brown, usually not blackish, thallus variously shaped, thallus anatomy paraplectenchymatous to densely reticulate with no vertical arrangement of the symbionts; not amphibious but possibly present on ± exposed siliceous or calcareous rock along the upper banks of rivers, dikes and lake shores .... Psorotichia A. Massalongo 1855

.H\VWRWKH6SHFLHVDQG6SHFLHV3UR¿OHV LQDOSKDEHWLFRUGHU Figure numbers in standard refer to photos, ¿gure numbers in italics to line drawings of involucrellum outlines.

Anisomeridium (Müller Argoviensis) M. Choisy 1928 Important references: Harada (1999), Harris (1990, 1995), Keissler (1936, 1937). Key to the species: 1a Thallus distinct, 30–50 m, usually with perithecia, pycnidia unknown, ascospores 9–15 × 3.6–7 m, amphibious species ............... A. carinthiacum E Thallus very thin, inconspicuous, conical pycnidia usually present, perithecia are rare in thalli from terrestrial habitats, unknown from riparian collections, ascospores (12) 14–20 (23) × (3) 4.5–5 (6) m, terrestrial (epiphytic) species, very rare among riparian vegetation in deeply shaded habitats ....... .............................................. A. polypori (Ellis & Everhart) M.E. Barr 1996

Anisomeridium carinthiacum (J. Steiner) R.C. Harris in Egan 1987 ¿J 10) Basionym: Arthopyrenia carinthiaca J. Steiner 1913. Synonyms: Paraphysothele carinthiaca (J. Steiner) Keissler 1937, Thelidium sublacteum Eitner 1911, Paraphysothele sublacteum (Eitner) Zschacke 1933, Paraphysothele kalenskyi Servít 1953, Ditremis carinthiaca (J. Steiner) R.C. Harris 1990, Anisomeridium japonicum Harada 1999.

Fig. 10. Anisomeridium carinthiacum: ×18.

36

.H\VWRWKHVSHFLHVDQGVSHFLHVSUR¿OHV

7KDOOXV whitish-grey, greenish-grey; 30–50 m thick; paraplectenchymatous but not subgelatinous; prothallus absent; photobiont yellowish-green or greenish, cells 5–18 m (TrentepohliaVSYHUL¿HG E\FXOWLYDWHGLVRODWHV Perithecia: immersed to semi-immersed; upper part exposed or covered by a thin thalline layer; 90–160 × 50–150 m; involucrellum 130–235 × 45–90 m, 12–70 m thick; asci 40–57 × 11–15 m; ascospores bi-seriate, uniseptate, cells of equal size or upper cell shorter and wider, 9–15 × 3.6–7 m, colourless. (FRORJ\ submerged or little above water level on siliceous rocks and pebbles in upland streams. Restricted to shaded localities. 'LVWULEXWLRQ very rare in humid mountain ranges, so far only known from a few localities in Germany (Hesse) and Austria (Styria, Carinthia). 6LPLODU VSHFLHV Pyrenocollema saxicola is a predominantly terrestrial species with larger ascospores (18–24 × 4–5 m), restricted to base rich substrata (usually limestone, but also on silty wood). Arthopyrenia inconspicua has extremely small perithecia (30–100 m) without involucrellum, a dark greenish-black thallus, and larger ascospores (12–19 × 4–6 m). Collemopsidium angermannicum, a freshwater lichen known from Nordic countries, is associated with cyanobacteria. It has a dark thallus, absent involucrellum and much larger ascospores. Other freshwater species of Anisomeridium have not been reported for Europe so far. Anisomeridium laevigatum has a thicker thallus (60–140 m), larger perithecia (170–320 m), asci (63–83 m) and ascospores (12–22.5 m). Anisomeridium SDFL¿FD has larger perithecia (240–440 m), asci (68–83 m) and ascospores (15–21.5 m). Anisomeridium distans has a scant to absent thallus, ascospores becoming 4-celled when older.

$VSLFLOLDA. Massalongo 1852 Important references: Clauzade & Roux (1985), Hafellner (1991), Pereira (1992), Purvis et al. (1992), Wirth (1995). Key to the species: 1a Medulla P+ orange, with stictic acid and ± norstictic acid, not in the submerged zone ............................................................................... $ODHYDWD E Medulla P-, with aspicillin, from terrestrial to submerged zones .............. 2 2a Asci usually with 6 ascospores, hymenium (80) 120–150 Pm high, pycnospores 6.5–12 PPORQJVWUDLJKWIURPÀXYLDOPHVLFWRWHUUHVWULDO]RQHV ...... ............................ A. caesiocinerea (Nylander ex Malbranche) Arnold 1886 E Asci usually with 8 ascospores, hymenium (110) 130–235 Pm high, pycnospores 10.5–14 or 26–40 PPORQJVWUDLJKWRUFXUYHGVXEPHUJHGWRÀXYLDO PHVLF]RQHGHWHUPLQDWLRQGLI¿FXOWZLWKRXWS\FQLGLD .............................. 3 3a Ascospores 22–35 × 14–18 Pm, m pycnospores 10.5–15 Pm long, straight .... .................................................................................................. A. aquatica m pycnospores 26–40 Pm long, slightly curved E Ascospores 29–34 × 18–21 Pm, ............................................................................................. A. supertegens

Aspicilia

37

$VSLFLOLDDTXDWLFD .|UEHU¿J 11) Synonyms: Lecanora aquatica (Körber) Hepp 1867, Pachyospora aquatica (Körber) Arnold 1869. 7KDOOXV grey greenish-grey, (light) brownish-grey; thick, rimose to areolate; surface smooth and even, cortex and medulla K-, P-, no lichen substances detected by t.l.c.; photobiont coccoid green algae (Trebouxia impressaYHUL¿HGE\ cultivated isolates and ITS-sequences, Coccomyxa according to Pereira 1992). Apothecia: immersed, hymenium (110) 150–200 m high; asci 8-spored; ascospores unicellular, 22–27 (35) × 14–18 m. Conidia: 12–15 × 1 m, straight. (FRORJ\ amphibious on periodically inundated rocks or in the splash water zone on stable siliceous substrates in cool upland streams. Best developed in full sunlight. 'LVWULEXWLRQ in high-montane to alpine regions; widely distributed in watercourses with stable bedrock in the Alps and Carpathians; elsewhere very rare and on the verge of extinction at lower altitudes. 6LPLODUVSHFLHV Aspicilia laevata has a P+ orange, K± yellow medulla (stictic acid and ± norstictic acid detected by t.l.c.) and smaller ascospores (13–23 m). Aspicilia caesiocinerea is a terrestrial species which is often present among riparian associations and also occurs in the splash water zone but never in sites with longer submersion periods. It usually has a lower hymenium (80–150 m, rarely up to 200 m), asci with predominantly six ascospores and smaller pycnospores (6.5–12 m). The medulla contains aspicilin (detected by t.l.c.).

Fig. 11. Aspicilia aquatica: ×15.

38

.H\VWRWKHVSHFLHVDQGVSHFLHVSUR¿OHV

$VSLFLOLDODHYDWD$FKDULXV $UQROG¿J 12) Basionym: Sagedia laevata Acharius 1809. Selected synonyms: Lecanora laevata (Acharius) Nylander 1866, Urceolaria laevata (Acharius) Duby 1830, Lecanora lusca Nylander 1872, Aspicilia lusca (Nylander) de Lesdain 1905, Aspicilia sylvatica Arnold 1887, Lecanora sylvatica (Arnold) Sandstede 1912. 7KDOOXV grey to olive or greenish-grey; thin to thick, smooth, continuous to cracked; medulla P+ orange, K± yellow; stictic acid detected by t.l.c., in some thalli also norstictic acid; photobiont coccoid green alga. Apothecia: emergent or immersed and crater like, up to 500 m in diameter; asci 8-spored; ascospores unicellular, 13–17 (23) × 9–13 m. Conidia: 18–25 × 0.8 m, straight. (FRORJ\ a terrestrial species on usually shaded siliceous rocks at humid sites, often part of the riparian vegetation but also amphibious in the upper splash water zone of streams in woodlands. 'LVWULEXWLRQ widely distributed but amphibious populations only locally abundant. 6LPLODUVSHFLHVAll other species of Aspicilia occurring in the splash water zone have a P- and K- medulla and lack stictic acid. Some thalli have an olive tinge which is usually not observed in the strictly amphibious A. aquatica and the terrestrial A. caesiocinerea.

Fig. 12. Aspicilia laevata: ×13.

Aspicilia

39

$VSLFLOLDVXSHUWHJHQV$UQROG¿J 13) Synonyms: Lecanora supertegens (Arnold) Zahlbruckner 1928, Lecanora gibbosa var. supertegens (Arnold) Migula 1926. Uncertain species: Aspicilia inornata Arnold 1877. 7KDOOXV greyish-green; thick, cracked to areolate, smooth to rather uneven; photobiont coccoid green alga. Apothecia: immersed in the thallus, 0.2–2.0 mm in diameter, discs black; hymenium 130–140 m high; epithecium green-olive, N+ emerald-green; asci 8-spored; ascospores unicellular, 17–24 × 10–16 m. Conidia: 31–37 × 1 m, slightly curved. (FRORJ\ amphibious on siliceous rocks in the splash water zone and at temporarily inundated sites. 'LVWULEXWLRQ a rare species preferring upland areas with oceanic climate. 6LPLODU VSHFLHV separated from Aspicilia aquatica by the wider ascospores, much longer and slightly curved conidia. Uncertain species: Aspicilia inornata differs from A. supertegens mainly in the uneven thallus and in colour nuances. Whether the former is a good species requires further studies.

Fig. 13. Aspicilia supertegens: ×9.

40

.H\VWRWKHVSHFLHVDQGVSHFLHVSUR¿OHV

Bacidina 9Č]GD Important references:3XUYLVHWDO 9Č]GD :LUWK  Notes: The key includes species of Bacidina and a selection of species from the similar genus Bacidia. The genera are separated by the structure of the exciple (paraplectenchymatous in Bacidina, prosoplectenchymatous in Bacidia) and the axial mass in the ascus tip (rounded in Bacidina, conical in Bacidia). The characters used in the key are easier to observe and also applicable for sterile specimens with pycnidia. Key to the species: 1a Apothecia present, thallus with or without pycnidia ................................. 2 E Apothecia absent, thallus with pycnidia only ........................................... 7 2a Hypothecium colourless to yellowish-brown ............................................ 3 E +\SRWKHFLXPUHGEURZQWRGDUNEURZQDSRWKHFLDÀDWRUFRQYH[ZKHQROG usually terrestrial species, occasionally extending inWRWKHÀXYLDO[HULF]RQH ................................................................................................................... 5 3a Apothecia small, up to 0.2 mm in diameter, with unicellular to 3-septate ascospores ..............................B. chloroticula 1\ODQGHU 9Č]GD & Poelt 1990 E Apothecia larger, 0.4–0.6 mm in diameter, with 4–8-septate ascospores .... 4 4a Apothecia frequent, ascospores 24–43 × 2–2.5 m, true exciple evident at ¿UVWEXWDSRWKHFLDVRRQEHFRPLQJFRQYH[DQGH[FLSOHWKHQRIWHQEHFRPLQJ indistinct, pycnidia immersed, rare, of two types (whitish or brown) with conidia > —PORQJIUHTXHQWLQWKHÀXYLDOPHVLFWRÀXYLDO[HULF]RQHV ... .................................................................................................. B. inundata E Apothecia rare, ascospores 14–28 × 3–4 m, true exciple persistent, pycnidia frequent, sessile, whitish-pubescent with conidia 4–5.7 m long, terrestrial species of humid overhangs, occasionally at the banks of streams ... ..................................... Bacidia carneoglauca (Nylander) A.L. Smith 1911 5a Ascospores 11–16 (20) m, apothecia black ................................................ .................................................... Bacidia trachona (Acharius) Lettau 1912 E Ascospores larger (17–55 m), apothecia variously coloured .................. 6 6a Epihymenium yellowish to colourless, apothecial disc light grey to greyEURZQRUGDUNUHGEURZQDSRWKHFLDOGLVF“ÀDW ........................................... ........................................... B. arnoldiana .|UEHU 9:LUWK 9Č]GD E Epihymenium olive, blue-green or ± colourless, apothecial disc grey-brown, black-brown, red-brown to black, occasionally convex when old ................ .............................................................. B. egenula1\ODQGHU 9Č]GD 7a Pycnidia ± immersed, whitish or brown with conidia > 10 m long, frequent LQWKHÀXYLDOPHVLFWRÀXYLDO[HULF]RQHV ................................. B. inundata E Pycnidia sessile, whitish-pubescent with conidia 4–5.7 m long (see also E .............................. Bacidia carneoglauca (Nylander) A.L. Smith 1911

Bacidina

41

Bacidina inundata )ULHV 9Č]GD¿J14) Basionym: Biatora inundata Fries 1822. Selected synonyms: Bacidia inundata (Fries) Körber 1855, Woessia inundata (Fries) Sérusiaux & Diederich 1998. 7KDOOXV grey-green, also yellowish-green; rimose-continuous to granular-warty, RIWHQ ZLWK H[WHQVLYH ZKLWH DQG ¿PEULDWH SURWKDOOXV 89 SKRWRELRQW FRFFRLG green alga, clustered in groups, 5–17 × 5–12 m. Apothecia:VHVVLOHÀDWWRFRQYH[±PPLQGLDPHWHUSDOHEURZQWRDOPRVWEODFNLVKWUXHH[FLSOHDW¿UVW distinct, sometimes becoming excluded with age; hymenium 45–55 m high, colourless to pale brown, K± purplish; hypothecium colourless, pale straw or pinkish-brown in upper part; ascospores 3 (7)-septate, 24–43 × 2.0–2.5 m. Pycnida: rare, 100–200 m in diameter, of two types: whitish, with conidia 23–55 × 0.5–1.0 m, or brownish with conidia 11–19 × 1 m in size. (FRORJ\ amphibious in the splash water zone, but also tolerating several months of being submerged; in shady to exposed streams, rivers and at lake margins on siliceous rocks; tolerant to moderate eutrophication and a rather strong competitor; occasionally growing as an epiphyte on other lichens or forming mechanical hybrids, especially with Verrucaria spp.; absent from strongly acidic waters (pH <  DOVRVHQVLWLYHWRDWPRVSKHULFDFLGL¿FDWLRQ 'LVWULEXWLRQ from lowland to (sub-) alpine areas throughout Central Europe. Notes: Bacidina inundata occasionally forms mechanical hybrids, e.g. with Verrucaria spp. (Thüs 2002). These mixed thalli have intermediate thallus characters. The clustered arrangement of the photobiont cells and the development of FKDUDFWHULVWLFDSRWKHFLDKHOSLQWKHLGHQWL¿FDWLRQRIBacidina inundata in these compound systems.

Fig. 14. Bacidina inundata: ×16.

42

.H\VWRWKHVSHFLHVDQGVSHFLHVSUR¿OHV

6LPLODU VSHFLHV There are several predominantly terrestrial species which are found occasionally in amphibious habitats. Bacidina arnoldiana and B. egenula have darker hypothecia and smaller thallus granules. Bacidia carneoglauca has ascospores 14–28 × 3–4 m in size and frequently forms pycnidia with small conidia (4–5.7 m long). Bacidina chloroticula has small apothecia (up to 0.2 mm in diameter) with a colourless exciple and unicellular, up to 3-septate ascospores. Bacidia trachona has small black apothecia and short ascospores (11–16 (20) m). Bacidina egenula, B. chloroticula and Bacidia trachona are reported to occur at similar sites like Bacidina inundata and are locally common in Upper Austria (F. Berger, pers. comm.) but may have been widely overlooked in amphibious habitats.

&DORSODFD Th. Fries 1860 nom. cons. Important references: Purvis et al. (1992), Wirth (1995). Key to the species LQFOXGLQJWKHDPSKLELRXVDQGDVHOHFWLRQRIULSDULDQVSHFLHV  1a Thallus yellow or orange ............................................................................. ............................. Caloplaca spp., terrestrial (riparian) species, not treated E Thallus not yellow or orange .................................................................... 2 2a Thallus thick, distinctly epilithic or lichenicolous .................................... 3 E Thallus thin and inconspicuous or ± endolithic (predominantly terrestrial species, a selection of riparian taxa is keyed out here) .............................. 4 3a Thallus grey to black, occasionally lichenicolous on other crustose lichens, without isidia, usually with apothecia ............... &DWURÀDYD var. VXEPHUVD E Thallus greyish, usually with a bluish tinge, never lichenicolous, with numerous minute isidia or blastidia, usually found sterile in freshwater habitats ........................................................................................... C. isdiigera 4a Ascospores with thick septum (> 25% of total length; 2.5–5 Pm), tips of paraphyses swollen (4–5 Pm) .... C. holocarpa (Hoffmann) A.E. Wade 1965 E Ascospores with thin septum (< 20% of total length; 1.5–3.5 Pm), tips of paraphyses moderately thickened (2–3 Pm) .............. C. arenaria (Persoon) Müller Argoviensis 1862 / subpallida H. Magnusson 1945.

&DORSODFDDWURÀDYD var.VXEPHUVD 1\ODQGHU 0DJQXVVRQ¿J 162) Basionym: Lecanora pyracea f. submersa Nylander 1855. Synonyms: Placodium pyraceum f. submersum (Nylander) A.L. Smith 1918, Caloplaca pyracea f. submersa (Nylander) Boistel 1931. 7KDOOXV grey to almost black, cracked to areolate, prothallus black; photobiont coccoid green alga. Apothecia:ÀDWFRQVWULFWHGDWWKHEDVH±—PLQGLameter, discs brownish-orange, with bright orange true exciple and usually an additional grey thalline exciple; ascospores polarilocular, 12–17 × 8–10 m, septum 5–7 m thick.

Collema

43

(FRORJ\amphibious in the splash water zone of streams and rivers on siliceous rocks, usually at well lit places. 'LVWULEXWLRQ rare but widespread. 6LPLODU VSHFLHV The terrestrial species Caloplaca holocarpa has an indistinct thallus, narrower ascospores and a thallus exciple is lacking. Caloplaca arenaria has yellowish coloured, smaller apothecia and narrower ascospores (5–8 m) with a thin septum (2–3 m). Caloplaca chlorina has larger apothecia (up to 1.5 mm) and thin ascospore septa (3–5 m).

&DORSODFDLVLGLLJHUD 9Č]GDµ¶¿J161) 7KDOOXV greyish, often with a bluish tinge, thin to thick, cracked, surface uneven, covered by minute spherical to coralloid blastidia or isidia (best seen under a hand lens); thallus and isidia K+ pale violet-red; photobiont coccoid green alga (Trebouxia VSYHUL¿HGE\FXOWLYDWHGLVRODWHV  Apothecia: unknown. (FRORJ\ a terrestrial species, occasionally in the uppermost part of the splash water zone and possibly better treated as an element of the riparian vegetation, but also found in regularly inundated sites together with Staurothele frustulenta. 8VXDOO\IRXQGDWHXWURSKLFDUWL¿FLDOEDQNVRIODUJHUVWUHDPVRUDWZHOOOLWSODFHV along river banks with a thin crust of silt. 'LVWULEXWLRQ only a few amphibious populations are known from Central Europe (river Elbe in and north of Hamburg, river Nahe east of Bad Münster am Stein, river Lauter north of Kaiserslautern) but certainly much overlooked. 6LPLODUVSHFLHV There is no other freshwater species with minutely isidiate/blastidiate thallus reacting K+ pale violet-red. Caloplaca isidiigera was placed in synonymy to C. chlorina by Coppins (2002). This step has not been followed by the authors of Central European checklists (e.g. Wirth et al. 2008).

&ROOHPD Weber ex F.H. Wiggers 1780 nom. cons. Important references: Degelius (1954, 1974), Jørgensen (2007a), Poelt (1969), Purvis et al. (1992), Wirth (1995). Key to the species: 1a Permanently submerged or drying out only very shortly, lobes narrow and UHSHDWHGO\ IXUFDWH FRQYH[ WR ÀDWWHQHG LVLGLD ODFNLQJ DSRWKHFLD XVXDOO\ present ............................................................................... C. dichotomum E Usually not permanently submerged but often found in the upper belt of ULYHUPDUJLQVHWFOREHVEURDGHQHGFRQFDYHWRÀDWWHQHGLVLGLDXVXDOO\SUHVent, apothecia rarely observed or unknown ............................................... 2

44

.H\VWRWKHVSHFLHVDQGVSHFLHVSUR¿OHV

2a 7KDOOXVOREHVFHQWUDOO\EHFRPLQJREVFXUHG E\GHQVHO\FURZGHG¿QDOO\FRUalloid isidia, lobes rather small, few to 10 mm wide, concave, somewhat undulating, apothecia unknown ........................................ &JOHEXOHQWXP E Lobes not obscured by crowded isidia, isidia globose or scale-like, abundant or sparse, lobes broadly lobate, usually > 5 mm wide, apothecia present or not ............................................................................................................. 3 3a Lobes usually ± undulating and pustulate, isidia sparse, globose, usually restricted to terrestrial habitats but also riparian along the margins of larger rivers ................................................................................... &IXVFRYLUHQV E Lobes radially folded, not undulating and pustulate, isidia numerous, scalelike, on the mossy base of trees, siliceous boulders in humid, shady places often along streams ................................................................ &ÀDFFLGXP

&ROOHPDGLFKRWRPXP (Withering) Coppins & J.R. Laundon 1984 ¿JV15–16) Basionym: Tremella dichotoma Withering 1776. Synonyms: &ROOHPD ÀXYLDWLOH (Hudson) Steudel 1824, Leptogium rivulare auct. brit. non (Acharius) Montagne, /LFKHQÀXYLDWLOLV Hudson 1778; for full synonymy see Degelius (1954, 1974). 7KDOOXV bright to dark green, sometimes glaucous, dull, epruinose, small, to 2.5 cm, but much larger when moist, roundish, loosely attached by rounded hapters or crowded rhizines, foliose, deeply lobate; lobes repeatedly furcate, tips GLVFUHWHRUSDUWO\LPEULFDWHVRPHWLPHVEURDGHQHGDWWKHWLSV ÀDWWHQHGRUFRQYH[ c. 0.5–1.5 (4) mm broad, 170–430 m thick when moist; photobiont Nostoc with rather short trichomes, crowded towards the thallus surface. Apothecia: numerous and crowded or sparse and scattered, sessile, with constricted base, roundish, plane, 0.7–1 mm wide; thalline margin rather thin, entire, persisting, smooth to minutely granulose or knobby, occasionally lobulate, not or only slightly promi-

Fig. 15. Collema dichotomum: well developed, richly fertile thallus, Scotland; ×5.

Collema

45

Fig. 16. Collema dichotomum: irregular thallus, sparingly fertile, Upper Austria; ×6. nent; disc plane, smooth, not glossy, dark to rather pale red; excip. prop. thick, euparaplectenchymatous, cells 8–20 m; hymenium c. 150 Pm tall, I+ blue, upper part yellowish to reddish brown; ascospores 8, sometimes fewer, ellipsoid or sometimes ovoid, poles acute or obtuse, 4-celled to usually submuriform, usually not constricted at septa, straight, hyaline, 20–30 × 8.5–13 m. Pycnidia:VXSHU¿cial, immersed or partly prominent, c. 300–550 m; pycnospores straight, bacilliform or slightly thickened at the ends, 5–6.5 (8) × 1.2–1.5 m. (FRORJ\ usually permanently inundated, on siliceous rock in unpolluted, clear, cold rivers or mountain streams; also in the splash zone of water courses very rarely drying out. 'LVWULEXWLRQ montane to alpine in Central Europe; known from the Alps (?), Upper Austria highest elevations of Black Forest and Vosgues, Ardennes, Tatra and Sudetes; extincted in the Eifel and Harz Mountains; occurs also in Great Britain and Ireland, Fennoscandia, Karelia (Russia), western Romania and North America. 6LPLODUVSHFLHV unlikely to be confused with any other lichen from aquatic habitats; the dark green, dichotomously branched rosettes with convex lobes and numerous apothecia are characteristic.

&ROOHPDÀDFFLGXP$FKDULXV $FKDULXV¿J17) Basionym: /LFKHQ ÀDFFLGXV Acharius 1795. Synonyms: for full synonymy see Degelius (1954, 1974). 7KDOOXV dark olive green to blackish, dull, epruinose, few to 6 cm across, ± roundish, loosely attached by hapters, foliose, ± deep and broadly lobate; lobes

46

.H\VWRWKHVSHFLHVDQGVSHFLHVSUR¿OHV

Fig. 17. &ROOHPDÀDFFLGXP: scale-like isidia; ×13 ÀDWWHQHGRUEHFRPLQJFRDUVHO\DQGLUUHJXODUO\IROGHGDQGXQGXODWHPDUJLQQRW VZROOHQ LVLGLD QXPHURXV VXSHU¿FLDO WR PDUJLQDO VTXDPLIRUP c. 0.2–0.5 mm, lobes c. 0.5–1.5 (3) cm broad, thin to membranaceous, 70–170 m thick when moist; photobiont Nostoc trichomes crowded towards the thallus surface. Apothecia: usually rare, sometimes numerous, scattered on surface, sessile, with constricted base, roundish, plane, to 1.5 (2.5) mm wide; thalline margin thin, ± entire, smooth to somewhat rugulose, not or only slightly prominent, persistent or disappearing in old, convex apothecia; disc plane, smooth, usually not glossy, pale to dark red, epruinose to rarely white pruinose especially in juvenile apothecia; excip. prop. rather thin, subparaplectenchymatous to rarely almost euparaplectenchymatous, cells 10.5 (17) m; hymenium c. 90–130 Pm tall, I+ blue, upper part yellowish to reddish-brown; ascospores 8, narrowly ellipsoid to fusiform, usually straight, ends acute, sometimes extended, 4–6-celled, septa not constricted, hyaline, (20) 26–34 (40) × 6–6.5 m, sometimes constantly larger 34–45 × 6.5–8.5 m. Pycnidia: laminal, immersed, globose, c. 100–210 m; pycnospores straight, bacilliform or slightly thickened at the ends, 4–4.5 (6) × 1– 1.5 (1.8) m. (FRORJ\ predominantly in shady, humid places, on usually mossy old trunks, mossy siliceous and more rarely calcareous rocks and boulders, in seepage tracks and rarely inundated in or at streams, rivers or lake shores; see Thüs (2002) for ecological details. 'LVWULEXWLRQ widely distributed in the northern hemisphere from sea level to mountain regions. 6LPLODUVSHFLHV Leptogium cyanescens has similar scale-like isidia, but the thallus is distinctly corticate and greyish.

Collema

47

&ROOHPDIXVFRYLUHQV :LWKHULQJ -5/DXQGRQ¿J18) Basionym: Lichen fuscovirens Withering 1776. Synonyms: Collema tuniforme $FKDULXV $FKDULXVµtunaeforme¶ IRUIXOOV\QRQ\P\VHH'HJHOLXV VXEµC. tunaeforme¶ DQG'HJHOLXVVXEC. tuniforme). 7KDOOXV dark olive green to blackish, usually dull, epruinose, (1) 3–5 (10) cm DFURVV“URXQGLVKRULUUHJXODU“DGQDWHWRWKHVXEVWUDWHDWWDFKHGE\FRQÀXHQW hapters or scattered rhizines, foliose, deeply and rather broadly lobate; lobes ± repeatedly and usually irregularly furcate, lobules 2–5 mm wide, (70) 110–200 (300) m thick when moist, imbricate or discrete, usually broadly canaliculate ZLWKDVFHQGLQJDQGXVXDOO\XQGXODWLQJ PDUJLQVUDUHO\OREXOHVPRUHÀDWDQGGHpressed, ± irregularly and minutely pustulate, margin not swollen, isidia numerRXV VXSHU¿FLDO WR PDUJLQDO XVXDOO\ JORERVH PP ZLGH UDUHO\ VRPHZKDW clavate or teretiform but not branched; photobiont Nostoc trichomes crowded towards the thallus surface. Apothecia: numerous or lacking, laminal, sessile, ¿QDOO\ZLWKFRQVWULFWHGEDVHEXWQRWVWLSLWDWHURXQGLVKWR± PPZLGH thalline margin thin, entire, smooth to isidiate, usually not prominent, generally persisting; disc ± plane, smooth, usually not glossy, dark red to reddish brown or brown, epruinose; proper exciple in central parts euparaplectenchymatous, 45–110 m thick, cells usually (6.5) 10–20 (30) m large, in marginal parts to 65 m thick with slightly smaller cells; hymenium 85–130 Pm tall, I+ blue, upper part yellowish to reddish-brown; ascospores 8, usually broad ellipsoid, straight, ends rounded to obtuse, rarely rather acute, submuriform with 3 transversal septa and 1 longitudinal septum, rarely eumuriform with > 3 transverse and 2–3 longitudinal septa, rarely 4-celled with no longitudinal septum, not or slightly constricted at septa, hyaline, 12–24 (28) × 6.5–13 (15) m. Pycnidia: laminal or marginal, immersed to slightly elevated, ± globose, 130–260 m; pycnospores straight, bacilliform or slightly thickened towards the ends, 4.5–6 × 1.5 m.

Fig. 18. Collema fuscovirens: ×2.

48

.H\VWRWKHVSHFLHVDQGVSHFLHVSUR¿OHV

(FRORJ\ usually on bare rock, rarely on pebbles or over mosses, predominantly on calcareous or calciferous rock, very rarely on acidic siliceous rock. This is not an aquatic or amphibious species but it occurs riparian along the upper belt of larger rivers where it is occasionally sprayed or inundated by nutrient rich and muddy water. The species is also known to grow seasonally inundated in the Alvar regions of Öland and Gotland. 'LVWULEXWLRQ widely distributed across the northern hemisphere with occurrences from sea level to alpine regions.

&ROOHPDJOHEXOHQWXP1\ODQGHUH[&URPELH 'HJHOLXV¿J19) Basionym: Leptogium glebulentum Nylander ex Crombie 1882. Synonyms: Collema alpinum Th. Fries 1866, Collema coralliferum Degelius 1944, Collema furvellum Räsänen 1940, Collemodium glebulentum (Nylander ex Crombie) Nylander ex Crombie 1886. 7KDOOXV dark olive green to blackish, sometimes more bright green or brownish, dull, epruinose, membranaceous, lighter and semi-transparent when moist, usually 3–6 cm across, roundish or irregular, closely to loosely attached by hapters, foliose, ± deeply lobate; lobes ± numerous and dense, rounded or usually extended, few to 10 mm wide, imbricate or ascending to irregularly folded and coarsely undulate, margin entire, not swollen, isidia always present, very numerRXVVXSHU¿FLDOWRPDUJLQDOWHUHWLIRUPWRHYHQWXDOO\EUDQFKHGDQGFRUDOORLG c. 0.1 mm broad, to 1.5 mm long, eventually densely crowded, covering central thallus parts and obscuring the thallus lobes, lobes thin, membranaceous, (50) 65–135 (200) m thick when moist; photobiont Nostoc. Apothecia: unknown. Pycnidia: very rare, often poorly developed.

Fig. 19. Collema glebulentum: foliose thallus with undulating lobes and crowded, erect isidia; ×10.

Dermatocarpon

49

(FRORJ\ on shortly to periodically inundated siliceous to slightly calciferous, occasionally somewhat soil-incrusted rocks and boulders along mountain streams and rivers or lake-shores as well as on steep rock faces moistened from seeping water; usually found in more moist situations than &ROOHPDÀDFFLGXP. 'LVWULEXWLRQ in Central Europe montane to alpine, very rare and scattered, southern Black Forest, Allgäu, Sudetes and Switzerland; also in Great Britain and Ireland, Fennoscandia, Iceland, Greenland and Novaya Zemlya.

Dermatocarpon Eschweiler 1824 Important references: Heiðmarsson (1996, 1998, 2001, 2003), Orange (1998), Poelt (1969). Key to the species: 1a Thallus multilobed, lobes 4–25 mm wide, with many holdfasts scattered on the lower surface, thallus turning bright green when wet¿J , medulla WXUQLQJUHGZLWK0HOW]HU¶V5HDJHQWXSSHUWKDOOXVVXUIDFHQHYHUSUXLQRVH ... ................................................................................................... 'OXULGXP E Thallus monophyllous, umbilicate, large (> 10 mm in diameter) or with VPDOO ±PP  VTXDPXORVH WKDOOL LQGLYLGXDO WKDOOL VRPHWLPHV GLI¿FXOW WR discern, wet thallus turning greenish or remaining greyish to brownish ¿J PHGXOODQRWWXUQLQJUHGLQ0HOW]HU¶V5HDJHQWXSSHUWKDOOXVVXUface pruinose or not ................................................................................... 2 2a Very rare species with small squamulose thalli, 2–7 mm diameter, typically IRUPLQJ GHQVH FRORQLHV DQG WKHQ LQGLYLGXDO WKDOOL GLI¿FXOW WR GLVFHUQ DQG giving the colony an areolate appearance, thallus pruinose or not ............... ........................................................................................ 'OHSWRSK\OORGHV E Widespread species with distinct foliose thalli, not forming dense colonies of areolate appearance, thalli small to large (2–50 mm or more) .............. 3 3a Ascospores 8–15 Pm, thallus very large, up to 50 mm in diameter ........... 4 E Ascospores 12–25 Pm, thallus small to large (up to 30 mm in diameter) ... 5 4a Thallus thick (400–700 Pm when wet), lower surface wrinkled to reticulate, amphibious in the splash water zone or at temporarily inundated sites in high montane regions ................................................................ 'DUQROGLDQXP E Thallus thin (210–460 Pm when wet), lower surface smooth, terrestrial species from high-montane to hilly lowland areas, occasionally part of the riparian vegetation on rock outcrops bordering rivers or in shady springs or headwaters with constant water levels and not or very rarely experiencing complete inundation ....D. miniatum auct. (but see note for D. arnoldianum) 5a Lower surface of thallus wrinkled, reticulate (but variable in some collection), usually > 20 mm (13–30 mm) in diameter, irregularly shaped and thin (0.29–0.35 mm), restricted to subalpine and alpine regions ... 'ULYXORUXP E /RZHU VXUIDFH RI WKDOOXV VPRRWK RU ¿QHO\ JUDQXORVH UDUHO\ ZLWK D IHZ wrinkles or folds, 3–12 (20) mm in diameter, ± rounded and thick (0.29– 0.35 mm), from low elevations to mountain range areas ... 'PHLRSK\OOL]XP

50

.H\VWRWKHVSHFLHVDQGVSHFLHVSUR¿OHV

'HUPDWRFDUSRQDUQROGLDQXP 'HJHOLXV¿J 20) Synonyms: Dermatocarpon miniatum (Linnaeus) W. Mann 1825 var. miniatum non auct. 7KDOOXV upper surface greyish to brown, lower surface light-coloured; thallus monophyllous, lobes 6–50 mm in size, (200) 400–700 m thick when wet; with RUZLWKRXWSUXLQDZLWKHSLQHFUDOOD\HURIDLU¿OOHGK\SKDHORZHUVXUIDFHXVXDOO\ smooth to rugose or slightly reticulate; medulla showing QRUHDFWLRQZLWK0HOW]HU¶V Reagent; photobiont coccoid green alga (Diplosphaera ( chodatii according to 5HKDNRYD   YHUL¿HG E\ RZQ FXOWLYDWHG LVRODWHV  Perithecia: 190–390 (520) × 170–350 (470) m; ascospores (8.5) 13–17 × 4.0–7.0 m. Pycnidia: rare, pycnospores 4–5 × 1 m. (FRORJ\ amphibious, in cold and usually well lit streams on slightly acidic to basic, stable rocks, often associated with D. rivulorum but preferring less frequently inundated sites. 'LVWULEXWLRQ scattered but locally abundant at subalpine to alpine sites in the Alps and Tatra. 6LPLODUVSHFLHV Dermatocarpon rivulorum has smaller and much thinner lobes, the lower surface is dark brown and distinctly reticulate. Taxonomic note: Dermatocarpon arnoldianum LVUHJDUGHGDVFRQVSHFL¿FZLWK D. miniatum var. miniatum by Heiðmarsson (2001), but he also stated that the type material has unusually thick thalli compared to terrestrial forms of D. miniatum. Specimens of D. miniatum with smooth lower surface and thin thalli were found in the vicinity of rivers and springs at much lower altitudes than D. arnoldia-

Fig. 20. Dermatocarpon arnoldianum: ×1.

Dermatocarpon

51

num (F. Berger in lit., D. Teuber in litt.). In Central Europe the species complex still requires more detailed studies.

'HUPDWRFDUSRQOHSWRSK\OORGHV 1\ODQGHU =DKOEUXFNQHU¿J21) Basionym: Endocarpon leptophyllodes Nylander 1876. Synonyms: Dermatocarpon lorenzianum Anders 1922, Dermatocarpon phonolithicum Anders 1928. 7KDOOXV upper surface grey, lower surface brown to dark brown; lobes small, 1–4 (7) mm in diameter, 170–420 m when wet; foliose but occasionally of almost squamulose appearance, monophyllous, forming packed colonies with DUHRODWH DSSHDUDQFH “ VPRRWK PHGXOOD QRW UHDFWLQJ ZLWK 0HOW]HU¶V 5HDJHQW photobiont coccoid green alga. Perithecia: 200–290 × 160–260 m; ascospores 13–21 × 5–8 m. (FRORJ\ terrestrial to amphibious in Nordic countries; uncertain whether occurring in amphibious habitats in Central Europe. 'LVWULEXWLRQ very rare, in Central Europe only known from two localities in France (Haute Vienne) and Czech Republic (Bohemia, Kltþ), rare but widespread in Nordic countries. 6LPLODUVSHFLHV The species is distinguished from juvenile thalli of other species of Dermatocarpon by the minute lobes, the compact aspect of the colonies, DQGWKHQHJDWLYHUHDFWLRQRIWKHPHGXOODZLWK0HOW]HU¶V5HDJHQW Dermatocarpon deminuens is a Nordic species which has larger and more scattered thallus lobes.

Fig. 21. Dermatocarpon leptophyllodes: ×11.

52

.H\VWRWKHVSHFLHVDQGVSHFLHVSUR¿OHV

'HUPDWRFDUSRQOXULGXP(Dillenius ex Withering) J.R. Laundon 1984 ¿J159) Basionym: Lichen luridus Dillenius ex Withering 1776. Selected synonyms: Dermatocarpon aquaticum (Weiss) Zahlbruckner 1901, 'HUPDWRFDUSRQ ÀXYLDWLOH (Weber) Th. Fries 1860, Dermatocarpon weberi (Acharius) W. Mann 1825. 7KDOOXV upper surface brownish to greyish, bright green when wet, lower surface light to dark brown; multilobed, with several holdfasts, lobes 4–25 mm; diameter of entire thallus from a few centimetres up to 30 cm; thallus 150–440 m WKLFN ZKHQ ZHW HSLQHFUDO OD\HU ZLWK DLU ¿OOHG K\SKDH RU FRPSUHVVHG K\SKDH ORZHUVXUIDFHVPRRWKWRVOLJKWO\UHWLFXODWHPHGXOODWXUQLQJUHGZLWK0HOW]HU¶V Reagent; photobiont coccoid green alga (Diplosphaera ( chodatii according to Rehakova (1968) and YHUL¿Hd by own cultured isolates). Perithecia: 210– 490 × 150–520 m; ascospores: 10.5–20 × 5.5–8.5 m. Pycnidia: frequent, pycnospores 3.5–6 × 1 m. (FRORJ\ amphibious in the splash water zone of streams and rivers on stable siliceous rocks, often in moderate shade. Absent from strongly acidic water bodies (pH < 5) and possibly also sensitive to air-borne acidic emission. 'LVWULEXWLRQ widespread but scattered from lowland to subalpine regions. 6LPLODUVSHFLHV All other Central European species of Dermatocarpon growing in watercourses are monophyllous (caution with dense colonies!), and the meGXOODGRHVQRWVWDLQZLWK0HOW]HU¶V5HDJHQW

'HUPDWRFDUSRQPHLRSK\OOL]XP 9DLQLR¿J22) Synonyms: Dermatocarpon bachmannii var. inundatum Klement 1922, Placidiopsis meiophylliza (Vainio) Servít 1956. 7KDOOXV upper surface grey to brown, lower surface dark brown to black; foliose, monophyllous, lobes usually ± circular, but margins often incised and occasionally with small lobule-like outgrows; lobes 4–15 (31) mm in diameter, 290–350 m thick when wet; not pruinose; with epinecral layer of compressed K\SKDHPHGXOODQRWUHDFWLQJZLWK0HOW]HU¶V5HDJHQWSKRWRELRQW FRFFRLGJUHHQ alga. Perithecia: 200–420 × 120–420 m; ascospores 11–20.5 × 5–10.5 m. (FRORJ\ amphibious on acidic and basic substrata but restricted to siliceous rock types; best developed at slightly shaded sites; often associated with Dermatocarpon luridum but tolerating longer submersion periods and often found intermingled between D. luridum in the lowest part of the zone with best growth of D. luridum or below that zone. 'LVWULEXWLRQ from low altitudes to high-montane sites; in Central Europe usually at localities with (sub-)oceanic climate.

Dermatocarpon

53

Fig. 22. Dermatocarpon meiophyllizum: ×11. 6LPLODUVSHFLHV Dermatocarpon rivulorum is a subalpine to alpine species with larger lobes which are much thinner. Dermatocarpon luridum is multilobed, turns EULJKWJUHHQZKHQZHWDQGWKHPHGXOODWXUQVUHGZLWK0HOW]HU¶V 5HDJHQW Dermatocarpon leptophyllodes is a very rare species with grey surface growing in crowded, very dense colonies of small squamulose thalli. Dermatocarpon deminuens is a Nordic species which differs by its pruinose thallus surface and an HSLQHFUDOOD\HURIDLU¿OOHGK\SKDH

'HUPDWRFDUSRQULYXORUXP(Arnold) Dalla Torre & Sarnthein 1902 ¿J 160) Basionym: Endocarpon rivulorum Arnold 1874. Synonym: Placidiopsis rivulorum (Arnold) Servít 1956. 7KDOOXV upper surface brown to dark brown, little changing when moistened, lower surface brown to brown-black; monophyllous, but sometimes forming crowded colonies, lobes 10–30 (44) mm in diameter, thin, 100–320 (520) m when wet; attached by a single holdfast; epruinose; epinecral layer occasionally ZLWK DLU¿OOHG K\SKDH ORZHU VXUIDFH GLVWLQFWO\ UHWLFXODWH PHGXOOD QRW UHDFWLQJ ZLWK0HOW]HU¶V5HDJHQWSKRWRELRQWFRFFRLGJUHHQDOJDDiplosphaera  chodatii in own cultivated isolates). Perithecia: 210–480 × 180–412 m; ascospores 14–26 × 5.5–10.5 m. (FRORJ\ amphibious in usually well lit, cold streams and rivers on stable siliceous rocks, best developed at sites with small changes in water levels. Tolerates several months of permanent submersion (Keller & Scheidegger 1994). 'LVWULEXWLRQ subalpine to alpine sites in the Alps and Tatry; widely distributed in Nordic countries.

54

.H\VWRWKHVSHFLHVDQGVSHFLHVSUR¿OHV

6LPLODU VSHFLHV Dermatocarpon luridum is multilobed with several holdfasts, and the medulla WXUQVUHGLQ0HOW]HU¶V5HDJHQWDermatocarpon arnoldianum has thicker lobes and a light, lower surface which is usually smooth. All other amphibious species of the genus have much smaller lobes.

Endocarpon Hedwig 1789 1RQHRIWKH&HQWUDO(XURSHDQVSHFLHVLVVSHFL¿FDOO\DGDSWHGWR IUHVKZDWHUKDELWDWV but some species have repeatedly been recorded from occasionally inundated sites. While most species are more frequent at dry terrestrial sites, Endocarpon psorodeum is locally common in temporarily inundated sites. It can develop very large colonies RQDUWL¿FLDOVXEVWUDWDDWULYHUEDQNVDQGDORQJODUJHUULYHUV $WOHDVWLQVRPHUHJLRQV it is likely that it is more frequent here than in any non inundated terrestrial habitat. Important references: Breuss (1990), Clauzade & Roux (1985), Molitor & Diederich (1997), Servít (1955), Vainio (1921), Wirth (1995). Key to the species: 1a Squamules with rhizines ..................................... E. pusillum Hedwig 1789 E Squamules lacking rhizines ....................................................................... 2 2a Squamule margins not ascending, imbricate (like roof tiles), lower surface light to dark ........................................................................... E. psorodeum E Squamule margins ascending, not imbricate, lower surface dark in the squamule center and pale margins ..... E. adscendens (Anzi) Müller-Argau 1881

Endocarpon psorodeum (Nylander) Th. Fries ex Blomberg & Forssell ¿JV23–25)

Fig. 23. Endocarpon psorodeum: banks of the river Rhine; ×18.

Endocarpon

55

Fig. 24. Endocarpon psorodeum: banks of the river Rhine; ×19. Basionym: Verrucaria psorodeum Nylander 1866. Synonyms: Verrucaria pallida *var. [?] psorodea Nylander 1866, Dermatocarpon psorodeum (Nylander) Vainio 1883. Uncertain species: Endocarpon latzelianum Servít 1955. 7KDOOXV upper surface brown, grey-brown, green-brown; lower surface light to dark squamulose; forming dense mats of imbricate squamules (like roof tiles); lower surface lacking rhizines; photobionts coccoid green alga. Perithecia: immersed in the thallus; periphyses 20–45 m long, exciple 150–200 m in diameter; hymenial algae rounded; asci 2-spored; ascospores muriform, 27–51 × 12–22 m.

Fig. 25. Endocarpon psorodeum: banks of the river Rhine; ×10.

56

.H\VWRWKHVSHFLHVDQGVSHFLHVSUR¿OHV

(FRORJ\ WHUUHVWULDOVSHFLHVEXWORFDOO\IUHTXHQWRQQDWXUDOURFNVDQGROGDUWL¿cial bank reinforcements at large rivers in the upper amphibious zone (inundation only for a few days each year), forming a distinct band between Staurothele frustulenta populations and even more pronounced terrestrial species like Lecanora muralis (Thüs 2006). River bank populations usually at fully sun-exposed sites. Terrestrial populations also reported from shaded sites (Spribille & Thiel 2007, as E. latzelianum). 'LVWULEXWLRQFKDUDFWHULVWLFHOHPHQWRIWKHULSDULDQOLFKHQÀRUDLQVWUHWFKHVRIWKH rivers Rhine and Neckar and to be expected elsewhere at similar sites. Also obVHUYHGDWSHULRGLFDOO\LQXQGDWHGURFNRXWFURSVLQZDWHU¿OOHGGROLQDVLQ(VWRQLD .RVWLYHUH.DUVW¿HOG  6LPLODU VSHFLHV Endocarpon adscendens has squamules with ascending margins, always dark coloured lower surface in squamule centres bordered by a pale marginal zone. This species is reported from amphibious sites in Luxembourg (Molitor & Diederich 1997), Great Britain and Ireland (Purvis et al. 1992). Uncertain species: Endocarpon latzelianum has been reported to differ in slightly shorter ascospores and periphyses as well as in smaller cortex cells (Breuss 1990). The two ascospores in the asci differ considerably in size with the lower one being much larger than the upper one. For the lower (larger) ascospores 40–50 × 14–18 m are reported in E. psorodeum and 30–38 × 12–14 m in E. latzelianum. In riparian specimens from Germany, however, we observed an intermediate ascospore size. Furthermore, the length of periphyses and the size of cortex cells often did not correlate with the ascospore size as suggested above for distinction of E. latzelianum and E. psorodeum. Genetic studies are required to check whether E. latzelianum is merely a poorly developed morph of E. psorodeum or whether the intermediate freshwater specimens constitute a third, yet undescribed taxon.

(SKHEHFries 1825 Important references: Henssen (1963a), Jørgensen (2007b). Key to the species: 1a 7KDOOXVODUJH¿ODPHQWRXV¿ODPHQWVORRVHO\DWWDFKHGWRWKHVXEVWUDWHDVFL 8-spored, ascospores 10–20 × 3.5–7 m ...................................... (ODQDWD E Thallus very small, appearing crustose, composed of densely aggregated, HUHFW WR DVFHQGLQJ PLQXWH ¿ODPHQWV IRUPLQJ FXVKLRQV RU DUHROHV WLJKWO\ attached to the substrate, asci polysporous, ascospores smaller, 4–6 (7) × 3–5 m ................................................................................. (PXOWLVSRUD

Ephebe

57

(SKHEHODQDWD /LQQDHXV 9DLQLR¿J 26) Basionym: Lichen lanatus Linneus 1753. Synonyms: Conferva atrovirens Dillwyn 1809, Ephebe intricata Lamy 1878, Ephebe lapponica Nylander 1875, Ephebe pubescens auct., Ephebe pubescens var. complicata Vainio 1881, Ephebeia cantabria Nylander 1875, Ephebeia martindalei Crombie ex Nylander 1883, Lichen intricatus Ehrhart ex Schrader 1799, Usnea intricata Ehrhart ex Hoffmann 1796. 7KDOOXV¿ODPHQWRXVEODFNLVKWRGDUNJUHHQLVKRUEURZQLVKXVXDOO\JORVV\GHcumbent, threads combed in one direction to rarely rosette-shaped in compressed forms, 2–3 cm wide, fusing and covering larger areas; threads terete, irregularly furcate with the branches loose and slightly bent to intricate, branches 70–40 m thick, at the basis to 200 m thick, usually smooth, rarely rough and with short side branchlets; hyphae elongated and parallel towards the tips of the branches, forming a central strand in basal parts, attached by small gelatinous holdfasts; photobiont Stigonema. Apothecia: pycnoascocarps, nowadays rarely observed, ODWHUDOPPZLGHDW¿UVWDVVPDOOVZHOOLQJVRQWKHWKUHDGVODWHUGLVFVKDSHG thalline margin smooth, persisting; disc blackish; excip. prop. DW¿UVWYHU\WKLQ later becoming distinct, hyaline to slightly brownish; hymenium I+ blue; asci cylindrical to slightly clavate, thin-walled, I-, 8-spored, ascospores simple, hyaline, ellipsoid, 10–20 × 3.5–7 m. Pycnidia: lateral, immersed to slightly elevated; pycnospores cylindrical, 2.5–4.5 × 1–1.5 m. (FRORJ\ on inclined rock faces frequently moistened by seeping water in usually exposed or occasionally in relatively shaded situations; also on boulders along mountain streams wetted by water spray or sometimes inundated by running water. 'LVWULEXWLRQ in Central Europe montane to alpine, widespread in the northern hemisphere.

Fig. 26. Ephebe lanata: thallus thread-like; ×13.

58

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6LPLODU VSHFLHV Ephebe perspinulosa and E. hispidula occur in Fennoscandia and may also be found on boulders along mountain streams etc. in Central Europe and the Alps. Both species differ from Ephebe lanata in the rosette-shaped thalli with short branches which are rough due to the presence of numerous spine-like side branchlets. Furthermore, Ephebe perspinulosa has a loose paraplectenchymatous thallus anatomy in old branches whereas in old branches of E. lanata the hyphae form a central strand. Ephebe hispidula has elongated and roundish hyphal cells in old thallus branches and 8–16 ascospores per ascus whereas E. lanata and E. perspinulosa have 8-spored asci.

(SKHEHPXOWLVSRUD('DKO +HQVVHQ¿J27) Basionym: Spilonematopsis multispora E. Dahl 1950. 7KDOOXV¿ODPHQWRXVEODFNLVKEURZQLQGLYLGXDOWKDOOL–0.6 mm wide forming small cushions, 1–2 mm wide, resembling cushions of Stigonema, single cushions IXVLQJWRFRYHUODUJHUDUHDVWKUHDGVDW¿UVWDGQDWHVRRQHUHFWEDVDOO\GLYLGHGLQ several equally thick branches, 1 mm long, 15–20 (40) m thick, side branches few; hyphae thin or angulate, few, running through photobiont gelatinous sheath in right angles; photobiont Stigonema. Apothecia: pycnoascocarps, always terminal, very small and ± globose, 0.1 mm wide; thalline margin thin, persisting, minutely spiny due to small branches originating from the margin; excip. prop. absent; hymenium 70–100 m high, I+ blue; asci cylindrical to slightly clavate or obclavate, thin-walled, I-, (12) 16-spored, ascospores simple, hyaline, subglobose, 4–6 (7) × 3–5 m; paraphyses eventually distinctly septate, c. 1.5 m thick. Pycnidia: terminal, ± globose; pycnospores cylindrical, 3 × 1 m. (FRORJ\ on moist siliceous rock and occasionally or periodically inundated along alpine streamlets; at the high elevation locality in the central Alps accom-

Fig. 27. Ephebe multisporaWKDOOXVZLWKGHQVHO\DJJUHJDWHGHUHFW¿ODPHQWV ×24.

Gonohymenia

59

panied by Ephebe lanata and Rhizocarpon lavatum; in a Swedish locality c. 10 m off a rocky lake shore accompanied by Ephebe hispidula, Placynthium pannariellum and P. rosulans. 'LVWULEXWLRQ so far two records exist from SW Greenland and two from northern Sweden. The species is reported here from eastern Tyrol suggesting an arctic-alpine to high montane distribution. 6LPLODU VSHFLHV Ephebe multispora resembles compact cushions of the freeOLYLQJ¿ODPHQWRXVF\DQREDFWHULXP Stigonema. The species may also be mistaken for Porocyphus coccodes or forms of P. rehmicus due to the minutely fruticose thallus forming dense cushions. Otherwise, E. multispora is unlikely to be conIXVHGZLWKDQ\RWKHUVSHFLHVRIWKHJHQXV,WLVUHDGLO\LGHQWL¿HGE\WKHSRO\VSRrous asci containing very small, subglobose spores and the small cushions with terminal, globose apothecia.

Gonohymenia J. Steiner 1902 Notes: Moreno & Egea (1992) suggested to synonymize Gonohymenia sensu Henssen under Lichinella Nylander and thereby created a large assemblage of VXSHU¿FLDOO\UDWKHUGLVVLPLODUVSHFLHVXQLWHGE\VLPLODULWLHVLQWKHWKDOOLQRFDUSRXV fruit bodies and ascus type. Gonohymenia was retained as a separate genus by Wirth (1994, 1995), Wirth et al. (1996), Scholz (2000), Hafellner & Türk (2001) and Santesson et al. (2004) whereas Nimis (1993) followed the concept of Moreno & Egea (1992) though with some hesitation. Recently, Jørgensen (2007b) reestablished the genus Thallinocarpon E. Dahl to accommodate the larger species of Gonohymenia (i.e. G. iodopulchra-group) which are said to differ in the fruit bodies from the type species of the genus, G. algerica J. Steiner. Indeed, the relationship and delimitation of the genera Lichinella, Gonohymenia and Thallinocarpon LVDGLI¿FXOWPDWWHUDQGDQRYHOWUHDWPHQWEDVHGRQDWKRURXJK UHYLVLRQLVQHHGHG Therefore, and for practical reasons Schultz (2005) adopted the concept of Moreno & Egea (1992) and treated all species present in SW North America under the single genus Lichinella. Here, the traditional concept of Gonohymenia (Henssen µ¶ LVXVHGVLQFH a possible placement in Thallinocarpon needs support from a novel revision of the concepts of the genera involved. Important references: +HQVVHQ µ¶  -¡UJHQVHQ E  0RUHQR Egea (1992), Schultz (2005).

Gonohymenia heppii0OOHU$UJRYLHQVLV +HQVVHQ¿J 28) Basionym: Omphalaria heppii Müller Argoviensis 1862. Synonym: Thyrea heppii (Müller Argoviensis) Lettau 1912. 7KDOOXV blackish to greyish pruinose, forming medium-sized rosettes or cushions 8–15 mm wide, fastened to the substrate by central umbilicus, rosettes soon becoming distinctly lobulate; lobes loosely adpressed to ascending, surface

60

.H\VWRWKHVSHFLHVDQGVSHFLHVSUR¿OHV

Fig. 28. Gonohymenia heppiiWKDOOXVIUDJPHQWZLWKÀDWHQHGGHHSO\EUDQFKHG ascending lobes; ×10. slightly rough due to the presence of small globose, isidia-like outgrowths ±—P DFURVV OREHV ±PP ZLGH QDUURZHG WRZDUGV WKH EDVH ÀDW WR convex, tips rounded with down curved margin; lobes ecorticate, 110–225 m thick, heteromerous, centrally with loose hyphal strand lacking photobiont cells 50–175 m thick, hyphal arrangement reticulate at the lobe periphery, 25–35 m thick, photobiont a single-celled cyanobacterium. Apothecia: thallinocarps, marginal to submarginal, usually visible as ± roundish to irregularly shaped, slightly ÀDWWRFRQYH[VZHOOLQJDWWKHOREHWLSVGLVFURXJKGRWWHG excip. prop. absent; asci 16–32-spored, ascospores simple, hyaline, broad ellipsoid, small, 5–9 × 3–5 m. Pycnidia: not observed. (FRORJ\ on sporadically submerged or sprayed calcareous rocks along river banks. 'LVWULEXWLRQ known from historical records in the river Arve close to Mornex and close to Boverniche at the river Dranse in the western Alps; also reported from the Llanos de la Larri, Huesca in the Spanish Pyrenees (Pereira 1992). Notes: According to Henssen & Jørgensen (1990) this taxon belongs to the Gonohymenia iodopulchra-group the species of which have recently suggested to be accomodated in Thallinocarpon E. Dahl (Jørgensen 2007b). Gonohymenia heppii is distinguished from G. iodopulchra by the somewhat larger, thinner lobes, perKDSVDUHVSRQVHWRWKHLQÀXHQFHRIUXQQLQJZDWHU

Gyalidea

61

*\DOLGHD /HWWDXH[9Č]GD Important references:3XUYLVHWDO 9Č]GD :LUWK  Key to the species: 1a Ascospores muriform ................................................................... *IULW]HL E Ascospores only with transverse septa ...................................................... 2 2a Epithecium colourless, exciple pale pink- or yellowish brown to creamFRORXUHGDSRWKHFLDOGLVFÀHVKFRORXUHG ..................................... *URVHROD E (SLWKHFLXPSDOHJUHHQEURZQH[FLSOHGDUNEURZQDSRWKHFLDOGLVFQRWÀHVK coloured .................................................................................................... 3 3a Apothecial disc obscure when dry, violet-grey when wet, 0.2–0.4 mm in diameter, ascospores 1 to rarely 3-septate .............................. G. diaphana E Apothecial disc pale orange to brown when dry, translucent when wet, 0.4–1.0 mm in diameter, ascospores 1–3-septate .............. *K\DOLQHVFHQV

*\DOLGHDGLDSKDQD .|UEHUH[1\ODQGHU 9Č]GD¿J29) Basionym: Biatora diaphana Körber ex Nylander 1856. Synonyms: Lecidea diaphana (Körber ex Nylander) Nylander 1857, Biatorina diaphana (Körber ex Nylander) Körber 1860, Catillaria diaphana (Körber ex Nylander) Lettau 1912. 7KDOOXV blackish; thin, cracked, ± gelatinous when wet; photobiont yellowish orange coloured green alga (Trentepohlia). Apothecia:VXSHU¿FLDO±—P LQGLDPHWHUGLVFÀDWGDUNEURZQWREODFNZKHQGU\YLROHWJUH\ZKHQZHWH[FLple thin, dark brown to black; hymenium 70–80 m high, colourless; ascospores 1 (3)-septate 8–15 × 5–6 m. (FRORJ\ amphibious on siliceous rocks in streams and rivers but also terrestrial in shady places (Palice et al. 2006).

Fig. 29. Gyalidea diaphana: ×32.

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.H\VWRWKHVSHFLHVDQGVSHFLHVSUR¿OHV

'LVWULEXWLRQ amphibions populations very rare in upland areas, known from areas along the Polish-Czech border and from Austria. 6LPLODUVSHFLHV Gyalidea fritzei has muriform ascospores.

*\DOLGHDIULW]HL6WHLQ 9Č]GD¿J149) Basionym: Gyalecta fritzei Stein in Cohn 1879. Synonym: Gyalecta lecideopsis var. fritzei 6WHLQ 9Č]GD 7KDOOXV rusty red to whitish; thin and often indistinct, smooth to cracked; photobiont yellowish-orange coloured green alga (Trentepohlia). Apothecia: sessile, perithecia-like, with small, pore-like disc when young, becoming deeply concave with age, 200–600 m in diameter, 100–200 m high, disc pale orange to dark brown; exciple brown; epithecium pale green-brown; hymenium 80–160 m high, pale green-brown in upper part; hypothecium pale brown; asci 76–120 × 25–45 m, 6–8-spored; ascospores muriform, 18–43 × 10–21 m. (FRORJ\ on sporadically wet and slightly basic, siliceous rocks. 'LVWULEXWLRQ rare species in upland areas. 6LPLODUVSHFLHV Other amphibious species off Gyalidea have 1–3 septate ascospores. Gyalidea lecideopsis is similar to G. fritzei but differs from the latter in the darker brown-green upper hymenium and the occurrence on dry calcareous rocks. Gyalidea roseola has a pale pink to yellowish brown or cream-coloured exciple and a colourless epithecium.

*\DOLGHDK\DOLQHVFHQV1\ODQGHU 9Č]GD¿J150) Basionym: Lecidea hyalinescens Nylander 1856. Synonyms: Bacidia hyalinescens (Nylander) Zahlbruckner 1927, Bilimbia hyalinescens (Nylander) Boistel 1926, Gyalecta hyalinescens 1\ODQGHU 9Č]GD 7KDOOXV grey to brown-grey; thin, smooth to cracked; photobiont yellowishorange coloured green alga (Trentepohlia). Apothecia:LPPHUVHGWRVXSHU¿FLDO XSWRPPLQGLDPHWHUGLVFÀDWWRFRQFDYHRUDQJHWREURZQZLWKSHUVLVWLQJ pale grey to brownish margin; hymenium 70–120 m high, colourless; asci 70–80 × 10–14 m; ascospores 3-septate, 12–24 × 5–7 m. (FRORJ\ amphibious on siliceous rocks at shaded sites. 'LVWULEXWLRQ very rare, extincted in Germany, old records from low mountainous localities with oceanic climate (Black Forest, Geroldsau). 6LPLODU VSHFLHV Ionaspis lacustris has a similar external appearance, but the exciple is indistinct, the ascospores are simple and the photobiont is a purely green-coloured Trebouxia (Asterochloris ( ). Gyalidea diaphana has a blackish thallus and most of the ascospores are uniseptate.

Hydropunctaria

63

*\DOLGHDURVHROD $UQROG 9Č]GD Basionym: Gyalecta roseola Arnold 1873. 7KDOOXV white; thin, smooth to uneven; photobiont yellowish-orange coloured green alga (Trentepohlia). Apothecia:VXSHU¿FLDOXSWR—PLQGLDPHWHUGLVF FRQFDYHSLQNLVKWRÀHVKFRORXUHGHSLWKHFLXPFRORXUOHVVWUXH H[FLSOHGLVWLQFW elevated, pale pink- or yellowish brown to cream-coloured; hymenium 90–120 m high; asci 70–100 × 15–25 m; ascospores muriform, 20–36 × 8–14 m. (FRORJ\ shaded, humid to damp places, occasionally in the splash water zone of streams or waterfalls. 'LVWULEXWLRQ a very rare species, only known from Austria and an uncertain record from Italy. 6LPLODUVSHFLHV other amphibious species of Gyalidea have a coloured epithecium, differently coloured apothecial disc, and a dark brown exciple.

Hydropunctaria Keller, Gueidan & Thüs 2008 Notes: This genus was recently segregated from the polyphyletic genus Verrucaria after multigene studies revealed a rather isolated phylogenetic position of the species of the new genus in the order Verrucariales (Gueidan et al. 2007). Hydropunctaria is characterized by the combination of small to medium-sized ascospores, a paraplectenchymatous and subgelatinous thallus which develops black columns or dots under certain ecological conditions and a distinctly coloured cortex when growing under well lit conditions. The black dots or columns make the genus very distinctive but morphs exist lacking this structure. For these forms a cross-reference was included in the Verrucaria key. Important references: Gueidan et al. (2007, 2008), Orange (2004), Thüs (2002), 9Č]GD  Key to the species: 1a Involucrellum apical or somewhat reaching further down (but rarely reaching the thallus base), if streching below, then soon laterally extending into the thallus and leaving an unprotected base of the exciple¿JV± , ascospores ± ellipsoid, occasionally almost circular, often one pole more acute than the other, 7.5–10.3 (16) Pm ...................................... +UKHLWKURSKLOD E Involucrellum reaching down to the thallus base, attached to the exciple or to the base of the exciple, distinction between involucrellum and exciple often impossible (¿JV ± , ascospores 9.5–16 (21.5) m, often with truncated poles ............................................................................ +VFDEUD

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Fig. 30

Fig. 32

Fig. 31

Fig. 33

Plate 3. Fig. 30–33. Shape of LQYROXFUHOOXP in Hydropunctaria. Fig. 30–31. H. rheitrophila. Fig. 32–33. H. scabra.

Hydropunctaria rheithrophila =VFKDFNH .HOOHU*XHLGDQ 7KV ¿JV 30–31 34–36) %DVLRQ\P Verrucaria rheithrophila =VFKDFNH6\QRQ\PVVerrucaria kernstockii =VFKDFNH Verrucaria minutipuncta(ULFKVHQVerrucaria minutipuncta Ilaeviorr (ULFKVHQ Verrucaria kernstockiiIminutipuncta (ErichVHQ 6HUYtW Verrucaria scotinodes =VFKDFNHVerrucaria sagedioides 6HUYtW 8QFHUWDLQVSHFLHVVerrucaria cinereolutescens =VFKDFNHVerrucaria rechingerii6HUYtW Verrucaria refugii 6HUYtWVerrucaria maurula 0OOHU $UJRYLHQVLVVerrucaria wolferi=VFKDFNH Thallus: ROLYH JUHHQ  \HOORZLVKJUHHQ \HOORZLVKEURZQ WR RUDQJHEURZQ RUDQJHRU\HOORZLVKEURZQWLQJHLQWHQVL¿HGE\VWRUDJH WKLQWRWKLFN±—P PXFKGHSHQGLQJRQOLJKWFRQGLWLRQVXVXDOO\FRQWLQXRXVEXWODUJHFUDFNVRFFXU DIWHUGURXJKWGDPDJH SURWKDOOXVDEVHQWDWIUHHWKDOOXVPDUJLQVEXWRFFDVLRQDOO\ GLVWLQFWEODFNOLQHVRUULGJHVEHWZHHQQHLJKERXULQJWKDOOLWKDOOXVDQDWRP\SDUDSOHFWHQFK\PDWRXVDQGVXEJHODWLQRXVPDUJLQVRIWKLQWKDOOLWUDQVSDUHQWZKHQZHW FDUERQLVHGGRWVRUFROXPQVLQWKHWKDOOXVXVXDOO\SUHVHQWEXW PXFKYDULDWLRQPD\ H[LVWLQDVLQJOHWKDOOXVDQGRFFDVLRQDOO\WKHGRWVRUFROXPQV DUHFRPSOHWHO\DEVHQWEODFNEDVDOOD\HUJHQHUDOO\DEVHQWRUSUHVHQWRQO\LQVPDOODUHDVRIWKHWKDOOXVFRUWH[\HOORZLVKEURZQWREODFNLVKEURZQRUDEVHQWSKRWRELRQW 'LODEL¿OXP or Heteroccoccus caespitosusRFFDVLRQDOO\HORQJDWHGFHOOVSUHVHQWDOJDOFHOOVLQ WKHWKDOOXV±—PLQGLDPHWHURIWHQDUUDQJHPHQWLQYHUWLFDO FROXPQVPerithecia: LPPHUVHGWRSURMHFWLQJGHSHQGLQJRQWKHKHLJKWRIWKHWKDOOXVSURMHFWLQJ SDUWVQDNHGH[FLSOH±—PLQGLDPHWHULQYROXFUHOOXPYHU\ YDULDEOHIURP DSLFDODQGOLWWOHGHYHORSHGWRODWHUDOO\VSUHDGLQJODWHUDOH[WHQVLRQ±—P EDVDOSDUWRIH[FLSOHJHQHUDOO\GLVWLQFWO\VHSDUDWHGIURPWKHLQYROXFUHOOXPOHDY-

Hydropunctaria

65

Fig. 34. Hydropunctaria rheitrophila: thallus surrounded by pale prothallus; ×16. ing a free space between the two structures; surface of involucrellum often “rough” or with small projections; asci 8-spored; ascospores 7.5–15 × 4.7–8.2 m, unicellular, not halonate. (FRORJ\ submerged or frequently wetted in the lower splash water zone on siliceous or calcareous rocks and pebbles. Characteristic component of permanently submerged vegetation. Typically with Verrucaria aquatilis and/or the red alga Hildenbrandia rivularis. Tolerant to a wide range of illumination and moderate eutrophic conditions in fast running streams but sensitive to silting, tuffa deposiWLRQRUFRPSHWLWLRQE\PDWWIRUPLQJF\DQREDFWHULD/HVVVHQVLWLYHWRDFLGL¿FDWLRQ than the otherwise ecologically similar V. aquatilis. 'LVWULEXWLRQ From sea-level to alpine areas. Absent from heavily eutrophicated streams but recovering in much of its former distribution area in Central Europe after water quality has increased due to improved sewage treatment. 6LPLODUVSHFLHV In Hydropunctaria scabra the involucrellum is generally reaching down to the thallus base with its lower part being closely adpressed to the exciple leaving no free space between the two structures. The perithecia are covered by a thallus mantle leaving only the apical part around the ostiole naked. A black basal layer is usually present and often massively developed, the cortex usually has an olive-greenish tinge and the ascospores are slightly larger. Verrucaria aquatilis has smaller and more rounded ascospores, a constantly thin, often blackish (occasionally brown or greenish) thallus and an involucrellum with smooth surface, reaching the lower exciple half or even the thallus base. Uncertain species: Several taxa have been described which differ from Hydropunctaria rheitrophila only in subtle details and most of these species are only known from their type localities. Additional material is needed to evaluate the character variability, and molecular studies may eventually help to achieve a ro-

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bust concept of the taxonomic status of the critical taxa involved. Verrucaria cinereolutescens is a yellowish species with spreading black involucrellum and ascospores 9–12 × 5–6.5 m in size. It lacks, however, any black dots in the thallus. Verrucaria refugii is very similar to V. cinereolutescens in its yellowishgreenish, thick thallus lacking black dots, completely immersed perithecia and a weakly developed apical involucrellum. On the other hand, V. refugii has small, globose ascospores (7–8 m in diameter) as present in V. aquatilis. It is only known from the type locality (streams close to lake “Steirer See” near the village Klachau, Austria at 1500 m altitude). Verrucaria rechingerii has small, but more elongated ascospores (7–10 × 5–7 m), a thick thallus and immersed perithecia and lacks black dots in the thallus like V. cinereolutescens. The structure of the involucrellum in authentic material kept at the Natural History Museum Vienna (Krypt. Exscicc. 1851) varies from apical to reaching the thallus base. A locally developed “black basal layer” (also depicted in Servít 1950) appears to consist of remnants of dead thalli which have been overgrown by younger thalli of Verrucaria spp. Verrucaria wolferi has a thallus up to 100 m thick and immersed perithecia with only the uppermost part of the involucrellum being naked. The involucrellum is closely adpressed to the exciple and stretches downwards almost to the base of the exciple. The perithecia are rather large and up to 350 m in diameter. The cortex consists of a coloured layer which is overlain by a ± distinctly separated colourless layer of dead hyphae. The thallus colour is bronze, and calcareous structures in the thallus are absent. It is only known from the type locality (Switzerland, Davos, inundated on calcareous rocks in the stream “Dorfbach” at 2000 m altitude). Verrucaria maurula is a poorly known taxon with black thallus, completely immersed perithecia and very small ascospores (6–8 × 4–6 m) similar to those of V. aquatilis. The species is only known from the type locality (Switzerland, on calcareous rocks in the Voirons Mountains close to Geneva). Authentic material could not be traced so far and no detailed information on the structure of the involucrellum is available rendering a revision of this taxon impossible at the moment.

Fig. 35–36. Hydropunctaria rheitrophila: black dotted thallus, rather indistinct in GU\VWDWH¿J35 FRQVSLFXRXVLQZHWVWDWH¿J36); ×43.

Hydropunktaria

67

+\GURSXQFWDULDVFDEUD 9Č]GD .HOOHU*XHLGDQ 7KV ¿JV 32–33, 154) Basionym: Verrucaria scabra9Č]GD 7KDOOXV black to olive; thick (110–140 m), surface often roughened by the tips of black columns, but also smooth when columns end below the cortex; thallus cortex olive-brown to olive-blackish or yellowish-brown; prothallus absent; thallus anatomy paraplectenchymatous and subgelatinous, only margins of young thalli occasionally becoming transparent when wet due to the presence of a black basal layer, basal layer much varying in size, rarely absent, often with black columns which may perforate the cortex; photobiont 'LODEL¿OXP, diameter of photobiont cells in the thallus 5.7–8 m, often arranged in vertical columns. Perithecia: completely immersed or immersed in ± projecting thallus warts; exciple 90–177 m in diameter; involucrellum reaching down to the thallus base and laterally spreading into the thallus, adjacent involucrella usually fusing or connected to the black basal layer, basal parts of involucrellum closely adpressed to WKHH[FLSOHOHDYLQJQRIUHHVSDFHRUUDUHO\ZLWKDWUDQVSDUHQWDUHDÀDQNLQJWKH exciple; asci 8-spored; ascospores unicellular, 9.7–18.5 × 5.2–9.4 m, not halonate. (FRORJ\ amphibious in the splash water zone or temporarily submerged on stable siliceous rocks, often in rapids or waterfalls. Tolerant to a wide range of illumination. 'LVWULEXWLRQ very rare but locally abundant in mountain regions; so far recorded from Slovakia (Tatra), Austria (Niedere Tauern, Ötztal), Switzerland, France (Massif Central) and Germany (Black Forest); also known from Iceland. 6LPLODU VSHFLHV Hydropunctaria rheitrophila lacks a continuous black basal layer, has smaller ascospores (7.5–15 m), and an involucrellum which is apical or, if reaching further below, never attached to the basal part of the exciple. The perithecia are either immersed or semi-immersed, but the protruding parts are always naked. Verrucaria madida is a very rare oceanic species with 4-spored asci, smaller ascospores (9–15 × 5.5–7.5 m) and a thinner greenish thallus without any black basal layer, carbonised black dots or columns in the thallus. Verrucaria pachyderma is an alpine species with larger perithecia, a much larger lenght/width-ratio of the ascospores (15–23.5 × 6–9 m), and the thallus never contains black dots or columns. Uncertain species of Hydropunctaria: see Verrucaria.

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+\PHQHOLD Krempelhuber 1852 Important references: Lutzoni & Brodo (1995), Poelt (1969). Key to the species: 1a On frequently inundated siliceous substrata, ascospores 6–12 (14) Pm, hyPHQLXP±—PKLJKWKDOOXVVPRRWKRU¿QHO\ULPRVHDSRWKHFLDOPDUJLQ distinctly raised ................................................................... H. cyanocarpa E Predominantly terrestrial species on calcareous substrata ......................... 2 2a Apothecia pale pink or pale brown ... H. epulotica (Acharius) Lutzoni 1995 E Apothecia dark bluish or blackish ............................................................. 3 3a Thallus r endolithic, ascospores 10–20 Pm long ........................................ ............................................. H. melanocarpa (Krempelhuber) Arnold 1887 E Thallus epilithic, rimose to areolate or powdery, ascospores 7–12 Pm long .......................................... H. heteromorpha (Krempelhuber) Lutzoni 1995

+\PHQHOLDF\DQRFDUSD$Q]L /XW]RQL¿J 148) Basionym: Aspicilia cyanocarpa Anzi 1862. Synonyms: Ionaspis cyanocarpa (Anzi) Th. Fries 1871, Lecanora cyanocarpa (Anzi) Nylander 1882. 7KDOOXV ZKLWLVKJUH\ WR SLQNLVK WKLQ WR WKLFN FRQWLQXRXV WR ¿QHO\ ULPRVH photobiont yellowish-orange to yellowish-greenish (Trentepohlia). Apothecia: semi-immersed (disc immersed but with distinct, ± prominent black margin), 300–500 m in diameter, disc blue-black; hymenium 85–115 m high, upper part blue-green, N+ red; ascospores unicellular, 6–14 × 5–8 m. (FRORJ\ amphibious on siliceous rocks. 'LVWULEXWLRQ a rare species in alpine to subalpine watercourses; also found in Scandinavia and across arctic regions. 6LPLODU VSHFLHV Ionaspis suaveolens has a lower hymenium (50–60 m), the exciple is not prominent and the upper hymenium is N-.

Ionaspis Th. Fries 1871 Notes: The key includes some species of Hymenelia which are very similar to Ionaspis in external appearance. The only species of Hymenelia from freshwater habitats in Central Europe is more similar to Ionaspis spp. than to any other lichen from such localities. Important references: Gilbert (2000), Lutzoni & Brodo (1994, 1995), Poelt (1969), Purvis et al. (1992), Wirth (1995). Key to the species: 1a Photobiont cells green (Trebouxia/Asterochloris / ) ..................... ,ODFXVWULV E Photobiont cells orange to (greenish-) yellow (Trentepohlia) ................... 2

Ionaspis

69

2a Apothecia pale pink or brown, upper part of hymenium colourless or pale brown to olive above, K- .......................................................................... 3 E Apothecia dark blue to black, upper part of hymenium blue-green, K+ violet or K- .......................................................................................................... 4 3a Hymenium 80–140 Pm high, predominantly terrestrial species, on calcareous substrata ....................... Hymenelia epulotica (Acharius) Lutzoni 1995 E Hymenium 60–80 PP KLJK ÀXYLDO PHVLF WR VXEPHUJH ]RQHV RQ VLOLFHRXV substrata ......................................................................................... I. odora 4a On siliceous substrata, apothecial margin not raised, upper hymenium N................................................................................................ ,VXDYHROHQV E On siliceous or calcareous substrata, apothecial margin raised or not, upper hymenium N+ red to mauve ...................................................................... 5 5a Ascospores 6–12 (14) PPWKDOOXVVPRRWKRU¿QHO\ULPRVHDSRWKHFLDOPDUgin distinctly raised, on frequently inundated siliceous substrata ................ ............................................................................... +\PHQHOLDF\DQRFDUSD E Ascospores (10) 15–17 (20) Pm, on calcareous substrata, predominantly terrestrial species ........................................................................................... 6 6a Thallus r endolithic .... Hymenelia melanocarpa (Krempelhuber) Arnold 1887 E Thallus epilithic, rimose to areolate or powdery .......................................... ............................. Hymenelia heteromorpha (Krempelhuber) Lutzoni 1995

,RQDVSLVODFXVWULV :LWKHULQJ /XW]RQL¿J 145) Basionym: Lichen lacustris Withering 1796. (Selected) Synonyms: Aspicilia lacustris (Withering) Th. Fries 1861, Lecanora gibbosa f. lacustris (Withering) Leighton 1871, Hymenelia lacustris (Withering) M. Choisy 1949, Lecanora lacustris (Withering) Nylander 1861. 7KDOOXV whitish, yellowish-ochre to reddish-orange or greenish; up to 400 m thick, continuous to rimose; photobiont green with ± spherical cells (Trebouxia, AsterochlorisJURXSYHUL¿HGE\FXOWLYDWHGLVRODWHVDQG,76VHTXHQFHV  Apothecia: immersed, concave, 150–600 m in diameter, disc pinkish, white to orange; margin (exciple) indistinct or slightly raised; hymenium 85–105 m high; ascospores unicellular, 13–24 × 6–11 m. (FRORJ\ amphibious on hard and stable siliceous substrata in cool, clear streams and rivers, from moderately shady to fully sun exposed sites; in neutral to moderately acidic waters (pH > 4.5–5). 'LVWULEXWLRQ widespread from lowland to high-montane regions, but absent or very rare in areas with brittle bedrock, strong acidic impacts (or acidic impact history) and high eutrophication; generally more frequent in cooler upland areas. 6LPLODUVSHFLHV Ionaspis lacustris is the only freshwater species of the genus in Central Europe with Trebouxia (Asterochloris ( )-photobionts (pure green colour) instead of yellowish-orange-green Trentepohlia cells. The photobiont colour changes after longer storage in the herbarium, but Trebouxia (Asterochloris ( ) cells are usually smaller (< 15 m) than those of Trentepohlia (10–18 m) and have thinner walls.

70

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Ionaspis odora $FKDULXVH[6FKDHUHU 7K)ULHV¿JV146, 169) Basionym: Gyalecta odora Acharius ex Schaerer 1826. Synonyms: Aspicilia odora (Acharius ex Schaerer) A. Massalongo 1852, Lecanora odora (Acharius ex Schaerer) Nylander 1857, Patellaria odora (Acharius ex Schaerer) Trevisan 1853, Pinacisca odora (Acharius ex Schaerer) Trevisan 1855, Ionaspis suaveolens f. odora (Acharius ex Schaerer) Arnold 1893. 7KDOOXV whitish, pinkish to olive-grey, yellowish-brown; rimose to areolate, thin to thick, K± violet; photobiont yellowish-orange to yellowish-green (TrentepohliaVSYHUL¿HGE\FXOWXUHGLVRODWHV¿J). Apothecia: immersed, 100–400 m in dia-meter, pale pink, yellowish-brown, orange-brown to olive-blackish; hymenium 50–80 m high, upper part colourless, yellowish-olive to blackish-olive, K+ violet, N-; ascospores unicellular, 8–14 × 5–6 m. Pycnidia: 50–80 m in diameter, with greenish, K+ violet wall. (FRORJ\ amphibious on hard and stable siliceous rocks in well lit streams. 'LVWULEXWLRQ alpine to high-montane regions; very rare outside the Alps and locally much declining due to forestation with spruce thickets in old meadow valleys. 6LPLODUVSHFLHVno other Central European Ionaspis has a K+ violet upper hymenium.

,RQDVSLVVXDYHROHQV (Fries) Th. Fries 1871 nom. cons. ¿JV 147, 168) Basionym: Gyalecta suaveolens Fries 1825 nom. cons. Synonyms: Ionaspis suaveolens (Schaerer) Th. Fries ex Stein 1874, Urceolaria suaveolens (Fries) Acharius 1826, Patellaria suaveolens (Fries) Trevisan 1852, Aspicilia chrysophana Körber 1855, Ionaspis chrysophana (Körber) Th. Fries 1871, Lecanora chrysophana (Körber) Nylander ex Stizenberger 1882. 7KDOOXV dark green, olive-brown, yellowish-brown or pinkish; thick and crackedareolate to thin and almost smooth and continuous; photobiont with large orange coloured cells (Trentepohlia sp.¿JYHUL¿HGE\FXOWXUHGLVRODWHV  Apothecia: immersed, 200–500 m in diameter, disc blackish; hymenium 50–60 m high, upper part blue-green, K-, N-, or N+ reddish, soon fading; ascospores unicellular, 6–11 × 5–7 m. Pycnidia: unknown, absent in all Central European specimens examined so far. (FRORJ\ splash zone of alpine and high-montane watercourses on stable siliceous substrates; also terrestrial on moist rocks. 'LVWULEXWLRQ Alps, Tatra, Sudetes, highest elevations of Black Forest and Vosgues. 6LPLODUVSHFLHV Hymenelia cyanocarpa is similar but differs in raised apothecial margins and the hymenial pigment, which gives a reddish colour in N.

Koerberiella

71

.RHUEHULHOODStein 1879 Important reference: Rambold et al. (1990).

.RHUEHULHOODZLPPHULDQD .|UEHU 6WHLQ¿J37) Basionym: Zeora wimmeriana Körber 1853. Synonyms: Lecanora wimmeriana (Körber) Poetsch 1872, Lecanora creatina Norman ex Th. Fries 1871, Lecidea creatina (Norman ex Th. Fries) Stizenberger 1876, Lecanora leucophyma Leighton 1871, Aspicilia leucophyma (Leighton) Hue 1912, Lecanora acceptanda Nylander 1879, Aspicilia acceptanda (Nylander) Arnold ex Hue 1912, Pertusaria littoralis Vainio 1881, Lecanora acceptanda var. littoralis (Vainio) Vainio 1883, Aspicilia littoralis (Vainio) Hue 1912, Lecanora littoralis (Vainio) Zahlbruckner 1928, Aspicilia leucophyma var. littoralis (Vainio) Räsänen 1939, Lecanorella josiae Frey 1924, Perspicinora leucophyma (Leighton) Riedl 1990. 7KDOOXV greyish, pale cream-coloured to pinkish-brown, light brownish; thallus of dispersed large granules or rimose-areolate, 200–1000 m thick, areoles/granules 300–100 m in diameter, usually isidiate, isidia unbranched, cylindrical to clavate, single or a few per areole; thallus C-, KC-, but gyrophoric acid detected LQ VPDOO TXDQWLWLHV E\ WOF DOVR DQ XQLGHQWL¿HG VXEVWDQFH LQ ODUJHU TXDQWLWLHV (RF-value 3–5 in solvent A); photobiont a coccoid green alga, spherical cells of 7–16 m in diameter in the thallus (Trebouxia impressa, YHUL¿HGE\LVRODWHGFXOtivates and ITS-sequences). Apothecia: rare, usually in the areas without true isidia, but developing from isidia-like protuberances, 0.6–1.4 mm in diameter, margin well developed and persistent; hymenium 110–140 m high; epihymenium brownish; asci 80–115 × 20–25 m; ascospores unicellular, 17–30 × 9–15 m. Pycnidia: immersed in the tips of isidia; pycnospores 3.5–5 × 1–1.5 m.

Fig. 37. Koerberiella wimmeriana: ×11.

72

.H\VWRWKHVSHFLHVDQGVSHFLHVSUR¿OHV

(FRORJ\ best developed in seepage tracks and small streams with only short periods of inundation, but also amphibious in the splash zone at or in larger and permanent watercourses; from moderately shady to fully sun-exposed locations on siliceous substrata. 'LVWULEXWLRQ high-montane to alpine localities in the Alps, very rare outside the Alps, e.g. in the Auvergne (France), Black-Forest (Germany), Krkonoše/Karkonosze (Czech Rep./Poland), Tatra (Poland/Slovakia). Also present in Scandinavia, Great Britain and generally widespread in the arctic to subarctic zones in the northern hemisphere. 6LPLODUVSHFLHV 6XSHU¿FLDOO\VLPLODULVLGLDWHVSHFLHVKDYHPRUHLVLGLDSHUDUHROH and contain different secondary metabolites (e.g. fatty acids in Pertusaria isidioides, stictic acid in Aspicilia corallophora, A. glomerulans, norstictic acid in A. mastrucata and/or the thallus reacts K+/KC+ red (Rimularia gibbosa, R. badioatra) or the isidia are branched. Although some of the species mentioned before are hygrophytic and may be found in seepage tracks none of them is regularly found at or in streams and rivers.

Lecidea Acharius 1803 Notes: None of the Central European species of Lecidea is restricted to freshwater habitats. Although several species may occasionally occur in the splash water zone of upland watercourses, most of them are much more frequent above the inundation zone. Furthermore, the small-scale distribution of Lecidea species at a riverbank usually shows no upper distribution limit, indicating that the species are bound to terrestrial lichen communities. Consequently, these species are not treated here in detail. An exception is Lecidea ahlesii which has one of its principal habitats in the vicinity of shady streams often being much more frequent in the immediate vicinity of watercourses than in neighbouring localities outside the GLUHFW LQÀXHQFH RI D ZDWHUFRXUVH ,W LV D W\SLFDO ULSDULDQ VSHFLHV ZKLFK XVXDOO\ DYRLGVLQXQGDWLRQEXWEHQH¿WVIURPWKHORFDOO\LQFUHDVHGDLUKXPLGLW\DWWKHERWtom of shady canyons. Important references: Purvis et al. (1992), Wirth (1995).

/HFLGHDDKOHVLL +HSS 1\ODQGHU¿J38) Basionym: Biatora ahlesii Hepp 1860. Synonyms: Mycobilimbia ahlesii (Körber) R. Santesson unpubl., Lecidea rusticula Nylander 1866, Biatora rusticula (Nylander) Walt. Watson 1953, Lecidea valentiorr Nylander 1877, Biatora valentior (Nylander) Walt. Watson 1953. 7KDOOXV grey to green, often ± shiny; thin, continuous to rimose; medulla I-, no lichen substances detected by t.l.c.; photobiont coccoid green alga. Apothecia: sessile, up to 800 m in diameter, red-brown to black, disc concave to slightly convex; true exciple prominent with dark red-brown edges, paler to colourless

Lempholemma

73

Fig. 38. Lecidea ahlesii: ×17. inside; hymenium 60–90 m high; hymenium and hypothecium with bluish-violet to brown-black, K+ green pigment; epihymenium colourless to pale yellowish or reddish-brown; asci of Porpidia-type (with a I+ blue, tube-like apical structure); ascospores unicellular, 12–17 × 5–9 m. (FRORJ\ at shaded sites with high air humidity, typically at the bottom of ravines on stable siliceous rocks, in Central Europe usually terrestrial or in the upper splash water zone but avoiding long periods of inundation. In Great Britain also reported from frequently inundated sites. 'LVWULEXWLRQ generally a rare species but locally rather frequent in areas with usually suboceanic climate.

/HPSKROHPPDKörber 185 Important references: Jørgensen (2007b), Poelt (1969), Schiman-Czeika (1988).

/HPSKROHPPDSRO\DQWKHV%HUQKDUGL 0DOPH¿J 39) Basionym: Lichen polyanthes Bernhardi in Schrader 1797. Selected synonyms: Lempholemma chalazanellum (Nylander) Zahlbruckner 1924, Lempholemma chalazanodes (Nylander) Zahlbruckner 1924, Lempholemma compactum Körber 1855, Lempholemma muelleri (Hepp) Zahlbruckner 1924, Lempholemma myriococcum (Acharius) Th. Fries 1860, Synalissa cancellata F. Wilson 1892. 7KDOOXV blackish or dark olive green, usually dull, small nodular granulose to spreading and to 5 cm in size, in spreading parts membranaceous, irregularly lobate at the margin and with irregular, often warty folds and ridges; attached by

74

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Fig. 39. Lempholemma polyanthes: ×15. scattered rhizohyphae; anatomy ecorticate and homoiomerous with very thin, irregularly arranged hyphae and scattered coiled trichomes of Nostoc photobiont. Apothecia: pycnoascocarps, few to abundant, often lumped on thallus folds or lobe margins, immersed to semi-immersed, small, 0.2–0.3 (0.5) mm wide, ± globose to eventually disc-shaped; thalline margin thick, smooth, persisting; discs at ¿UVWVPDOODQGSXQFWLIRUP¿QDOO\RSHQÀDWWRVOLJKWO\GHSUHVVHG\HOORZLVKRU reddish brown, dull; excip. prop. thin, hyaline; epihymenium yellowish brown, K-; hymenium 100–130 (140) m high, hyaline, I+ blue; subhymenium hyaline; asci 8-spored, thin-walled, I-, ascospores simple, hyaline, broad ellipsoid to subglobose, (7.5) 10–17.5 (20) × 7.5–12.5 (15) m, old ascospore walls to 2.5 (5) m thick, otherwise thin. Pycnidia: immersed to slightly elevated on lobe margins and thallus folds, simple, hyaline, narrow-pyriform to ovoid; pycnospores cylindrical, 2–3 × 1 m. (FRORJ\ usually in sheltered, humid situations on mosses over rock, occasionally along river banks and on old, crumbling walls. 'LVWULEXWLRQ from lowland to alpine regions and widespread in arctic to temperate regions of the northern hemisphere; also in Australia. Notes: Lempholemma polyanthes has a fairly wide ecological amplitude usually growing on mosses or debris in moist, terrestrial habitats. However, forms described as Lempholemma (Collema, Physma) muelleri have been reported from semi-inundated localities at the rivers Arve and Rhine as well as a few small rivers in the Alps.

Leptogium

75

Leptogium (Acharius) Gray 1821 Important references: Guttová (2000), Jørgensen (1994, 2007a), Jørgensen & James (1983), Sierk (1964). Key to the species: 1a Thallus large and distinctly foliose, lobes rather broad, (1) 1.5–5 mm wide ... 2 E Thallus crustose, squamulose, subfruticose or foliose to rosette-shaped with smaller lobes, 0.2–1.5 mm wide ................................................................. 5 2a Apothecia abundant, isidia lacking, lobes broadly rounded, incised, periodically inundated at the base of trees or boulders covered by mosses or soil along eutrophic watercourses .................................................... /ULYXODUH E $SRWKHFLDUDUHWRDEVHQWLVLGLDSUHVHQWRUQRWOREHPDUJLQVODFHUDWH¿PEULate, broadly rounded or undulate ............................................................... 3 3a ,VLGLD ODFNLQJ OREH PDUJLQV ODFHUDWH¿PEULDWH XVXDOO\ QRW LQXQGDWHG EXW often found on mossy boulders or mosses on soil along rivers and streams .............................................. L. lichenoides (Linnaeus) Zahlbruckner 1924 E Isidia present, scale-like or granulose to coralloid, lobe margins broadly rounded or undulate .................................................................................. 4 4a Isidia scale-like, laminal, lobes light to medium grey, adpressed, translucent when wet, on the mossy basis of trees or on mossy boulders along watercourses, occasionally inundated ........................................... L. cyanescens E Isidia globose to coralloid, clustered, lobes dark grey to brownish, undulate, ascending, in seepage tracks and moist siliceous rock, very rarely indundated .............................. L. magnussonii Degelius & P.M. Jørgensen 1994 5a Thallus crustose, inundated on pebbles etc. in mountain streams .../DTXDOH E Thallus squamulose, small-lobate to rosette-shaped or dwarf-fruticose, frequently to very rarely inundated, substrate various ................................... 6 6a Thallus with ± cylindrical branchings ....................................................... 7 E Thallus squamulose to small-lobate and rosette-shaped ............................ 8 7a Thallus forming dense fruticolous cushions composed of minute cylindrical branches c. 0.15 mm wide; usually on mossy bark of trees, rarely on rocks or soil, also along river margins but usually not inundated .......................... ...................................................... L. teretiusculum (Wallroth) Arnold 1892 E Thallus rosette-shaped, branches subcylindrical, furcate, loosely adpressed to somewhat ascending, 0.2–0.35 mm wide; temporarily indundated on rock along mountain rivers ..................................................... /VXEWRUXORVXP 8a 7KDOOXVURVHWWHVFRPSRVHGRIÀDWWHQHGWRFRQFDYHOREHV±PPZLGH ....................................................................................................... /ULYDOH E Thallus forming adnate rosettes with rigid, elongated lobes, in seepage tracks on calcareous rock and temporarily inundated along larger rivers, in lowland Fennoscandia also on lake shores ................................ /SOLFDWLOH

/HSWRJLXPDTXDOH$UQROG 30-¡UJHQVHQ¿J40) Basionym: Leptogium pusillum var. aquale Arnold 1873. 7KDOOXVEODFNLVKFRQWLQXRXVWKLQDQG¿OPOLNHRUSDUWO\JUDQXORVHDQG EHFRPing rimose and somewhat thickened, to 70 m thick; anatomy paraplectenchy-

76

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Fig. 40. Leptogium aquale: ×15. matous throughout; photobiont Nostoc in short trichomes, cells small, 3–4 m. Apothecia: common, to 0.5 mm wide, globose; thalline margin thin, 45–50 m thick, apically thinner and receding; disc brownish, concave; excip. prop. prominent, 35–40 m thick, subparaplectenchymatous, hyaline but apically pale reddish brown, sometimes becoming crenulate; hymenium hyaline, c. 200 m high, I+ blue, epihymenium pale reddish brown; ascospores broad-ellipsoid, submuriform to muriform, 5–6 (7)-septate, (26) 30–45 (48) × 10–14 Pm. Pycnidia: rare, very small, c. 0.1 mm wide, ± immersed in thallus, pycnospores 3.5–4 × 1.2– 1.5 m, dumbbell-shaped. (FRORJ\ on ± inundated calcareous pebbles in mountain streams. 'LVWULEXWLRQ rare and scattered in subalpine regions in the Alps but possibly with wider distribution; recently reported from Sweden (Nordin 2002a). Notes: characterized by the very thin thallus of dark colour with Nostoc photobionts, small apothecia with prominent, pale proper margin and large submuriform to muriform ascospores.

Leptogium cyanescens5DEHQKRUVW .|UEHU¿J 41) Basionym: Collema cyanescens Rabenhorst 1845. Synonyms: Collema tremelloides var. caesium Acharius 1810, Leptogium caesium (Acharius) Vainio 1890, Leptogium tremelloides auct.; for full synonymy see Sierk (1964). 7KDOOXV light to medium grey, usually dull, translucent when wet, smooth to somewhat wrinkled, 1–5 (8) cm in size, adnate, foliose, subdichotomously to irregularly lobate; lobes elongate, ± plane to slightly roughened, discrete, 2–5 mm wide, 35–110 m thick, tips rounded, entire to dentate or irregularly incised and

Leptogium

77

Fig. 41. Leptogium cyanescens: foliose thallus with scale-like isidia; ×8. LVLGLDWHLVLGLDGHQVHODPLQDOWRPDUJLQDOF\OLQGULFDOWRÀDWWHQHGRFFDVLRQDOO\ becoming lobulate, simple to eventually branched, lobes corticate, upper and lower cortex consisting of single layer of ± isodiametric cells, 5–8 m in size; photobiont Nostoc as clearly visible trichomes. Apothecia: rare, laminal, sessile to substipitate, 0.5–2 mm; thalline margin greyish or cream-coloured, entire or isi-diate; disc slightly concave to plane or slightly convex, usually reddish brown to pale brownish; hymenium 95–130 m high, I+ blue; ascospores 8 per ascus, hyaline, ellipsoid to fusiform, submuriform with 3 transverse and 0–1 longitudinal septum, 18–23 × 6–10 m. Pycnidia: not observed. (FRORJ\ on mossy bark towards the tree base and on moist, mossy or naked siliceous or basic, more rarely calcareous rock, also sporadically or periodically inundated at river banks; ecologically similar to &ROOHPDÀDFFLGXP. 'LVWULEXWLRQ in central Europe rare and predominantly in sub- to high montane, oceanic regions; distributed world wide. 6LPLODUVSHFLHV Leptogium cochleatum is similar. In the British Isles it is said to occur on sheltered trunks of old deciduous trees and mossy boulders especially close to streams. It differs from Leptogium cyanescens in the lack of isidia and rather thick, striate lobes.

/HSWRJLXPSOLFDWLOH$FKDULXV /HLJKWRQ¿J 42) Basionym: Lichen plicatilis Acharius 1795. Selected synonyms: Collema hydrocharum (Acharius) Acharius 1810, Collema plicatile (Acharius) Acharius 1810, Leptogium cataclystum var. ÀXFWXDQV (Krempelhuber) Zahlbruckner 1924, Leptogium hydrocharum (Acharius) Zahlbruckner 1924, Leptogium plicatile var. hydrocharum (Acharius) A.L. Smith 1918.

78

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Fig. 42. Leptogium plicatile: thallus lobes wrinkled, partly with apothecia and small, globose proliferation; ×3. 7KDOOXV dark brown-black, up to 5 cm wide, usually forming closely adpressed rosettes, lobes rather short and thick, heavily wrinkled, tips rounded, up to 3 mm long and 350 m thick, strongly swelling when wet, surface uneven, rigid and with isidia-like warts, indistinctly corticate, photobionts clustered towards thallus surface, medulla thick; photobiont Nostoc in short trichomes. Apothecia: fairly common, marginal or laminal, to 1.5 mm wide; thallus margin prominent; disc concave, dark brownish; hymenium 135–160 m high, I+ blue; ascospores 8 per ascus, hyaline, ellipsoid, muriform, 20–25 (30) × 8–16 m. Pycnidia: unknown. (FRORJ\ usually on calcareous rocks moistened by seeping water, also on conFUHWHERXOGHUVDORQJWKH5LYHU'DQXEHÀXVKHGE\ZDYHDFWLRQ (F. Berger, pers. comm.); in lowland Fennoscandia also in the submerged zone of lakes. 'LVWULEXWLRQ from lowland to montane and subalpine regions; widespread in Europe but amphibious populations rather local, e.g. in Austria along the Danube (F. Berger, pers. comm.). Notes: According to Jørgensen (1994) the status of Leptogium subtorulosum as a taxon distinct from L. plicatile is in need of further studies.

/HSWRJLXPULYDOH 7XFNHUPDQ¿J43) Synonym: Polychidium rivale (Tuckerman) Fink 1935. 7KDOOXV often dark emerald green, 0.5–2 cm wide, usually closely adnate but also VSUHDGLQJ RQWR PRVVHV IROLRVH WR VTXDPXORVH FRPSRVHG RI ÀH[XRXV OREXOHV RU squamules, lobules 0.2–1.5 mm wide, smooth and glossy or dull, usually tightly adpressed with occasionally ascending margins, becoming crowded and imbricate, tips

Leptogium

79

Fig. 43. Leptogium rivale: thallus small-lobed with some apothecia; ×16. roundish, isidia absent; lobules 50–150 m thick, paraplectenchymatous throughout, distinctly corticate on both sides with cortex formed by prosoplectenchymatous cells; photobiont Nostoc (?) in short trichomes. Apothecia: rare, laminal, sessile, to 1 mm ZLGHDW¿UVWJORERVHWKDOOXVPDUJLQSDOHUWKDQWKHDSRWKHFLDO GLVFGLVFFRQFDYH dark brownish; hymenium 100–150 m high, I+ blue; ascospores 8 per ascus, hyaline, ellipsoid, muriform, 20–39 (46) × (6) 9–14 m. Pycnidia: not observed. (FRORJ\ on mossy siliceous boulders in fast running, cold mountain streams as well as in lakes; in western North America usually permanently inundated on bare rock. 'LVWULEXWLRQ montane to subalpine regions; in Europe known only from an old collection in western Romania but recently reported from localities in the western Carpathians in southern Poland and in the Giant Mountains/Krkonoše in the Czech Republic (Guttová 2000, Palice, pers. comm.) as well as from the Serre de Estrela in Portugal (van den Boom & Jansen 2002); otherwise occurring widespread in the Rocky Mountains, Sierra and Cascade Range regions in western North America reaching south-eastern Alaska. Notes: Jørgensen (1994) and Guttová (2000) point to slight differences in the shape of the thallus and apothecia as well as in the spore size between the recently discovered European specimens and the material from North America; further studies needed.

/HSWRJLXPULYXODUH$FKDULXV 0RQWDJQH¿J44) Basionym: Lichen rivularis Acharius 1799. Synonyms: Gabura rivularis (Acharius) O. Kuntze 1891, Leptogiopsis rivulare (Acharius) Trevisan 1880, Leptogium crenatellum Tuckerman 1859, Leptogium sernanderi Du Rietz 1922.

80

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Fig. 44. Leptogium rivulare: thallus broadly lobate with laminal apothecia; ×11. 7KDOOXV greyish blue, dull, reaching few cm in size, ± closely adnate, foliose; OREHVGXOOVPRRWKÀDWLQFLVHG ±PPZLGH±—PWKLFNWLSVURXQded, isidia absent, lobes distinctly corticate on both sides; photobiont Nostoc in trichomes. Apothecia: numerous, laminal, sessile to substipitate, to 1 mm wide; thalline margin greyish blue, narrow, entire, often becoming excluded; disc slightly concave to plane or slightly convex, pale reddish to pale brownish; hymenium 75–105 m high, I+ blue; ascospores 4 per ascus, hyaline, ellipsoid, 3 (4)-septate to submuriform, (15) 16–20 (22) × (6) 7–8 (10) m. Pycnidia: not observed. (FRORJ\ growing periodically submerged on exposed roots and stems along the margins of sluggish rivers and ponds, occasionally also on rocks. 'LVWULEXWLRQ obviously rare, historical records from the River Seine in northern France; also in Finland, Sweden, Estonia and the Great Lakes region in North America.

/HSWRJLXPVXEWRUXORVXP (Nylander ex Stizenberger) Degelius 1954 ¿J45) Basionym: Collema subtorulosum Nylander ex Stizenberger 1882. 7KDOOXV dark olive green or brownish, dull, often incrusted by silt, loosely adpressed, forming narrow-lobed, deeply branched rosettes; lobes almost cylindriFDOWRVOLJKWO\ÀDWWHQHGDQGVRPHZKDWNQRWW\“GLFKRWRPRXVO\EUDQFKHG±PP long, c. 0.2–0.35 mm thick, surface slightly uneven, lobe tips usually somewhat lifted, central lobes becoming ascending and sometimes with globose to subcylindrical isidia or accessory lobules, otherwise loosely adpressed; lobes at least partially distinctly corticate, cortical cells isodiametric, c. 5 m; photobiont Nostoc in short, twisted trichomes, not distinctly crowded towards the lobe periphery, hyphae irregularly arranged, cells somewhat elongated to angulate, to 15 × 3–

Leptogium

81

Fig. 45. Leptogium subtorulosum: thallus rosette with ± cylindrical, radiating lobes; ×13. 5 m, medulla rather compact, thallus anatomy in general resembling that of L. magnussonii VHH -¡UJHQVHQ   ¿J E  DFFRUGLQJ WR -¡UJHQVHQ   close to L. plicatile VHH -¡UJHQVHQ   ¿J E  Apothecia: apparently very rare and not studied so far. Pycnidia: not studied so far. (FRORJ\ semi-inundated on mostly calcareous boulders along river banks; the specimens seen were often accompanied by 3\UHQRFDUSRQÀRWRZLDQXP and various pyrenocarpous lichens forming a community resistant to at least moderate silting. 'LVWULEXWLRQ so far only known from the river Arve in the western Alps and seen among material of Placynthium tantaleum and 3\UHQRFDUSRQ ÀRWRZLDQXP collected 50 years ago in the Lech ravine in southern Bavaria and observed in fertile condition (!) in type material of Psorotichia riparia Arnold (=Porocyphus rehmicus (A. Massalongo) Zahlbruckner) collected at the bank of the Danube in Lower Bavaria c. 150 years ago; also recorded from the stream Langbathseebach in Upper Austria (F. Berger, pers. comm.); two further records of similar material exist from the rivers North Tyne and Eden in England (Gilbert & Giavarini 1997 [sub L. massiliense], Jørgensen 1994); also in the Azores. 6LPLODUVSHFLHVLeptogium subtorulosum belongs to the aggregate of L. plicatile, but the status of the species is still unsettled (Jørgensen 1994). The characteristic thallus shape of L. subtorulosum PD\EHFDXVHGE\WKHLQÀXHQFHRIUXQning water. Leptogium massiliense is similar because of the cylindrical, furcate branches. It differs, however, in the non-aquatic habitat. Leptogium teretiusculum has been reported from the river Mosel by Diederich & Sérusiaux (2000). It is, however, a predominantly terricolous or corticolous species. In Scandinavia, Leptogium plicatile occurs on sometimes submerged calcareous rock and on acidic rock irrigated by calciferous water (Jørgensen 2007a, Santesson et al. 2004). In Central Europe, however, this species is usually reported from terrestrial habitats only (e.g. Wirth 1995).

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/RERWKDOOLD(Clauzade & Cl. Roux) Hafellner 1991 Important references: Hafellner (1991).

/RERWKDOOLDPHODQDVSLV $FKDULXV +DIHOOQHU¿J 46) Basionym: Parmelia melanaspis Acharius 1803. Synonyms: Lecanora melanaspis (Acharius) Acharius 1810, Lichen melanaspis (Acharius) Wahlenberg 1812, Parmularia melanaspis (Acharius) Räsänen 1939, Placodium melanaspis (Acharius) Link 1833, Placolecanora melanaspis (Acharius) Räsänen 1943, Squamaria melanaspis (Acharius) Elenkin 1904, Aspicilia melanaspis (Acharius) Poelt & Leuckert 1973. 7KDOOXV grey to whitish, changing colour to green when wet; thick, margins with “HORQJDWHGUDGLDWLQJOREHVOREHVRQO\ORRVHO\¿[HGWRVXEVWUDWXP±PP wide, with pale lower surface; medulla P-, K-, photobiont a coccoid green alga. Apothecia: semi-immersed to almost sessile, up to 1.8 mm in diameter, with prominentWKDOOLQHH[FLSOHGLVFÀDWWRFRQYH[UHGEURZQWREODFNEURZQ t HSLWKHFLXP red-brown, N-; hymenium 60–75 m high; ascospores unicellular, 10–13 × 6–10 m. Conidia 4.5–6 × 1 m. (FRORJ\ amphibious on stable boulders in the splash water zone of usually very turbulent streams, e.g. in rapids; heliophilous. 'LVWULEXWLRQ scattered in high-montane to alpine areas, usually above the timber line. 6LPLODU VSHFLHV Otherwise similar amphibious species of Aspicilia are tightly adpressed to the substratum, never have radiating elongated marginal lobes, and the apothecia are more immersed. The conidia are longer in Aspicilia and the chemistry is different. Lobothallia radiosa is a terrestrial species with thin lobes being tightly adpressed to the substratum and a P+ orange, K+ red thallus (norstictic acid detected by t.l.c.). It is often found in the immediate vicinity of rivers

Fig. 46. Lobothallia melanaspis: ×6.

Phaeophysica

83

on stable rocks or technical buildings where it is not inundated or inundated only GXULQJH[FHSWLRQDOKLJKÀRRGV,WPD\ EHUHIHUUHGWRDVDULSDULDQOLFKHQPhaeophyscia endococcina is a foliose species with rhizines on the lower surface, K+ purple medulla and larger ascospores (17–24 m).

Phaeophyscia Moberg 1977 Important references: Moberg (1977), Purvis (1992), Wirth (1995). Key to the species: 1a Isidia or soralia lacking, usually with apothecia, restricted to amphibious sites ..................................................................................... P. endococcina E Isidia or soralia present, rarely with apothecia, species with wide ecological amplitude, only occasionally in freshwater habitats .................................. 2 2a Lobe margins (later also in the centre of the lobe) with dark isidia, soralia absent .................................................. P. sciastra (Acharius) Moberg 1977 E Lobe margins lacking isidia, with patches of soralia in lobe centre .......... 3 3a Lobes up to 0.3 mm wide, dark brown to black-brown ................................ .............................................................. P. nigricans (Flörke) Moberg 1977 E Lobes to 1.2 mm wide, grey to brown or greenish ....................................... ........................................................... P. orbicularis (Necker) Moberg 1977

Phaeophyscia endococcina .|UEHU 0REHUJ¿J47) Basionym: Parmelia endococcina Körber 1866. 7KDOOXV upper site brownish-olive-grey, lower side black with black rhizines, thallus foliose, adpressed to the substratum, rosette-like with narrow lobes 0.3–1 mm wide; medulla white or here and there with orange-red pigment (skyrine, K+

Fig. 47. Phaeophyscia endococcina: ×11.

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purple); isidia and soralia absent; photobiont coccoid green alga (Trebouxia impressa YHUL¿HG E\ FXOWXUHG LVRODWHV DQG ,76VHTXHQFHV  Apothecia: frequent, sessile, up to 1 mm in diameter; ascospores uniseptate, 17–24 × 7–11 m. (FRORJ\ on stable siliceous substrata in the splash water zone at moderately shaded to sunny localities. 'LVWULEXWLRQ mainly in upland areas, in Central Europe very rare outside the Alps and Carpathians; widespread in Nordic countries. 6LPLODUVSHFLHV Other species of Phaeophyscia or Physcia occurring at streams and rivers have soredia or isidia. Lobothallia melanaspis is a crustose species lacking rhizines on the pale coloured lower surface, K- medulla and smaller ascospores (10–13 m).

3ODF\QWKLXP (Acharius) Gray 1821 Important references: Czeika & Czeika (2007), Foucard (1990, 2001), Gyelnik (1940), Henssen (1963b), Jørgensen (2007c), Poelt (1969). Key to the species: 1a 7KDOOXVZLWKGLVWLQFWO\HI¿JXUDWHPDUJLQRUUDGLDWLQJPDUJLQDOOREXOHV .... 2 E 7KDOOXVQRWPDUJLQDOO\HI¿JXUDWHGLVWLQFWO\UDGLDWLDOOREXOHV DWPDUJLQVDEsent ............................................................................................................ 5 2a Marginal lobules canaliculate to cylindrical, 0.05–0.2 mm wide, thallus dark olive to blackish, centrally densely papillose, usually on moist rock (seepage WUDFNVHWF EXWSRVVLEO\DOVRUHSRUWHGIURPWHPSRUDULO\ÀXVKHG URFNV ......... ...................................................... P. asperellum (Acharius) Trevisan 1869 E 0DUJLQDOOREXOHVGLVWLQFWO\ÀDWWHQHGRURQO\VOLJKWO\FRQYH[QHYHUF\OLQGULcal, 0.1–0.5 mm wide, thallus dark to greyish-olive, not papillose ........... 3 3a 7KDOOXV ROLYH WR JUH\LVKEURZQ PDUJLQDO OREXOHV GLVFUHWH GLVWLQFWO\ ÀDWtened and tightly adpressed, 0.2–0.5 mm wide, thallus eventually covered with scale-like isidia, sometimes fertile ............................... 3ÀDEHOORVXP E Thallus olive, marginal lobules contiguous to somewhat imbricate, usually not GLVWLQFWO\ÀDWWHQHGDQGWLJKWO\DGSUHVVHGWKDOOXVODFNLQJLVLGLDRULISUHVHQW isidia distinctly elongated and not scale-like, apothecia present or not ........ 4 4a 0DUJLQDOOREXOHVXVXDOO\GLVWLQFWO\HORQJDWHGDQG¿QHO\VWULDWH± PP long, 0.1–0.25 (0.3) mm wide, thallus centrally with abundant cylindrical to VOLJKWO\ÀDWWHQHGHORQJDWHGLVLGLDLQCentral Europe so far only known in sterile condition ................................................................ 3SDQQDULHOOXP E Marginal squamules or lobules rather short and often slightly wider than ORQJ¿QHO\VWULDWHRUQRW± PPORQJ±PPZLGHWKDOOXV centrally becoming thick and subsquamulose or remaining thin especially in water, isidia lacking, apothecia usually present, often with pale margin and brownish to reddish disc ........................................................... 3URVXODQV 5a Ascospores 4–5-celled, 19–35 (46) × 3.5–6 m, apothecia usually rather big, 0.3–1 (1.5) mm wide, dark bluish prothallus usually absent or very indistinct .............................................................................. 3GROLFKRWHUXP

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E Ascospores 1–2-celled, shorter, apothecia usually 0.3–1 mm wide, bluish prothallus present or not ........................................................................... 6 6a Prothallus distinct, usually pale-blue, thallus composed of densely aggregated papillae or consisting of more discrete upright granules, growing on calcareous or siliceous rock, ascospores (9) 10–12 (15) × (5) 6–7 (8) m ... ................................................................................................ 3WDQWDOHXP E Prothallus absent, thallus thin, forming a uniform to irregularly cracked to eventually areolate crust, no discrete upright granules, growing exclusively on siliceous rock, ascospores (12) 15–19 (21) × 5–7 (8) m ....................... ............................................................................. P. nigrum f. crustaceum

3ODF\QWKLXPGROLFKRWHUXP1\ODQGHU 7UHYLVDQ¿J48) Basionym: Pannaria dolichotera Nylander 1861. Selected synonyms: Lecothecium dolichoterum (Nylander) Blomberg & Forssell 1880, Lecothecium pluriseptatum Arnold 1870, Pannaria melantera Stirton 1879, Pannularia melantera (Stirton) Crombie 1894, Parmeliella melantera (Stirton) A.L. Smith 1918, Placynthium dolichoterum f. melanterum (Stirton) Gyelnik 1939, Placynthium dolichoterum var. pluriseptatum (Arnold) Servít 1937, Placynthium pluriseptatum (Arnold) Arnold 1889. 7KDOOXV prothallus usually lacking or thin and inconspicuous, pale to dark blue; thallus dark olive-green or brownish-olive, dull, 1–3 (4) cm wide, crustose, usually rimose-areolate; areoles subpapillose, 0.15–0.8 mm wide, 0.15–0.5 (1) mm WKLFNDQJXODWH“LVRGLDPHWULFÀDWWRVOLJKWO\FRQYH[VXUIDFHURXJKRUZDUW\WR uneven, lower surface dark with blackish hypothallus; upper cortex indistinct and thin, olivaceous, thallus otherwise paraplectenchymatous throughout, distinct medulla lacking, upper thallus parts with living photobionts only to 0.15 mm high, lower cortex dark, paraplectenchymatous, supporting dark rhizohyphae;

Fig. 48. Placynthium dolichoterum: thallus crustose, surface of areoles rough; ×18.

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photobiont Scytonema-like, trichomes ± vertical or coiled, much split. Apothecia: usually present, sessile, quite large, (0.3) 0.35–1 (1.5) mm wide, margin EODFNHQWLUHDQGXVXDOO\JORVV\HYHQWXDOO\VLQXRVHDW¿UVWVOLJKWO\SURPLQHQW ¿QDOO\UHFHGLQJWKDOOLQHPDUJLQDEVHQWGLVFDW¿UVWVOLJKWO\FRQFDYHVRRQ“ÀDW DQG¿QDOO\VOLJKWO\WRGLVWLQFWO\FRQYH[EODFN excip. prop. dark brownish purple, 65–70 m thick; hymenium 75–80 (110) m high; epihymenium dark dirty greyish blue green; ascospores hyaline, narrow-ellipsoid, 4 (5)-celled, (19) 20–35 (46) × 3.5–6 Pm, sometimes spirally twisted in ascus. Pycnidia: 90–150 m wide, pycnidial wall usually bluish; pycnospores bacilliform, (4) 5–6 × 1 m. (FRORJ\ in humid areas on occasionally inundated siliceous or calcareous boulders along mountain streams, also on mosses and inclined rock faces long moistened from seeping water; in Fennoscandia on well-lit, moist, schistose rock overgrowing mosses. 'LVWULEXWLRQ montane to alpine, Alps, Vosgues, Black Forest and High Tatra; also known from Scandinavia (not amphibious there) and Scotland. 6LPLODUVSHFLHV Placynthium tantaleum and P. nigrum f. crustaceum are distinguished by the presence of smaller, 2-celled ascospores.

3ODF\QWKLXPÀDEHOORVXP7XFNHUPDQ =DKOEUXFNQHU¿J 49) Basionym: 3DQQDULDÀDEHOORVD Tuckerman 1862. Synonyms: Anziella adglutinata (Anzi) Gyelnik 1939, Lecothecium adglutinatum Anzi 1862, Placynthium adglutinatum (Anzi) Trevisan 1869, Placynthium pannariellum f. sparsum Gyelnik 1940, Pterygium adglutinatum (Anzi) Nylander 1882.

Fig. 49. 3ODF\QWKLXPÀDEHOORVXPWKDOOXVPDUJLQHI¿JXUDWHZLWKFORVHO\DGSUHVVHG lobules; ×23.

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7KDOOXV prothallus absent; thallus greyish-brown to olive, rosettes to 1–2 (4) cm ZLGHPDUJLQGLVWLQFWO\HI¿JXUDWHZLWKFORVHO\DGSUHVVHGÀDWOREHVWRPPORQJ and 0.2–0.3 (0.5) mm wide, lobe tips broadened, to 0.8 mm wide, thallus cenWUDOO\ZLWKDUHROHV±PPZLGHDQG PPWKLFN“FRYHUHGE\ÀDWUDUHO\ cylindrical isidia, isidia to 1 × 0.1–0.3 mm in size; thallus lobes 65–150 m thick with hyphae mainly running parallel; lower surface and rhizines bluish black; photobiont Scytonema-like, trichomes stretched or coiled towards the upper thallus surface. Apothecia: to 1.5 mm wide, margin black, entire; thalline margin absent; disc brown or blackish; excip. prop. 90–140 m thick, marginally with long-celled hairs; hymenium 80–140 Pm high; epihymenium dark green or brown; asci (4) 8-spored, ascospores narrow ellipsoid, hyaline, (3) 4-celled, (10) 13.5–18.5 (20) × 4–6 (8) Pm. Pycnidia: c. 0.2 mm wide, ostiole dark green; pycnospores slightly dumbbell-shaped, 4–5 × 1 m. (FRORJ\ growing temporarily inundated on siliceous, often mineral rich rocks along cold mountain streams and lake shores below high water level; also in drainage lines long wetted from seeping water. 'LVWULEXWLRQ (high)montane to subalpine and very rare in Central Europe, known from the Vosgues, southern Black Forest, W Sudetes, Bohemian Forest and rare in the Alps; fairly widespread in Fennoscandia, also British Isles and North America. 6LPLODUVSHFLHV Placynthium pannariellum is similar but has an olivaceous thalOXVDQGHORQJDWHGF\OLQGULFDOWRVOLJKWO\ÀDWWHQHGLVLGLD

3ODF\QWKLXPQLJUXP f. crustaceum Hepp in Harmand ex Gyelnik ¿J 50) 7KDOOXV prothallus absent; thallus dark olive to greyish-brown, dull, to 5 cm in size, 0.1–0.2 mm thick, 0.6 (1) mm thick in the center, thinning out towards the margin, otherwise uniformly crustose to rimose-areolate to eventually areolate; areoles irregularly shaped, 0.8–1.6 mm, surface smooth to somewhat uneven, composed of very densely aggregated, erect papillae or minute lobules, isidia absent; distinct upper cortex lacking, thallus otherwise paraplectenchymatous or densely reticulate with hyphae ± in vertical rows, cells ± isodiametric or somewhat elongated especially in the upper photobiont layer, medulla lacking or very thin and indistinct, lower cortex or 1–2 (3) rows of small, roundish to slightly angulate hyphal cells; photobiont Scytonema, trichomes split, ± vertically arranged. Apothecia: usually present, immersed to semi-sessile, 0.32–0.75 (1) mm ZLGHPDUJLQEODFNLVKHQWLUH DW¿UVWVOLJKWO\SURPLQHQWHYHQWXDOO\VRPHZKDWreFHGLQJWKDOOLQHPDUJLQDEVHQWGLVFDW¿UVWFRQFDYHVRRQ“ÀDWDQG¿QDOO\VOLJKWly convex, smooth, dull, blackish; excip. prop. dark bluish green, 50–65 (75) m thick; hymenium 70–85 m high; epihymenium dark bluish green; ascospores hyaline, ellipsoid, (0) 1-septate (12) 15–19 (21) × 5–7 (8) Pm, cells sometimes unequal. Pycnidia: not observed. (FRORJ\ usually long inundated on siliceous rocks along mountain streams; according to Keller & Scheidegger (1994) ecologically similar to Staurothele clopimoides and Koerberiella wimmeriana.

88

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Fig. 50. Placynthium nigrum f. crustaceum: thallus crustose, rimose with some apothecia; ×16. 'LVWULEXWLRQ so far recognized only a few times in the Swiss Alps but corresponding specimens were found in Scandinavia as well (P.M. Jørgensen, pers. comm.); to be expected in suitable sites in the Alps and high mountainous regions and among gatherings of either Placynthium nigrum or P. tantaleum from corresponding localities. 6LPLODU VSHFLHV Placynthium nigrum f. crustaceum was recently accepted as distinct from the main form (Czeika & Czeika 2007). It differs from the usually terrestrial P. nigrum f. nigrum and the semi-aquatic P. tantaleum in the lacking prothallus and the 2-celled, slightly larger ascospores. Unlike cited in Czeika & Czeika (2007), WKH¿UVWYDOLGGLDJQRVLVIRU P. nigrum f. crustaceum was provided by Gyelnik (1938).

3ODF\QWKLXPSDQQDULHOOXP1\ODQGHU +0DJQXVVRQ¿J 51) Basionym: Pterygium pannariellum Nylander 1859. Synonyms: Lecothecium pannariellum (Nylander) Blomberg & Forssell 1880, ?Placynthium pannariellum f. conferciens (Nylander) Räsänen 1936, ?Pterygium conferciens Nylander 1875. 7KDOOXV prothallus absent or indistinct, bluish, visible only between marginal squamules or lobules; thallus dark olive-green or olive-brown, thallus forming VPDOOURVHWWHV± FPZLGHZLWKGLVWLQFWO\HI¿JXUDWHPDUJLQVPDUJLQDOVTXDPXOHVRUOREXOHVÀDWRUVOLJKWO\FRQYH[XVXDOO\GLVWLQFWO\HORQJDWHGDQG¿QHO\ striate, 0.4–1 (1.5) mm long, 0.1–0.25 (0.3) mm wide, 75–80 m thick, tips URXQGHGVOLJKWO\REWXVHRU¿QHO\FUHQDWHWKDOOXVFHQWUHEUHDNLQJLQWRLUUHJXODUO\ shaped squamules or areoles 1–2.5 mm wide and 0.4–0.8 mm thick, soon disVROYHGLQWRDEXQGDQWQDUURZHORQJDWHGVOLJKWO\ÀDWWHQHGLVLGLDORZHUVXUIDFH

Placynthium

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Fig. 51. Placynthium pannariellumWKDOOXVPDUJLQHI¿JXUDWHZLWKORVHO\DGSUHVVHG lobules, centrally with elongated, ascending isidia; ×14. blackish with distinct dark hypothallus; upper cortex thin, olivaceous, composed of a single row of small cells, medulla thick, whitish, composed of densely aggregated, robust, somewhat elongated hyphae, periclinally arranged or fan-shaped towards the surface, lower cortex dark, paraplectenchymatous supporting dark rhizohyphae; photobiont Scytonema-like but trichomes short, much split and contorted towards the thallus surface. Apothecia: not seen in Central European material, sessile, 0.5–0.8 (1) mm wide, margin black, entire; thalline margin absent; GLVFDW¿UVWFRQFDYHVRRQ“ÀDWRU¿QDOO\VOLJKWO\FRQYH[EDUNEURZQLVKRUGDUN reddish brown or almost black; hymenium to 115 m high; epihymenium dark greenish blue brown; ascospores hyaline, 4-celled, 15–20 (21) × (3) 4–5 (6) Pm. Pycnidia: not observed. (FRORJ\ usually submerged on siliceous rocks along rivers, lakes and lake outlets. 'LVWULEXWLRQ colline to subalpine, Alps and Carpathians; also known from the Pyrenees, Les Causses, Provence, North Wales and North Scotland; widely distributed in Fennoscandia, also in Greenland. 6LPLODU VSHFLHV 3ODF\QWKLXP ÀDEHOORVXP is very similar and distinguished by the very closely adpressed marginal lobes and scale-like isidia.

3ODF\QWKLXPURVXODQV 7K)ULHV =DKOEUXFNQHU¿J 52) Basionym: Lecothecium corallinoides ssp. rosulans Th. Fries 1863. Synonyms: Lecidea microphylla var. radiata Wahlenberg 1812, Lecothecium corallinoides var. rosulans (Th. Fries) Hellbom 1871, Placynthium pannariellum var. rosulans (Th. Fries) Degelius 1943, Placynthium pannariellum var. squamulosum Räsänen 1936.

90

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Fig. 52. Placynthium rosulansWKDOOXVPDUJLQHI¿JXUDWHZLWKVKRUWEURDGOREXOHV centrally areolate with some apothecia; ×9. 7KDOOXV prothallus partly distinct, dark blue to dark greenish; thallus dark olivegreen or olive-brown, sometimes with ochraceous tinge, often with silky gloss, thallus 0.5–3 cm wide, sometimes larger, forming small, squamulose rosettes ZLWKGLVWLQFWO\HI¿JXUDWHPDUJLQVPDUJLQDOOREXOHVXVXDOO\¿QHO\VWULDWHVOLJKWO\ FRQYH[ RU DOPRVW ÀDW VRPHZKDW HORQJDWHG WR RIWHQ VOLJKWO\ ZLGHU WKDQ ORQJ ± PPORQJ± PPZLGHWLSV¿QHO\FUHQDWHXVXDOO\FORVHO\ attached, thallus centre breaking into irregularly shaped squamules or areoles 0.4–1.6 mm wide and 0.1–0.4 mm thick, thallus surface rough or granulose but GLVWLQFWLVLGLDDEVHQWWKDOOLIRUPHGXQGHUWKHLQÀXHQFHRIUXQQLQJZDWHUWKLQQHU lower surface blackish with distinct dark hypothallus; upper cortex thin, olivaceous, medulla thick, whitish, composed of densely aggregated, robust, elongated hyphae, periclinally arranged or fan-shaped towards the surface, lower cortex dark, paraplectenchymatous, supporting abundant dark rhizohyphae; photobiont Scytonema-like, trichomes usually stretched. Apothecia: usually present, sessile, 0.4–0.8 (1) mm wide, margin blackish, often with a dark bluish or brownish grey WLQJH HQWLUH DW ¿UVW UDWKHU WKLFN DQG SURPLQHQW HYHQWXDOO\ UHFHGLQJ WKDOOLQH PDUJLQDEVHQWGLVFDW¿UVWFRQFDYHVRRQ“ÀDWRU¿QDOO\VOLJKWO\FRQYH[EDUN brownish or dark reddish brown, rarely almost black; excip. prop. dark bluish green, (60) 70–80 (90) m thick; hymenium c. 125 m high; epihymenium dark greyish brown; ascospores hyaline, cylindrical to narrow ellipsoid, (2) 4-celled, (10) 12–18 (20) × (4) 5–6 Pm. Pycnidia: c. 150 m wide, ostiole dark blue; pycnospores 5–7 × 1 m. (FRORJ\ on occasionally inundated siliceous or calcareous rocks along lake shores and mountain streams as well as in seepage tracks on inclined rock faces. 'LVWULEXWLRQ colline to alpine, apparently very rare in Central Europe, Tyrol; also in western Romania, Sweden and Finland.

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91

6LPLODU VSHFLHV Placynthium pannariellum is very similar. It differs in the slightly thinner marginal lobules and the presence of abundant cylindrical to VOLJKWO\ ÀDWWHQHG HORQJDWHG LVLGLD 3ODF\QWKLXP ÀDEHOORVXP GLIIHUV LQ WKH ÀDW PDUJLQDOOREXOHVDQGWKHSUHVHQFHRIVKRUWÀDWWHQHGWRVFDOHOLNHFHQWUDOLVLGLD

3ODF\QWKLXPWDQWDOHXP +HSS +XH¿J 163) Basionym: Biatora corallinoides var. tantalea Hepp 1867. Synonyms: Pannularia nigra f. tantalea (Hepp) Stizenberger 1882, Placynthium diblastum Gyelnik 1940, Placynthium nigrum var. tantaleum (Hepp) Arnold 1873, Racoblenna tantalea (Hepp) Trevisan 1869. 7KDOOXV prothallus usually very distinct, pale to rarely dark-blue, often whitish at the outermost margin; thallus light to dark brownish-olive, dull, sometimes with pale lime crystals between the areoles, thallus 2–4 cm wide, c. 0.5 mm thick, centrally to 1.5 mm thick, crustose, granulose to rimose-areolate; areoles eventually 1–1.5 (2) mm wide, usually even, composed of erect papillae or minute lobules and thus appearing papillose, papillae 0.1–0.2 mm thick, densely to rather loosely aggregated, isidia absent; distinct upper cortex lacking, otherwise paraplectenchymatous, hyphae ± in vertical rows, cells ± isodiametric or somewhat elongated especially in the upper photobiont layer, medulla lacking or very thin and indistinct with isodiametric hyphal cells, lower cortex or 1–2 (3) rows of small, roundish to slightly angulate hyphal cells; photobiont Scytonema-like, trichomes ± vertically arranged, split and sometimes coiled. Apothecia: sometimes abundant, immersed to semi-sessile, 0.32–0.75 (1) mm wide, margin black, enWLUHURXQGLVKWRHYHQWXDOO\VLQXRVHDW¿UVWVOLJKWO\SURPLQHQWHYHQWXDOO\UHFHGLQJWKDOOLQHPDUJLQDEVHQWGLVF DW¿UVWFRQFDYH VRRQ“ÀDW DQG¿QDOO\VOLJKWO\ convex, smooth, dull, blackish; excip. prop. dark bluish green, 50–70 m thick; hymenium 100–110 m high; epihymenium bright bluish green; ascospores hyaline, ellipsoid, (0) 1-septate, (9) 10–12 (15) × (5) 6–7 (8) Pm. Pycnidia: 70–135 m wide, ostiole bluish green; pycnospores (4) 6–8 × 1 m. (FRORJ\ on inundated or persistently wetted calcareous or siliceous rocks and boulders in or along mountain rivers and streams. 'LVWULEXWLRQ montane to subalpine, scattered in the mountains of Central Europe and in the Alps; also in the Pyrenees, Scandinavia, Canada and northern USA. 6LPLODUVSHFLHV Henssen (1963b) treated Placynthium tantaleum as a variety of P. nigrum differing from the latter in the simple to 1-septate, broad ellipsoid ascospores. This view has not been adopted in the recent European literature.

92

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3RO\EODVWLDA. Massalongo 1852 nom. cons. Notes: This genus is separated from Thelidium only by the presence of additional longitudinally septate ascospores. However, in several species of Thelidium ascospores with longitudinal septa may occasionally be observed among the majority of exclusively transversely septate ascospores. According to recent molecular studies the species with a thallus mantle covering the perithecium surface and brownish ascospores form a monophyletic group for which the name Sporodictyon$0DVVDORQJRLVDYDLODEOH6DYLüHWDO 7KHRQO\DPSKLELous species of this genus in Central Europe is S. cruentum (=Polyblastia cruenta), but others are ± hydrophytic and can be found in the vicinity of cataracts. Terrestrial representatives of Polyblastia have been shown to form a polyphyletic JURXS6DYLüHWDO 0ROHFXODUGDWDIRU several Central European freshwater species of Polyblastia, however, are still missing and further changes in the circumscription of the genus are to be expected. Important references: Clauzade & Roux (1985), Gueidan et al. (2007, 2008), 6DYLüHWDO =VFKDFNH  .H\WRWKHVSHFLHVLQFOXGLQJWKHRQO\ aquatic species of Sporodictyon from WKHVWXG\UHJLRQ  1a Perithecia covered by a thin thallus mantle at least when young, ascospores muriform, brownish when old (subgen. Sporodictyon sensu Zschacke 1934), thallus epilithic on siliceous rocks ....................... Sporodictyon cruentum E Perithecia naked, thallus epi- or semi-endolithic, ascospores partly with transversal septa only or with a few longitudinal septa in the middle of the ascospores, always colourless (subgen. Thelidioides sensu Zschacke 1933), RQ FDOFDUHRXV RU VLOLFHRXV URFNV VRPHWLPHV GLI¿FXOW WR GLVWLQJXLVK IURP Thelidium, see also there) ......................................................................... 2 2a Ascospores 22–32 m long ..................................................... P. peminosa E Ascospores > 32 m long ......................................................................... 3 3a Perithecia completely immersed in the thallus and the substratum, involucrellum lacking ............................................................................. 3ODYDWD E Perithecia at least with upper third raised above the surrounding thallus and the substratum, with distinct apical involucrellum ........................ 3ULYDOLV

3RO\EODVWLDODYDWD Zschacke 1933 Synonym: Amphoroblastia lavata (Zschacke) Servít 1954. 7KDOOXV olive-greenish grey; thin, with powdery surface, semi-endolithic; photobiont coccoid green alga. Perithecia: completely immersed, only the black ostiole visible, up to 250 m in diameter, with black-brown exciple; asci clavate; ascospores XQLFHOOXODUDW¿UVW¿QDOO\ZLWK±WUDQVYHUVDOVHSWDDQGHYHQWXDOO\ with one additional ± inclined longitudinal septum, 32–35 × 14–15 m, colourless. (FRORJ\ on calcareous rocks on the bottom of an ephemeral stream, drying out during summer.

Polyblastia

93

'LVWULEXWLRQ only known from the type locality, the stream Balea near the village Vulkan in the Southern Carpathians (Romania).

3RO\EODVWLDSHPLQRVD1\ODQGHU $=DKOEUXFNQHU¿J53) Basionym: Verrucaria peminosa Nylander 1866. Synonym: Polyblastia anziana f. peminosa Servít 1954. 7KDOOXV reddish-grey, greenish when wet; 200 m high, rimose to areolate; paraplectenchymatous, cortex 15–20 m high, hyphal cells in cortex and algal layer mostly vertically, in the medulla horizontally arranged; photobiont coccoid green alga, with dispersed cells, rounded, 4–8 m in diameter, partly elongated, 9 × 3–4 m. Perithecia: immersed, 1–2 per areole, exciple 200– 300 × 300 m, involucrellum stretching down to the thallus base, blackishbrown near the apex, pale brown below; asci 80–90 × 26–25 m; ascospores mostly unicellular but partly with 2–5 transversal septa or muriform, colourless, 22–32 × 10–16 m. (FRORJ\ amphibious on periodically inundated siliceous rocks. 'LVWULEXWLRQ only known from a few localities in the Sudetes and the Carpathians (Poland: Karkonosze/Giant Mountains, Romania: Transylvania). Also reported from Scandinavia. 6LPLODU VSHFLHV Because of the high variability of the ascospore septation among specimens of Polyblastia peminosa, WKH LGHQWL¿FDWLRQ LV YHU\ GLI¿FXOW Eitner (in Zschacke 1933) suggested that one of his specimens from a stream between “Kamme” and the locality “Kleine Teiche” may represent a mechanical

Fig. 53. Polyblastia peminosa: ×19.

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hybrid of Verrucaria laevata (?=Verrucaria praetermissa) and Sporodictyon cruentum. Zschacke (1933) however found unicellular, 2–4 celled as well as muriform ascospores in a single perithecium of another specimen. These earlier observations give an impression of the high variability of ascospore septation in the material.

3RO\EODVWLDULYDOLV$UQROG =VFKDFNH¿J54) Basionym: Thelidium rivale Arnold 1871. 7KDOOXV yellowish (clay-like) whitish to pale-grey; semi-endolithic, continuous, cortex lacking; algal layer up to 130 m high, prosoplectenchymatous, with oil-cells in central to lower parts, fungal cells in the algal layer 4–4.5 m wide; hyphal cells in the lowest parts generally smaller (2–3 m wide), oilcells larger, up to 10 m and partly aggregated, photobiont coccoid green alga. Perithecia: predominantly immersed, only uppermost parts raised above the thallus, up to 600 m in diameter; exciple black, 450 × 400 m; involucrellum distinct (50–75 m thick) and reaching far below but not laterally protruding into the thallus; asci 8-spored, clavate; ascospores with 3–6 transversal and some additional longitudinal septa, colourless, 48–76 × 18– 30 m. (FRORJ\ amphibious on calcareous or lime incrusted rocks. 'LVWULEXWLRQ only known from a few localities in Tyrol, Austria.

Fig. 54. Polyblastia rivalis: ×14.

Porina

95

Porina Müller Argoviensis 1883 nom. cons. Important references: Hafellner & Kalb (1995), Keissler (1937), Purvis et al. (1992), Wirth (1995). 1a At least some ascospores with longitudinal septa ..................................... 2 E Ascospores with transversal septa only ..................................................... 3 2a Ascospores with 5 (6) transversal and 1–3 longitudinal septa ......................... ................................................................................................. P. interjungens E Ascospores with 7 transversal and 1–2 longitudinal septa ...... P. guentheri 3a Perithecia pinkish, reddish or orange, at least at their tips ........................ 4 E Perithecia black throughout ...................................................................... 5 4a Ascospores 3-septate ............................................................... 3OHFWLVVLPD E Ascospores 6–7-septate ............................................................ 3DKOHVLDQD 5a Ascospores 3-septate ............................................................... 3FKORURWLFD E Ascospores 4–7-septate ............................................................................. 6 6a Ascospores 5–6 Pm wide, perithecia < 700 Pm ...................... P. guentheri E Ascospores up to 10 Pm wide, perithecia up to 800 Pm ............. P. grandis

3RULQDDKOHVLDQD .|UEHU =DKOEUXFNQHU¿J55) Basionym: Segestrella ahlesiana Körber 1865. Selected synonyms: Segestria septemseptata Hepp ex Zwackh 1862, Porina septemseptata (Hepp ex Zwackh) Swinscow 1962. 7KDOOXV pale olive-brown, greenish-grey, pale-yellowish; a continuous crust or irregularly cracked; photobiont a yellowish-orange coloured green alga (Trentepohlia). Perithecia: VHPLLPPHUVHGDW¿UVWPRVWO\FRYHUHGE\DWKDOOXVPDQWOH

Fig. 55. Porina ahlesiana: ×24.

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±—PLQGLDPHWHUDSH[¿UVWUHGGLVKHYHQWXDOO\EODFNLVKLQYROXFUHOOXP yellow, pink to reddish, reddish colour intensifying when wetted; asci 90 × 15–18 m, ascospores (5) 7-septate, 30–70 × 6–15 m. (FRORJ\ at shady, humid sites, amphibious in the splash water zone on siliceous rocks in streams and small rivers. 'LVWULEXWLRQ very rare, old records from localities with oceanic climate in Germany (Heidelberg, Pforzheim), also recorded from Austria (Spitz a.d. Donau), Great Britain and Scandinavia. 6LPLODUVSHFLHV Porina lectissima has smaller, 18–30 (40) m, 3 (6)-septate ascospores.

3RULQDFKORURWLFD $FKDULXV 0OOHU$UJRYLHQVLV¿J56) Basionym: Verrucaria chlorotica Acharius 1810. Synonyms: Sagedia chlorotica (Acharius) A. Massalongo 1852, Pyrenula chlorotica (Acharius) Trevisan 1853, Segestrella chlorotica (Acharius) Branth & Rostrup 1869, Segestria chlorotica (Acharius) Hellbom 1870, Arthopyrenia chlorotica (Acharius) H. Olivier 1884, Pseudosagedia chlorotica (Acharius) Hafellner & Kalb 1995, Trichothelium chloroticum (Acharius) R.C. Harris 1995. 7KDOOXV brownish-olive to blackish-green, sometimes bright-green translucent when wet; a thin crust with smooth to granular surface; photobiont a yellowishorange coloured green alga (Trentepohlia). Perithecia: semi-immersed to sessile, 200–300 m in diameter, at least upper part naked and shiny black; involucrellum reaching the thallus base, black; exciple brown-black in outer part, centre ± colourless; ascospores 3-septate, 15–32 × 4–6 m.

Fig. 56. Porina chlorotica: ×19.

Porina

97

(FRORJ\ a terrestrial species on siliceous substrata at shady and humid sites, but also in the splash zone of running water bodies or in small upstream parts which are inundated only for short periods. Often a pioneer species colonizing even small pebbles, but also persistent on old large boulders. 'LVWULEXWLRQ widespread and frequent at terrestrial sites, amphibious populations much rarer but locally frequent. 6LPLODUVSHFLHV Porina aenea is an epiphytic species without greenish colour, smaller perithecia (100–300 m) and ascospores 13–17 (24) m. The exciple in P. aenea is ± uniformly colourless except eventually present brownish inclusions.

Porina grandis .|UEHU =DKOEUXFNQHU¿J 57) Basionym: Sagedia grandis Körber 1863. Synonyms: Porina guentheri var. grandis (Körber) Swinscow 1962, Pseudosagedia grandis (Körber) Hafellner & Kalb 1995. 7KDOOXV olive-greenish, pale ash-grey to blackish, often with purple or reddish WLQJHWKLQFRQWLQXRXVWR¿QHO\ULPRVHSKRWRELRQWD\HOORZLVKRUDQJHFRORXUHG green alga (Trentepohlia). Perithecia: semi-immersed or at least basal parts covered by a thallus mantle, apex naked, 500–800 m in diameter; involucrellum black, purple to violet, especially in lower parts; exciple colourless to straw-coloured; asci 100–250 m; ascospores 5–7 septate, 25–50 × 6–10 m. Pycnidia: rare in Central European material, black; microconidia 4.5 × 0.7–1.0 m. (FRORJ\ on stable siliceous boulders in the splash water zone at or in shaded streams.

Fig. 57. Porina grandis: ×20.

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'LVWULEXWLRQ very rare in high mountain ranges (Black Forest, Germany; Giant Mountains, Poland & Czech Republic). 6LPLODUVSHFLHV Porina guentheri has narrower ascospores (up to 6 m) and perithecia 400–700 m in diameter. Whether Porina grandis should be separated from P. grandis at species rank or merely represents a morph or subspecies of the variable species Porina guentheri s.l. requires further studies. According to Keissler (1937) the perithecial wall of P. guentheri reacts N- or C-, whereas in P. grandis it is N+ or C+ violet. This observation received little attention by other DXWKRUVDQGQHHGVWREHFRQ¿UPHGE\VWXGLHVRQDODUJHUQXPEHU RIVSHFLPHQV

Porina guentheri )ORWRZ =DKOEUXFNQHU¿J58) Basionym: Verrucaria guentheri Flotow 1850. Synonyms: Pseudosagedia guentheri (Flotow) Hafellner & Kalb 1995, Trichothelium guentheri (Flotow) R.C. Harris 1995, Verrucaria koerberi Flotow ex Körber 1855 in syn., Sagedia koerberi (Körber) Körber 1855, Segestria koerberi (Körber) Hellbom 1870, Amphoridium koerberi (Körber) A. Massalongo 1855, Spermatodium koerberi (Körber) Trevisan 1860. 7KDOOXV olive-greenish to pale-grey, often with purple or reddish tinge; thin, FRQWLQXRXVWR¿QHO\ULPRVHSKRWRELRQWD\HOORZLVKRUDQJHFRORXUHGJUHHQDOJD (Trentepohlia). Perithecia: semi-immersed or at least basal parts covered by a thallus mantle, apex naked, 200–700 m in diameter; involucrellum black, purple to violet; exciple colourless to straw-coloured; asci 90–150 × 8 m; ascospores 5–7 septate, 20–45 × 5–6 m. Pycnidia: rare in Central European material, black; microconidia 4.5 × 0.7–1.0 m. (FRORJ\ on stable siliceous boulders in the splash water zone at or in shaded streams.

Fig. 58. Porina guentheri: ×12.

Porina

99

'LVWULEXWLRQ rare species in mountain ranges with usually oceanic climate. 6LPLODU VSHFLHV Porina grandis has broader ascospores (up to 10 m), larger asci (150–250 m) and perithecia up to 800 m in diameter. Whether this taxon should be treated as a separate species or as a subspecies of P. guentheri requires further consideration. Porina interjungens has constantly 5 transversal septa (eventually with additional longitudinal septa) and slightly smaller ascospores than P. guentheri.

Porina interjungens (Nylander) Zahlbruckner 1922 Basionym: Verrucaria interjungens Nylander 1872. Synonyms: Pseudosagedia interjungens (Nylander) Hafellner & Kalb 1995, Verrucaria interseptula Nylander 1881, Porina interseptula (Nylander) A.L. Smith 1911. 7KDOOXV ROLYHJUHHQLVK WR SDOHJUH\ WKLQ FRQWLQXRXV WR ¿QHO\ ULPRVH SKRWRbiont a yellowish-orange coloured green alga (Trentepohlia). Perithecia: up to 700 m in diameter, semi-immersed or at least basal parts covered by a thallus mantle, apex naked; involucrellum black, purple to violet; exciple colourless to straw-coloured; ascospores 5-septate, eventually with 1–3 additional longitudinal septa, 20–25 (30) × 6–10 m. (FRORJ\ amphibious in the splash water zone of streams on siliceous rocks in exposed to shaded sites. 'LVWULEXWLRQ mainly in nothern countries (Scandinavia, British Isles), only historic records from the Harz Mountains (Saxony-Anhalt, Scholz et al. 2004). 6LPLODUVSHFLHV Porina guentheri is very similar in thallus and perithecia morphology, but the ascospores are usually larger (up to 45 m) with up to seven transversal but no longitudinal septa.

3RULQDOHFWLVVLPD )ULHV =DKOEUXFNQHU¿J 59) Basionym: Segestria lectissima Fries 1825. Synonyms: Sagedia lectissima (Fries) Hepp 1860, Verrucaria lectissima (Fries) Nylander 1853, Arthopyrenia lectissima (Fries) H. Olivier 1884, Sphaeromphale lectissima (Fries) Flotow 1839, Verrucaria erysiboda Taylor 1836, Verrucaria rubiginosa Taylor 1836, Verrucaria irrigua Taylor 1836 non Zschacke 1927. 7KDOOXV olive-greenish, olive-brown, brownish-orange, yellowish-brown; rather thick, continuous crust or irregularly cracked; photobiont a yellowish-orange coloured green alga (Trentepohlia). Perithecia: semi-immersed, mostly enveloped by a thallus mantle, 400–500 m; involucrellum pink to reddish, reddish colour intensifying when wetted; asci 70–100 × 5–8 m; ascospores 3 (6)-septate, 18–30 (40) × 4–8 m. Pycnidia rounded, yellowish-red, to brown; pycnospores 3–4.5 × 0.5–1 m.

100

.H\VWRWKHVSHFLHVDQGVSHFLHVSUR¿OHV

Fig. 59. Porina lectissima: ×17. (FRORJ\ best developed in shady seepage tracks, but also found on stable siliceous boulders in the splash water zone along streams, tolerating limited periods of submersion in clean and cool upland watercourses. 'LVWULEXWLRQ scattered in mountain ranges below the timber line, locally abundant but missing in large areas. 6LPLODU VSHFLHV Porina ahlesiana has larger (30–50 m), (6) 7-septate ascospores.

Porocyphus Körber 1855 Important references: Henssen (1963a, 1974), Jørgensen (2007b). Notes: In some species of Porocyphus the cyanobacterial photobiont largely reWDLQVLWV¿ODPHQWRXVVKDSHZKHUHDVLQRWKHUVSHFLHVWKHSKRWRELRQWDSSHDUVWREH VLQJOHFHOOHG$FFRUGLQJWR:LUWK EDVDOKHWHURF\WHVDUHGLI¿FXOWWRREVHUYH but usually present in the two species treated here. Key to the species: 1a 7KDOOXVDUHRODWHDUHROHVÀDWDQGVPRRWKWRZDUW\RUJUDQXORVHXVXDOO\ZLWK abundant vertical outgrowths, outgrowths 50–90 m thick, 100–300 (400) m high, excip. prop. 15–25 (30) m thick, usually remaining pale ... P. coccodes E Thallus granulose to irregularly areolate, areoles never smooth, usually warty to granulose, sometimes dissolved into minute coralloid outgrowths, outgrowths 20–50 m thick, 50–200 (300) m high, excip. prop. (20) 35–55 (65) m thick, usually distinctly brownish coloured ............................... P. rehmicus

Porocyphus

101

Porocyphus coccodes )ORWRZ .|UEHU¿J 60) Basionym: Collema coccodes Flotow 1850. Selected synonyms: Collema pyrenopsoides Nylander (Nylander) Nylander 1855, Collemopsis lecanopsoides (Nylander) Crombie 1874, Gabura pyrenopsoides (Nylander) O. Kuntze 1891, Homopsella aggregatula Nylander 1887, Lecanora pyrenopsoides Nylander 1853, Porocyphus areolatus (Flotow) Körber 1855, Porocyphus cataractarum Körber 1865, Porocyphus furfurellus (Nylander) Forssell 1885, Porocyphus vivariensis Couderc in Harmand 1912, Psorotichia coccodes (Flotow) Arnold 1897, Psorotichia lecanopsoides (Nylander) Boistel 1903, Psorotichia pyrenopsoides (Nylander) Forssell 1885, Pyrenopsis lecanopsoides Nylander 1863. 7KDOOXV black to blackish-green-olive, crustose-areolate, areoles ± angulate to irregular, sometimes granulose, 0.25–1.5 (2.5) mm wide, 0.2–0.5 (1) mm thick, surface usually uneven, rarely smooth, warty to granulose and usually with vertical outgrowths or proliferations and thus coralloid, outgrowths ± cylindrical, 0.1–0.3 (0.4) mm high, 50–90 m thick; attached by rhizohyphae originating from sometimes indistinct paraplectenchymatous basal layer; thallus anatomy paraplectenchymatous, hyphal cells small, roundish to ± angulate, densely reticuODWHWRSDUDSOHFWHQFK\PDWRXVSKRWRELRQWD¿ODPHQWRXVF\DQREDFWHULXP5LYXODriaceae), but trichomes often strongly splitted and appearing single-celled, basal heterocytes visible or not. Apothecia: pycnoascocarps, sessile and laminal on areoles or terminal on vertical outgrowths, 1–3 (5) per areole, up to 0.3 (0.4) mm wide, thalline margin thin, entire, 30–40 m wide, sometimes with minute proOLIHUDWLRQVHYHQWXDOO\UHFHGLQJGLVFDW¿UVWSXQFWLIRUPDQGGHSUHVVHGODWHURSHQ DQG “ ÀDW GDUN UHGGLVK EURZQ excip. prop. distinct only in mature apothecia when the thalline margin recedes, apically yellowish brown otherwise hyaline, often visible as a light ring when mature apothecia are moistened, basally 10–15 m thick, otherwise 15–25 (30) m wide, apically usually distinctly widened; remnants of conidiophores sometimes present between hymenium and exciple (i.e. former pycnidial wall) in juvenile apothecia; epihymenium faintly reddish-brown; hymenium (100) 110–130 m high, hyaline, usually I+ blue; paraphyses thin, 1.5–2 m thick, indistinctly septate, sparingly branched and anastomosing, apical cells slightly thickened, to 3 m wide; asci 8-spored; ascospores simple, hyaline, broad ellipsoid, (9) 11–17.5 (18) × 7–12 (13) m. Pycnidia: immersed to slightly elevated, globose to broad pyriform, 0.075–0.125 mm in size; pycnospores short cylindrical to almost globose, 2–2.5 × 1–1.5 m. (FRORJ\ on usually exposed, acidic to slightly calciferous rock faces moistened by seeping water and semi-aquatic along streams, rocky lake shores and water falls, also known from fresh water tidal zones along the River Elbe, occasionally on old walls etc. in urban places. 'LVWULEXWLRQ from lowland into alpine regions and widespread in the northern hemisphere but only rarely recorded. A form deviating from typical Porocyphus coccodes growing ± submerged in a shallow stream in the Ardennes was reported by Sérusiaux et al. (1999). According to the brief description this species belongs into a different genus.

102

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Fig. 60. Porocyphus coccodes: thallus areolate, surface with coarse, vertical proliferations and some apothecia; ×19. 6LPLODUVSHFLHV According to Henssen (1974), Porocyphus coccodes LVGLI¿FXOW to distinguish from P. kenmorensisHVSHFLDOO\ZKHQJURZLQJXQGHUWKHLQÀXHQFH of running water. In fact, Porocyphus kenmorensis LVFRQ¿QHGWRURFNVDORQJWKH margins of rivers and lakes. It is known so far from the British Isles and Finland but should be expected at suitable habitats in Central Europe. Porocyphus rehmicus is very similar to P. coccodes DQG SUREDEO\ RQO\ D PRGL¿FDWLRQ RI WKH ODWWHU (Henssen 1963a).

Porocyphus rehmicus $0DVVDORQJR =DKOEUXFNQHU¿J 61) Basionym: Psorotichia rehmica A. Massalongo 1856. Selected synonyms: Collemopsis riparia (Arnold) Trevisan 1880, Porocyphus byssoides Hepp 1857, Porocyphus globulosus Couderc in Crozals 1908, Porocyphus riparius (Arnold) Körber 1865, Psoropsis rehmica (A. Massalongo) Nylander in Zwackh 1883, Psorotichia geophila Hy 1893, Psorotichia globulosa A. Massalongo in Arnold 1885 nom. nud., Psorotichia riparia Arnold 1859, Psorotichia rufescens Hy 1893, Pyrenopsis riparius (Arnold) Nylander 1859. 7KDOOXV blackish to dark-greenish, crustose, granulose to irregularly areolate, areoles 0.3–1 (1.5) mm wide, 0.2–0.5 (1) mm thick, surface usually uneven, warty to granulose, never smooth, often areoles becoming dissolved into ± erect, coralloid granules, 50–200 m long, 20–50 m thick, attached to the substrate by rhizohyphae; thallus anatomy with irregularly arranged roundish mycobiont cells, appearing ± paraplectenchymatous; photobiont (Rivulariaceae) usually with short and strongly splitted trichomes but basal heterocytes sometimes visible. Apothecia: pycnoascocarps, semi-immersed to soon sessile, 1–3 (8) per DUHROHDW¿UVWDSSHDULQJJORERVHHYHQWXDOO\GLVFVKDSHGWRPPZLGHWKDOOLQe margin entire, 20–50 m wide, eventually receding and brownish proper margin

Porocyphus

103

Fig. 61. Porocyphus rehmicus: thallus granulose-areolate, partly with minute, very narrow proliferations and some apothecia; ×19. EHFRPLQJGLVWLQFWGLVFDW¿UVWSXQFWLIRUPDQGGHSUHVVHGODWHURSHQDQG“ÀDW rather pale to dark reddish brown; excip. prop. basally 10–20 m wide, apically DW¿UVWUDWKHULQGLVWLQFWEXWHYHQWXDOO\WKLFNHQHGDQG± —PZLGHDSLcally yellowish brown, otherwise hyaline, often visible as a pale ring around the disc when mature apothecia are moistened, in juvenile apothecia remnants of conidiophores often present between hymenium and exciple (i.e. former pycnidial wall); epihymenium pale or faintly reddish brown; hymenium 90–130 m high, hyaline, I+ blue; paraphyses very thin, indistinctly septate, usually unbranched, apical cells slightly thickened, to 3 m wide; asci 8-spored, ascospores simple, hyaline, globose to broad ellipsoid, 11–16.5 × (5.5) 7–13 m. Pycnidia: immersed to slightly elevated, globose to broad pyriform, 0.01–0.125 mm in size; pycnospores short cylindrical, 2.5 × 1 m. (FRORJ\ on exposed, usually calciferous rock, occasionally inundated on boulders along rivers, also on old walls, predominantly in nutrient enriched sites; usually considered to prefer drier places than the preceding species but also known from semi-aquatic habitats along larger rivers such the Danube as well as from historical collections along the shore of Lake Zurich. 'LVWULEXWLRQ from lowland to mountainous regions, according to historical collections probably fairly widespread in Central Europe but nowadays rarely recorded. Records from outside Europe should be treated cautiously. 6LPLODUVSHFLHV Porocyphus rehmicus LVSUREDEO\MXVWDPRGL¿FDWLRQRIP. coccodes (Henssen 1963a). Porocyphus leptogiella (Nylander) L.T. Ellis from the British Isles is very similar to P. rehmicus. According to Ellis (1981) it is distinguished from P. rehmicus only by the slightly smaller ascospores and vague differences in the shape of the paraphyses. Henssen (1963a) placed Porocyphus byssoides Hepp and P. riparius (Arnold) Körber in synonymy to P. rehmicus. Van

104

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den Boom et al. (1998) realized that P. byssoides might be separated from P. rehmicus s. str. on account of the smaller ascospores. The spore size given in the present treatment matches that of P. byssoides given by van den Boom et al. (1998) and given for P. rehmicus s.l. by Henssen (1963a). Like Porocyphus byssoides, P. riparius was described from a periodically inundated locality along the Danube in Lower Bavaria and the type material is heavily incrusted with silt. Porocyphus riparius is somewhat deviating from characteristic specimens of P. rehmicus. Further taxonomic as well ecological studies are needed to clarify the status of the taxa involved. Pterygiopsis neglectaVXSHU¿FLDOO\UHVHPEOHVPorocyphus rehmicus and P. coccodes. It differs, however, in the coccoid cyanobacterial photobiont, the lack of pycnoascocarps, a thin proper exciple, a persisting thallus margin and eventually umbonate apothecial discs.

Porpidia Körber 1855 Notes: Many PorpidiaVSHFLHVDUHW\SLFDOULSDULDQOLFKHQV7KH\EHQH¿WIURPWKH higher moisture in the vicinity of watercourses and at the bottom of small canyons but usually avoid longer inundation. An exception is P. hydrophila which is restricted to amphibious habitats and tolerates longer periods of submersion. Porpidia soredizodes, although also widespread at exclusively terrestrial sites is described here in detail, because this species is a very characteristic component of the splash water communities in certain mountain ranges. On a small scale, the occurrence of P. soredizodes often shows an upper as well as a lower boundary depending on the limited submersion resistance of the species and the preference of high air humidity and splash water supply (Ried 1960a, 1960b, Wirth 1972). In shady habitats P. glaucophaea and P. albocaerulescens can be found frequently in the splash water zone. Determination of sterile specimen usually requires t.l.c. for reliable results. Important references: Fryday (2005), Gilbert (2000), Ried (1960a, b), Wirth (1995). Key to the species: 1a Thallus with soralia ................................................................................... 2 E Thallus lacking soralia .............................................................................. 7 2a Thallus lacking apothecia ......................................................................... 3 E Thallus with apothecia .............................................................................. 6 3a Thallus mostly orange, thick ........................................................................ .......................... 3ÀDYRFRHUXOHVFHQV (Hornemann) Hertel & Schwab 1984 E Thallus not orange, thin or thick ............................................................... 4 4a Thallus white, never with bluish tinge, containing glaucophaeic acid ......... ........................................... P. glaucophaea (Körber) Hertel & Knoph 1984 E Thallus grey, with or without bluish tinge, lacking glaucophaeic acid ...... 5 5a 7KDOOXVFRQVSLFXRXVVRUDOLDJUH\,“EOXHFRQWDLQLQJFRQÀXHQWLFDFLGUHstricted to terrestrial zone, but often on boulders bordering small streams .... ............................................... P. tuberculosa (Smith) Hertel & Knoph 1984 E Thallus thin, often inconspicuous, soralia distinct, often with bluish tinge, ,FRQWDLQLQJVWLFWLFDFLGWHUUHVWULDOWRÀXYLDOPHVLF]RQHV .... 3VRUHGL]RGHV

Porpidia

105

6a Apothecial disc pruinose, thallus thick, soralia often variously shaped, usually along thallus cracks, thallus K-, P-, containing glaucophaeic acid ........ ........................................... P. glaucophaea (Körber) Hertel & Knoph 1984 E Apothecial disc epruinose, thallus thin, soralia immersed, rounded, thallus K+ yellowish, P+ orange, containing stictic acid ................. 3VRUHGL]RGHV 7a Epihymenium bright emerald-green, apothecial disc epruinose ... 3K\GURSKLOD E Epihymenium not bright emerald-green, apothecial disc pruinose or epruinose .......................................................................................................... 8 8a Apothecia whitish pruinose, r immersed, thallus whitish cream-coloured or whitish-grey ............... P. albocaerulescens (Wulfen) Hertel & Knoph 1984 E Apothecia epruinose, sessile, thallus whitish or grey to brown or ochre ... 9 9a Apothecia to 0.55–1.6 mm in diameter, apothecial margin thin (0.07–0.17 mm), ascospores 12–20 × 6–10 Pm, common from lowland to alpine areas ............. ............................................ P. crustulata (Acharius) Hertel & Knoph 1984 E Apothecia 1.0-3.8 mm in diameter, apothecial margin thick (0.13–0.25 mm), ascospores 17–25 (30) × 7–12 (14) Pm, usually in montane to alpine areas ................................... P. macrocarpa (de Candolle) Hertel & Schwab 1984

3RUSLGLDK\GURSKLOD )ULHV +HUWHO 6FKZDE¿J 62) Basionym: Lecidea hydrophila Fries 1822. Selected synonyms: Lecidea macrocarpa var. hydrophila (Fries) Th. Fries 1874, Lecidea convexa var. hydrophila (Fries) Th. Fries 1874, Huilia hydrophila (Fries) Hertel 1975, Haplocarpon hydrophilum (Fries) V. Wirth 1972. 7KDOOXV whitish, whitish cream-coloured, light-grey, often partly with orange or UXVW\WLQJHWKLQFRQWLQXRXVWR¿QHO\ULPRVHQROLFKHQSURGXFWVGHWHFWHGE\WOF photobiont coccoid green alga. Apothecia: frequent, sessile, 0.5–1.5 (2.5) mm LQ GLDPHWHU ZLWK GLVWLQFW DQG SURPLQHQW H[FLSOH DSRWKHFLDO GLVF ÀDW RU VOLJKWO\

Fig. 62. Porpidia hydrophila: ×15.

106

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concave when young, later becoming convex; epithecium 12–25 m high, bright emerald-green; hymenium 80–120 (140) m high, upper part bluish-green; periphyses tightly glued; ascospores (14) 17–25 × 5.5–10 m. Pycnidia: rare in Central European populations, but frequent in thalli from northwestern Europe. (FRORJ\ on stable rocks in and at streams with neutral to acidic soft water, usually in the splash water zone but also tolerating short periods of complete submersion. 'LVWULEXWLRQ mostly in oceanic regions at low elevations (often < 700 m); a rare species in Central Europe, populations in the Eifel and Harz Mountains have much decreased and are on the verge of extinction. 6LPLODUVSHFLHV No other species of Porpidia has a bright emerald-green epihymenium. Lecidella stigmatea and L. carpathica are terrestrial species and also have a blue-green or green epithecium. However, the ascospores are shorter (11– 17 m), the paraphyses become easily separated when sections are gently pressed, and the thallus is usually K+ yellow. The outermost parts of the exciple are black, while the exciple is transparent or red-brownish towards the centre.

3RUSLGLDVRUHGL]RGHV/DP\H[1\ODQGHU -5/DXQGRQ¿J 63) Basionym: Lecidea crustulata var. soredizodes Lamy ex Nylander 1883. Synonyms: Lecidea crustulata ssp. soredizodes (Lamy ex Nylander) anon., Haplocarpon soredizodes (Lamy ex Nylander) V. Wirth 1972, Huilia soredizodes (Lamy ex Nylander) Hertel 1980. 7KDOOXV JUH\LVK EXW RIWHQ ZLWK DQ RUDQJH WLQJH WKLQ FRQWLQXRXV RU ¿QHO\ ULmose; soralia ± circular, whitish to greenish, but often also with bluish tinge, immersed to slightly convex, P+ orange, K+ yellowish, I-, stictic acid detected by

Fig. 63. Porpidia soredizodes: ×19.

Pseudarthopyrenia

107

t.l.c.; photobiont coccoid green alga (Asterochloris ( VSYHU¿HGE\FXOWXUHGLVRlates and ITS-sequence data). Apothecia: very rare, sessile, up to 1 mm in diaPHWHUDSRWKHFLDOGLVFÀDWHSUXLQRVHRIWHQEURZQLVKZKHQZHWK\PHQLXP± 150 m high. Ascospores: 12.5–20 × 6.5–10 m. (FRORJ\ a terrestrial species with wide ecological amplitude and in some regions a characteristic element of splash water communities, especially on well buffered substrata (e.g. basalt); rather shade tolerant. 'LVWULEXWLRQ amphibious populations scattered all over Central Europe but more often in regions with neutral to slightly basic siliceous bedrock. 6LPLODUVSHFLHV Porpidia tuberculosa has a thicker thallus with a I+ blue medulla and irregular soralia lacking a bluish tinge. This species is often found close to watercourses favoured by the higher air humidity, but it is sensitive to inundation and occurs usually well above the splash water zone. Other sorediate species of Porpidia have thicker thalli with a whitish colour and a deviating chemistry.

Pseudarthopyrenia Keissler 1935 Note: This a poorly known genus which is in need of revision. No specimens have been traced by the authors and the details for the species treated here are based on the descriptions given by Servít (1955). Important references: Servít 1955. Key to the species: 1a Ascospores 25–30 m long, asci clavate, thallus with thin black basal layer (up to 25 m) .......................................................................... 3ED\HULDQD E Ascospores 14–16 m long, asci cylindric, thallus without black basal layer ................................................................................................... 3ULYXODULV

3VHXGDUWKRS\UHQLDED\HULDQD Servít 1955 7KDOOXVGDUNROLYHHSLOLWKLFFRQWLQXRVWR¿QHO\ULPRVHVXUIDFHXQHYHQYHUUXcose; cortex absent; hyphae in all parts of the thallus paraplectenchymatous; algal layer 30–40 m high, photobiont a bluish-green coloured cyanobacterium , 3–4 m in diameter; medullary layer (algal free lower part of thallus) up to 20 m high, colourless to brown; black basal layer up to 25 m. Perithecia: hemispherical in shape, immersed to semi-immersed in the thallus; up to 180 m in diameter, exposed apical part without thalline mantle 100–150 m wide; exciple 90 m in diameter, in lower parts colourless and 12 m thick, with elongated cells, 4 × 0.7 m; upper part brown-black and up to 30 m thick, exciple and hymenium I-; paraphyses sparingly branched, with cell lumina 0.8 m wide; asci clavate, 70–75 × 20–22 m, I+ brown-red, apical wall 5–7 m, ascospores in up to three rows, colourless, uniseptate, upper cell slightly wider than the lower one; slightly constricted at the septum, 25–30 × 10–12 m, cell wall 0.5 m thick.

108

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(FRORJ\ on gneiss in a river. 'LVWULEXWLRQRQO\NQRZQIURPWKHW\SHORFDOLW\LQWKH&]HFK5HSXEOLF&KRWƟERĜ  at 500 m. 6LPLODUVSHFLHV The thallus of Collemopsidium angermannicum is very similar, but the perithecia differ in the richly branched, anastomosing paraphyses, larger asci (100 m long) and smaller ascospores (17–26 m) (Nordin 2002b). Servít (1955) points to the similarity with the parasitic species Didymella brunii, which is said to grow on the thallus of Verrucaria species with green algae and furthermore differs from P. bayeriana by an olive-greenish tinge of the exciple. According to Servít there is no indication that the photobiont cells in the thallus of P. bayeriana belong to a host thallus. The thallus of the type material appeared to be in good condition and he concluded, that P. bayerina is a lichenized fungus.

3VHXGDUWKRS\UHQLDULYXODULV Servít 1955 7KDOOXVGDUNROLYHHSLOLWKLFFRQWLQXRXVRUSDUWO\YHU\¿QHO\ULPRVHJHODWLQRXV when wet, surface uneven; cortex hardly discernable; algal layer up to 40 m high, paraplectenchymatous, with fungal cell of 1–2 m in diameter, photobiont a cyanobacterium, cells in small aggregated groups to uniformly dispersed, 1–3 m in diameter or elongated and 4 × 2 m in size. Perithecia: hemispherical, 80–150 m in diameter, XSSHUSDUWVQDNHGÀDQNVFRYHUHGE\DWKLQWKDOOLQHPDQWOHH[FLSOH 80–150 m in diameter, basal part almost colourless to brownish, 12 m thick, with elongated cells (3–5 × 0.7 m), upper parts brown to black, up to 40 m thick; paraphyses sparingly branched, with cell lumina 0.7 m wide; exciple and hymenium I-; asci cylindrical 50–60 × 12–15 m, I+, dark red; apical wall thin, 1 m thick; ascospores in 1–2 rows, most ascospores uniseptate with the upper cell slightly wider than the lower one and slightly constricted at the septum, some ascospores unicellular, 14–16 × 4–6 m, cell wall 0.3 m. (FRORJ\ on schist in a river. 'LVWULEXWLRQRQO\NQRZQIURPWKHW\SHORFDOLW\LQWKH&]HFK5HSXEOLF5XGRKRĜt (U]JHELUJH3ĜtVHþQLFH DWP 6LPLODUVSHFLHV The thallus of Collemopsidium angermannicum is very similar, but the perithecia differ in the richly branched, anastomosing paraphyses, much larger asci (100 m long) and slightly larger ascospores (17–26 m). Servít (1955) points to the similarity of P. rivularis with the parasitic species Didymella martinatiana which has narrower ascospore cells.

Pterygiopsis

109

Pterygiopsis Vainio 1890 Important references: +HQVVHQ µ¶  +HQVVHQ  -¡UJHQVHQ   Jørgensen (1990, 2007b). Key to the species: 1a Thallus forming an evenly spreading rimose to areolate crust, apothecia 0.3–0.5 (0.6) mm wide, disc usually expanded, black, thallus margin to 100 m wide ............................................................................. 3FRQFRUGDWXOD E 7KDOOXVDW¿UVWWKLQDQGULPRVHVRRQEHFRPLQJLUUHJXODUO\DUHRODWHHYHQWXDOO\ dissolved into irregular granules, apothecia 0.2–0.3 (0.4) mm wide, disc very small, usually < 100 m wide, dark reddish-brown, eventually umbonate and then pale-brownish, thallus margin usually 40–60 m wide ........ 3QHJOHFWD

3WHU\JLRSVLVFRQFRUGDWXOD (Nylander) P.M. Jørgensen 2007 ¿JV 164, 171) Basionym: Pyrenopsis concordatula Nylander 1875. Synonyms: ?Collemopsis assimulans Nylander 1876, Collemopsis coracodiza Nylander 1878, ?Psorotichia assimulans (Nylander) Forssell 1885, Psorotichia coracodiza (Nylander) Forssell 1885, Pterygiopsis coracodiza (Nylander) Henssen 1990, Pyrenopsis amphibia unpubl. herb. name, ?Pyrenopsis assimulans (Nylander) Blomberg & Forssell 1880. 7KDOOXV dark blackish-brown, dull, crustose, soon rimose to rimose-areolate, HYHQWXDOO\DUHRODWHPDUJLQVQRWHI¿JXUDWHDUHROHVLUUHJXODUO\ VKDSHGZLWKVKDUS HGJHV±PPZLGH±PPWKLFNSODQHVXUIDFH¿QHO\ZDUW\JLYLQJ the thallus a furfuraceous appearance; attached by gelatinous basal layer; thallus anatomy paraplectenchymatous with the photobionts arranged in vertical rows, hyphal cells small, roundish to broad ellipsoid or angulate, c. 3–5 m; photobiont a single-celled cyanobacterium, cells small, 8–10 (11) m, gelatinous sheath 2–2.5 m thick, yellowish brown¿J . Apothecia: semi-immersed to sessile, VLQJOH¿OOLQJWKHHQWLUHDUHROH± PPZLGHWKDOOLQHPDUJLQHQWLUHÀDW to slightly prominent, persisting, c. —P WKLFN GLVF DW ¿UVW SXQFWLIRUP EXW VRRQRSHQHGEODFNVOLJKWO\GHSUHVVHGRUÀDWROGGLVFVPD\EHFRPHURXJKDQG umbonate; excip. prop. thin, 10–15 m thick, apically slightly widened; epihymenium dingy yellowish brown, K+ pale reddish purple, N-; hymenium 100 (130) m high, hyaline, I+ blue; paraphyses thin, septate, branched and anastomosing, apical cells slightly thickened, c. 3 × 3–5 m; asci 8-spored, thin walled, ± cylindrical to narrowly clavate, I-; ascospores simple, hyaline, broad ellipsoid to ellipsoid, (8) 10–15 (17) × (6) 7–9 (10) m. Pycnidia: immersed, c. 100–125 m; pycnospores small ellipsoid, 3–4 × 1 m. (FRORJ\ on irrigated, siliceous or schistose rock, in seepage tracks and semiinundated along rocky river banks. 'LVWULEXWLRQ rarely observed in the Flora region and perhaps overlooked or lumped under more common species; from lowland to mountainous regions, reported here from Krkonoše Mts. (Giant Mts.) in Northern Bohemia, also in Brit-

110

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tany, central France, Wales, Scotland, southern Norway, southern and middle Sweden and southern Finland as well as the easternmost foothills of the Pyrenees (Pereira 1992). 6LPLODUVSHFLHV Pterygiopsis lacustris differs from P. concordatula in the thin, continuous crustose thallus, the bluish-green epihymenium which reacts N+ purple and the I- reaction of the hymenium. It grows submerged for most of the year in the lower lichen zone in rocky lake shores in southern Sweden and on the British Isles. The species is easily overlooked and may be expected in similar habitats in continental Europe (Jørgensen 1990).

3WHU\JLRSVLVQHJOHFWD(ULFKVHQ 06FKXOW] 7KVLQHG¿J 64) Basionym: Forssellia neglecta Erichsen 1940. 7KDOOXVEODFNLVKGXOOFUXVWRVHDW¿UVWYHU\WKLQDQGULPRVHVRRQLUUHJXODUO\ DUHRODWHVRPHWLPHVGLVVROYHGLQWRLUUHJXODUJUDQXOHVPDUJLQVQRWHI¿JXUDWHDUHoles 0.2–0.4 (0.5) mm wide, 0.1–0.25 mm thick, surface soon rough and uneven; attached by gelatinous basal layer; thallus anatomy paraplectenchymatous, symbionts arranged in ± distinct vertical rows, hyphal cells small, roundish to broad ellipsoid or angulate, (3) 4–7 m; photobiont a single-celled cyanobacterium, cells small, 7–10 (11) m, gelatinous sheath 2–2.5 m thick, yellowish brown. Apothecia: semi-immersed to sessile, 1–3 per areole, 0.2–0.3 (0.4) mm wide, thalline margin entire, persisting, slightly prominent, ±—PZLGHGLVFDW¿UVW punctiform, later opened but remaining very narrow, usually < 100 m wide, GDUN UHGGLVK EURZQ GHSUHVVHG DQG ÀDW HYHQWXDOO\ XPERQDWH DQG WKHQ SDOH brownish; excip. prop. hyaline, thin, 10 m thick, apically slightly widened; epihymenium pale or slightly yellowish-brown, K-, N-; hymenium 100–125 m

Fig. 64. Pterygiopsis neglecta: thallus granulose-areolate, apothecia with umbonate dicsc and thin thalline margin; ×27.

Pyrenocarpon

111

high, hyaline, I+ blue or I-; paraphyses straight, thin, septate, sparingly branched and anastomosing, apical cells slightly thickened, c. 3 × 3–5 m; asci 8-spored, thin walled, ± cylindrical to narrowly clavate, I-; ascospores simple, hyaline, broad ellipsoid to ellipsoid, (10) 12–16 (17) × 7.5–10 (12) m. Pycnidia: immersed, c. 50–60 m wide; pycnospores small ellipsoid, 2.5–3 × 1.5 m. (FRORJ\ on boulders (slack, granite, limestone, concrete etc.) in the fresh water tidal zone along the River Elbe in and around Hamburg, experiencing regular inundation for approximately 1–3 hours during high tide; usually associated with Staurothele frustulenta, Caloplaca isidiigera and C. subpallida/arenaria. 'LVWULEXWLRQ so far only from the type collection and a few recent collections from the bank of the River Elbe in Hamburg, Wedel and Mojenhörn, but certainly overlooked and to be expected at similar fresh water tidal habitats of the rivers Weser and Ems etc.; possibly also on suitable substrata of inland dikes along larger rivers and lakes (?). Notes: Forssellia neglecta is a forgotten taxon which will be dealt with in detail in a forthcomming publication wherein its new name, Pterygiopsis neglecta, will formally be proposed. Pterygiopsis neglecta can be (and perhaps used to be) mistaken for Porocyphus ssp. However, unlike Porocyphus, the species has a very thin proper exciple, a persisting thalline margin, possesses robust paraphyses and has eventually umbonate apothecial discs. In general, Pterygiopsis neglecta shows the growth habit of species of Psorotichia. However, typical representatives of the latter genus do not have umbonate apothecial discs and usually do not occur in aquatic or amphibious situations. Psorotichia frustulosa was recently observed growing abundantly on exposed revetments of Lake Michigan in Chicago (Illinois, USA). At this locality the lichen experiences occasional inundation. Similar collections exist from the dikes of the IJsselmeer in The Netherlands.

Pyrenocarpon Trevisan 1855 Notes: Pyrenocarpon is a monotypic genus when the unrelated species “Pyrenocarpon montinii$0DVVDORQJR 7UHYLVDQ´LVH[FOXGHG7KHJHQXVKDVDI¿QLWLHV to Psorotichia and perhaps Lemmopsis. Important references: Jørgensen (2007b), Jørgensen & Henssen (1990).

3\UHQRFDUSRQÀRWRZLDQXP+HSS 7UHYLVDQ¿J65) Basionym: 9HUUXFDULD ÀRWRZLDQD Hepp 1853. Synonyms: %LDWRUD ÀRWRZLDQD (Hepp) Th. Fries 1861, &ROOHPRSVLVÀRWRZLDQXV (Hepp) Nylander 1873, Montinia ÀRWRZLDQD (Hepp) A. Massalongo 1855, 3RURF\SKXVÀRWRZLDQXs (Hepp) B. Stein 1879, 3VRURWLFKLD ÀRWRZLDQD (Hepp) Müller Argoviensis 1872, Pyrenocarpus ÀRWRZLDQXV (Hepp) Trevisan 1860, 3\UHQRSVLVÀRWRZLDQD (Hepp) Nylander 1858, 7KHORFKURD ÀRWRZLDQD (Hepp) A. Massalongo 1855, ?Thrombium thelostomum (Acharius ex Harriman) A.L. Smith 1911, ?Pyrenula umbonata Acharius 1810.

112

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Fig. 65. 3\UHQRFDUSRQÀRWRZLDQXP: thallus very thin, small-areolate and with numerous, conical to hemispherical apothecia; ×17. 7KDOOXV brownish olive to rarely almost blackish, crustose, minutely areolate to granulose or effuse especially towards the margin; areoles indistinct or very VPDOO± PPZLGH“DQJXODWHWKLQÀDWRUVOLJKWO\FRQYH[VRPHWLPHV ZLWK VOLJKWO\ OLIWHG PDUJLQV VXUIDFH XVXDOO\ VPRRWK RU ¿QHO\ JUDQXORVH attached by rhizohyphae originating from the lower surface; thallus anatomy paraplectenchymatous, hyphal cells small, roundish to broad ellipsoid or angulate, c. 3–5 m; photobiont (apparently?) a single-celled cyanobacterium, cells to 10 m wide, gelatinous sheath thin, yellowish brown. Apothecia: conical to KHPLVSKHULFDOHYHQWXDOO\GLVFVKDSHGDQGVHVVLOHZLWKEURDGEDVLVVLQJOH¿OOLQJ entire areole, 0.15–0.4 (0.6) mm wide; thallus margin smooth, persisting, not SURPLQHQW ±PP ZLGH EXW DSLFDOO\ WKLQQHU GLVF DW ¿UVW SXQFWLIRUP DQG hardly opened, 0.08–0.15 mm wide, eventually open, 0.25–0.35 (0.4) mm wide, GLVFVOLJKWO\GHSUHVVHGRUÀDWGDUNUHGGLVKEURZQWREODFNLVK ROGGLVFVPD\EHcome rough and umbonate; excip. prop. distinct from the beginning, small-celled subparaplectenchymatous, 10–15 m wide at the margin and at the basis, apically c. 25–35 m wide, hyaline but apically reddish or yellowish brown, slightly paler than the apothecial disc; epihymenium faintly reddish brown; hymenium 125–150 m high, hyaline, I+ faintly blue, sometimes indistinct or bluish colouration rapidly fading; subhymenium extending downwards as small stipe; paraphyses thin, septate, branched and anastomosing, apical cells c. 3 × 5 m; asci 8-spored, very thin walled, ± cylindrical to narrowly clavate, I-; ascospores simple, hyaline, broad ellipsoid to ellipsoid, 10–22.5 × 7–10 m, walls thin. Pycnidia: small, immersed, with pale ostiole, c. 75 m in size; pycnospores c. 3 × 1 m. (FRORJ\ on limestone and various calciferous, rarely acidic rocks along mountain rivers, streams and water falls, periodically to sporadically inundated; in the type collection (Hepp exs. no. 92) and material from the Lech ravine (Bavaria) usually accompanied by Placynthium tantaleum, Leptogium subtorulosum and various pyrenocarpous lichens such as Staurothele solvens s.l..

Pyrenocollema

113

'LVWULEXWLRQ known only from a few, old records in the Alps and Sudetes but perhaps overlooked and to be searched for; also known from Scandinavia, Extremadura (Spain) and Serra da Estrela (Portugal).

3\UHQRFROOHPDReinke 1895 Notes: The circumscription of the genus and species is far from being settled and in need of revision. The thallus is sometimes hard to discern and may contain several different algal taxa. It has been questioned whether these loose associations can be referred to as lichens or simply represent opportunistic algicolous fungi JURZLQJLQDPL[HGELR¿OPZLWKGLIIHUHQWDXWRWURSKV0ROLWRU 'LHGHULFK  Important references: Egan (1987), Grube & Ryan (2002), Harris (1995), Molitor & Diederich (1997), Purvis et al. (1992). Key to the species: 1a Ascospores r oblong, the lower cell not markedly narrowed ..... P. caesium E Ascospores ovoid to ovoid-fusiform, the lower cell markedly narrowed .... 2 2a Ascospores 8–12 m wide, perithecia immersed, only the apex exposed .... ........................................................................................ P. tichothecioides E Ascospores 4–7 m wide, perithecia semi-immersed, upper half exposed .... .................................................................................................... 3VD[LFROD

3\UHQRFROOHPDFDHVLXP (Nylander) R.C. Harris in Egan 1987 Basionym: Verrucaria caesia Nylander 1853. Synonyms: Leiophloea caesia (Nylander) Trevisan 1860, Arthopyrenia caesia (Nylander) Zahlbruckner 1921, Pseudarthopyrenia caesia (Nylander) Keissler 1937. 7KDOOXVJUH\LVKWRUHGGLVKEURZQLPPHUVHGWRVXSHU¿FLDOSKRWRELRQWFHOOV\HOlowish (Trentepohlia) or blue-green (cyanobacteria). Perithecia: semi-immersed; exciple 250–400 m in diameter; hamathecial hyphae richly branched and anastomosing; involucrellum absent; ascospores generally uniseptate but occasionally with additional pseudosepta; both cells of the ascospores of similar shape and size, 20–30 × 9–12 m. (FRORJ\ amphibious on calcareous rocks. 'LVWULEXWLRQ a rare species in freshwater habitats, only known from a few localities in France and Switzerland, but also in Nordic countries. 6LPLODU VSHFLHV Pyrenocollema saxicola has smaller perithecia (100–200 m) and ascospores with a lower cell being distinctly narrowed towards the apex.

114

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3\UHQRFROOHPDVD[LFROD$0DVVDORQJR 5&+DUULV¿J66) Basionym: Arthopyrenia saxicola A. Massalongo 1855. Synonyms: Spermatodium saxicolum (A. Massalongo) Trevisan 1860, Sagedia saxicola (A. Massalongo) Krempelhuber 1861, Verrucaria saxicola (A. Massalongo) Crombie 1876, Leiophloea saxicola (A. Massalongo) Riedl 1969, Pyrenocollema saxicola (A. Massalongo) Coppins 1992, Naetrocymbe saxicola (A. Massalongo) R.C. Harris 1995. Uncertain species: Sagedia massalongiana Hepp 1857, Segestrella massalongiana (Hepp) Beltramini 1858, Segestria massalongiana (Hepp) Trevisan 1860, Spermatodium massalongianum (Hepp) Trevisan 1860, Naetrocymbe massalongiana (Hepp) R.C. Harris 1995. 7KDOOXV whitish-grey, greenish-grey, bluish-grey or reddish-grey; thin or ± endolithic, 60–120 m, semi-endolithic thalli with pyriform oil cells in lower parts, K-, C-; photobiont cells (yellowish-) green, dispersed or in short chains, with thick cell walls (Trentepohlia), occasionally additional cells of other eucaryotic algae or cyanobacteria present, some populations being non-lichenized but algicolous (Molitor & Diederich 1997). Perithecia: semi-immersed; exciple black (100) 180–200 m in diameter; hamathecial hyphae richly branched and anastomosing; involucrellum absent; asci (49) 75–100 × (11.5) 18–25 m; ascospores with 1–3 septa, both cells of similar shape and size, 18–25 × 4–7.5 m. (FRORJ\ a terrestrial species but occasionally at amphibious sites on ± shaded limestone and wood in the splash water zone (Germany, North Rhine-Westphalia, banks of the small river “Anger” in the city of Ratingen). 'LVWULEXWLRQ This is a rare, possibly overlooked species. The amphibious populations reported from Luxembourg (Molitor & Diederich 1997) may refer to a different taxon as these specimens are described as non-lichenized algicolous fungi without a differentiated lichen thallus.

Fig. 66. Pyrenocollema saxicola: ×16.

Pyrenocollema

115

6LPLODU VSHFLHV Pyrenocollema tichothecioides has wider ascospores and the perithecia are less exposed. Pyrenocollema caesium is similar in thallus appearance but has larger perithecia (200–400 m), and the spore cells are of equal size. The determination of the above cited collection from North Rhine-Westphalia to P. saxicola remains uncertain since the asci are much larger in the lichenized forms than in the non-lichenized specimens reported from Luxembourg (Molitor 'LHGHULFK DQGLQWKHW\SHPDWHULDO+DUULV 7KHJHQHULFDI¿OLDWLRQ as well as the species delimitation of P. saxicola are in need of revision. Uncertain species: Naetrocymbe massalongiana is separated from Pyrenocollema saxicola by the larger asci and ascospores (Harris 1995). An amphibious OLFKHQL]HGVSHFLPHQIURP1RUWK5KLQH:HVWSKDOLD+7KV:E ¿WVEHWWHU with this description although the size of the asci exceeds even the maxima given by Harris (1995). Molitor & Diederich (1997) discussed the distinctiveness of the two taxa and concluded to place N. massalongiana in synonymy to P. saxicola with the remark that still more data are necessary to solve the problems connected with species delimitation in this poorly collected group.

3\UHQRFROOHPDWLFKRWKHFLRLGHV (Arnold) R.C. Harris in Egan 1987 Basionym: Arthopyrenia tichothecioides Arnold 1869. Synonyms: Xanthopyrenia tichothecioides (Arnold) Bachmann 1919. 7KDOOXVgreyish; epilithic, up to 350 m high; algal layer only 30 m high; medulla prosoplectenchymatous, 320 m high, colourless; photobiont cells (yellowish-)green, dispersed or in short chains, with thick cell walls (Trentepohlia sp.), occasionally cells of other eucaryotic algae or cyanobacteria present. Perithecia: immersed with only the apex exposed; hamathecial hyphae richly branched and anastomosing; involucrellum absent; asci 50–60 × 10 m; ascospores uniseptate, lower cell markedly narrowed towards the apex, 18–30 × 8–12 m. (FRORJ\on carbonaceous rocks, found at terrestrial sites as well as (temporarily?) inundated in springs. 'LVWULEXWLRQ This is a poorly known, possibly overlooked species. Records have been published from Germany (Thuringia, Sommerleite), Austria (North Tyrol) and Italy (South Tyrol), but some of these records refer to terrestrial populations. $PSKLELRXVSRSXODWLRQVKDYHEHHQFRQ¿UPHGIURP$XVWULDRQO\ (spring close to Ochsenalm/Waldrast, leg. Arnold). 6LPLODUVSHFLHV Pyrenocollema saxicola has narrower ascospores, and the perithecia are semi-immersed in the thallus. Pyrenocollema caesium is similar in thallus appearance but has equally sized ascospore cells.

116

.H\VWRWKHVSHFLHVDQGVSHFLHVSUR¿OHV

Pyrenopsis Nylander 1858 nom. cons. Important references:+HQVVHQµ¶ -¡UJHQVHQE -¡UJHQVHQ  Henssen (1990). Notes: This is one of the most problematic genera in crustose cyanobacterial lichens. Approximately 60 species have been described but very little is known about species boundaries and much work remains to be done in order to clarify the taxonomy at species level and to evaluate the generic placement of discordant elements. As a consequence, the species concepts adopted here are provisional and changes are to be expected in the future. Likewise, any remarks on distribution and ecology of the species remain sketchy. Key to the species: 1a Ascospores (5) 6–8 (10) × 5–6 m, thallus more irregular, granulose to irregularly areolate, areoles irregularly shaped, uneven, apothecia subsessile ................................................................................................. P. sanguinea E Ascospores larger, (9) 10–15 (18) × (6) 7–10 (11) m, thallus continuous, VSUHDGLQJ HYHQWXDOO\ GLVWLQFWO\ DUHRODWH DUHROHV ÀDW WR VOLJKWO\ FRQYH[ smooth, at the margin thallus more granulose, apothecia immersed to slightly elevated ............................................................................ 3VXEDUHRODWD

Pyrenopsis sanguinea$Q]L¿JV67, 170) Synonyms: Psorotichia sanguinea (Anzi) Jatta 1900, Pyrenopsis rhodosticta auct. medieur. p.p. non (Taylor) Müller Argoviensis 1888. 7KDOOXV reddish to dark reddish-brown, rarely almost black, scattered to conÀXHQW FUXVWRVH LUUHJXODUO\ JUDQXORVH WR DUHRODWH DUHROHV LUUHJXODUO\ VKDSHG WR roundish, (0.25) 0.3–1.0 (1.2) mm, 01–0.4 mm thick, usually slightly to distinctly convex, surface of areoles usually distinctly granulose and uneven, attached to the substrate by hyaline gelatinous basal layer; thallus anatomy densely reticulate, hyphal cells 3–4 (6) × 1.5–3 m in size, hyphae running around photobiont cells; photobiont a coccoid cyanobacterium with thick red gelatinous sheath¿J 170), cells single or in pairs, 10–20 m with sheath, 6–13 m without sheath. Apothecia:S\FQRDVFRFDUSVXVXDOO\IHZUDUHO\DEXQGDQWDW¿UVWVHPLLPPHUVHG soon sessile with broad base, 0.25–0.4 mm wide; thalline margin granulose, perVLVWLQJ±—PWKLFNGLVFDW¿UVWSXQFWLIRUPVRRQRSHQHGDQGÀDWGDUNUHG brown or pale to dark brownish; excip. prop. very thin to virtually absent, apically sometimes to 10 m wide, hyaline; hymenium hyaline, 70–75 m high, I+ deep blue; epihymenium faintly yellowish brown; asci clavate, 40–50 × 10–13 m, outer wall deep blue in I, 8-spored, ascospores simple, hyaline, broad ellipsoid to subglobose, (5) 6–8 (10) × 5–6 m. Pycnidia: semi-immersed in small thallus warts, simple, ± pyriform, c. 50–70 × 90–100 m; pycnospores ellipsoid, 3 × 1–1.5 m. (FRORJ\ on acidic rock in rather exposed situations, in seepage tracks and sporadically inundated at the margin of mountain streams; in moist or wet situations often together with Ephebe lanata.

Pyrenopsis

117

Fig. 67. Pyrenopsis sanguinea: thallus irregularly areolate, surface of areoles rough and uneven, few apothecia; ×18. 'LVWULEXWLRQ montane to alpine, known from southern Black Forrest and Tyrol but perhaps more widely distributed in Central Europe.

3\UHQRSVLVVXEDUHRODWD1\ODQGHU¿JV68, 165) Synonyms: Collemopsis subareolata (Nylander) Malbranche 1881, Pyrenopsis fuliginea var. subareolata (Nylander) Boistel 1903, Pyrenopsis haematopsis sensu auct. brit., Pyrenopsis rhodosticta sensu auct. medieur. p.p. non (Taylor) Müller Argoviensis 1888. 7KDOOXV UHGGLVK WR GDUN UHGGLVKEURZQ VRPHWLPHV DOPRVW EODFN FRQÀXHQW crustose, sometimes spreading across several centimetres; in juvenile stages thallus thin and granulose to small-areolate, becoming thicker and soon distinctly rimose-areolate to areolate, areoles irregularly shaped, (0.2) 0.3–0.8 (1.0) mm ZLGH±PPWKLFNDQJXODWHÀDWWRVOLJKWO\FRQYH[VPRRWKWR¿QHO\JUDQXlose, attached to the substrate by hyaline, gelatinous basal layer; thallus anatomy paraplectenchymatous, hyphal cells 3–4 m; photobiont a coccoid cyanobacterium with red gelatinous sheath, cells single or in pairs, 7–10 m with sheath, 3–6 m without sheath. Apothecia: pycnoascocarps, usually rare, sometimes abundant, 1 (2–3) per areole, immersed to semi-immersed, small, 0.25–0.4 mm ZLGHWKDOOLQHPDUJLQSHUVLVWLQJVPRRWKÀDWRUVOLJKWO\HOHYDWHGQRWSURPLQHQW WR—PWKLFNGLVFDW¿UVWSXQFWLIRUPDQGGHSUHVVHGEHFRPLQJVOLJKWO\RSHQHG but remaining depressed with only slightly elevated thalline margin, brownish, rarely appearing blackish; excip. prop. lacking; hymenium hyaline, 90–95 m high, I+ deep blue; epihymenium colourless to very faintly yellowish-brown; asci narrow clavate to clavate, (40) 50–75 × 12–18 m, outer wall deep blue in I, 8-spored; ascospores simple, hyaline, broad ellipsoid , (9) 10–15 (18) × (6) 7–10 (11) m. Pycnidia: immersed, simple, ± pyriform, c. 90–100 × 70–75 m; pycnospores ellipsoid, c. 3 × 1–1.5 m.

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Fig. 68. Pyrenopsis subareolata: thallus rimose-areolate to areolate, surface rather smooth, fully mature apothecia with small, opened discs; ×20. (FRORJ\ similar to Pyrenopsis sanguinea. 'LVWULEXWLRQ known from southern Black Forest, Vosgues and Alps; also known from Scandinavia; records from oceanic regions may belong to Cryptothele rhodosticta (Taylor) Müller Argoviensis. Uncertain species: Pyrenopsis cleistocarpa (Müller Argoviensis) Forssell is known from a single collection at the banks of the river Dranse in the Swiss Alps. The locality suggests that the species grows inundated at least during high water periods. The thallus of P. cleistocarpa is dark reddish brown, very thin and minutely granulose.

5KL]RFDUSRQde Candolle 1805 Important references: Feuerer (1991), Fryday (2000), Gilbert (2000), Wirth (1995). Key to the species: 1a Thallus yellow .................... R. geographicum (Linnaeus) de Candolle 1805 E Thallus brown or grey ............................................................................... 2 2a Ascospores with one septum, usually with black prothallus ........................ ............................................................................................. 5EDGLRDWUXP E Ascospores with more than one septum, prothallus absent or indistinct ..... 3 3a Ascospores submuriform, colourless, rare species in subalpine to alpine sites ..................................................................................... 5DPSKLELXP E Ascospores muriform, colourless to brown (at least old ones), from lowlands to alpine sites ............................................................................................ 4

Rhizocarpon

119

4a Ascospores 30–42 Pm long, hymenium 120–200 Pm thick, medulla K-, usually amphibious ......................................................................... 5ODYDWXP E Ascospores 20–33 Pm long, hymenium 70–130 Pm thick, medulla K-/K+ yellow, usually terrestrial ................................. R. reductum Th. Fries 1874

5KL]RFDUSRQDPSKLELXP )ULHV 7K)ULHV¿J 69) Basionym: Lecidea amphibia Fries 1822. Synonyms: Abacina amphibia (Fries) Norman 1853, Buellia amphibia (Fries) H. Olivier 1901, Rhizocarpon obscuratum f. amphibium (Fries) Schade 1933. 7KDOOXV grey; areolate, areoles 0.2–0.5 mm; prothallus absent; medulla I-; photobiont coccoid green alga. Apothecia: immersed, black; exciple thin (< 0.03 mm), hardly visible; hymenium 150–200 m high; epihymenium dilute red-brown (K+ purple); hypothecium mid- to dark-brown; asci 8-spored, 85– 100 × 25–30 m; ascospores submuriform, hyaline, halonate, 23–27 × 10.5– 12 m. (FRORJ\ amphibious on siliceous rocks in cold streams. 'LVWULEXWLRQ a rare species at subalpine to alpine sites in the Alps. 6LPLODUVSHFLHV Rhizocarpon lavatum has apothecia with a distinct exciple and muriform ascospores. The terrestrial R. distinctum has a I+ blue medulla. Rhizocarpon badioatrum has an extended black prothallus and distinctly brown coloured ascospores.

Fig. 69. Rhizocarpon amphibium: ×13.

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5KL]RFDUSRQEDGLRDWUXP(Flörke ex Sprengel) Th. Fries 1874 ¿JV 70–71) Basionym: Lecidea badioatra Flörke ex Sprengel 1821. Selected synonyms: Buellia badioatra (Flörke ex Sprengel) Körber 1855, Catocarpus badioater (Flörke ex Sprengel) Arnold 1872. 7KDOOXV brown with black prothallus; areolate, areoles scattered on the prothallus in young thalli or at the thallus margins, forming a compact thallus when older; areoles thick, up to 2 mm in diameter; two chemotypes: medulla K-/P- or medulla K+/P+ (stictinic acid and norstictinic acid in varying quantities detected by t.l.c); photobiont coccoid green alga. Apothecia: semi-immersed to immersed, up to 1 mm in diameter, black, discs epruinose; epihymenium red-brown to dark brown, K+ red; ascospores uniseptate, soon becoming brown, 26–36 × 13– 16 m. (FRORJ\ a terrestrial species but also often amphibious in the splash water zone of cold upland streams on stable siliceous rocks, preferably at sun-exposed sites. 'LVWULEXWLRQ amphibious populations restricted to high altitudes and a frequently found at subalpine to alpine sites. 6LPLODU VSHFLHV The amphibious species Rhizocarpon lavatum has an indistinct prothallus, muriform ascospores and prominent apothecia with a distinct margin. Rhizocarpon jemtlandicum is a terrestrial species with a K- epihymenium.

Fig. 70. Rhizocarpon badioatrum: central part of thallus with apothecia; ×18.

Rhizocarpon

121

Fig. 71. Rhizocarpon badioatrum: thallus margin with scattered areoles on extended black prothallus; ×18.

5KL]RFDUSRQODYDWXP )ULHV +D]VOLQVN\¿J72) Basionym: Lecidea lavata Acharius ex Fries 1827. Synonyms: Lichen petraeus var. obscuratus Acharius 1810, Buellia obscurata (Acharius) H. Olivier 1884, Rhizocarpon obscuratum f. lavatum (Acharius ex Fries) Stein 1879, Rhizocarpon petraeum var. lavatum (Acharius ex Fries) Körber 1855, Buellia lavata (Acharius ex Fries) Monguillon 1900, Phalodictyum obscuratum (Acharius) Clements 1909, Rhizocarpon obscuratum (Acharius) A. Massalongo 1852 non auct., Diplotomma obscuratum (Acharius) Jatta 1900.

Fig. 72. Rhizocarpon lavatum: ×13.

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7KDOOXV EURZQ UHGEURZQ RFKUH JUH\ FRQWLQXRXV ¿QHO\ ULPRVH WR FUDFNHG prothallus indistinct; medulla K-, P-; photobiont coccoid green alga (Trebouxia sp. in own cultivated isolates). Apothecia: distinctly raised above the thallus with prominent margin, 800–1000 m in diameter; hymenium 120–200 Pm high; exciple close to the hymenium > 40 Pm high; ascospores muriform, colourless or faintly brown with age, 30–42 × 14–18 Pm. (FRORJ\ amphibious in the splash water zone of clean streams with neutral to moderately acidic water (pH > 5), on stable siliceous rocks from rather shady to sun exposed sites. 'LVWULEXWLRQ widespread but scattered from lowlands to alpine areas. 6LPLODUVSHFLHV Rhizocarpon reductum is a terrestrial species with smaller, immersed apothecia, a lower hymenium (70–130 Pm) and smaller ascospores (20–32 × 9–15 Pm). Rhizocarpon amphibium is a rare subalpine to alpine species with submuriform, colourless ascospores. Rhizocarpon badioatrum has uniseptate, brown ascospores and a black prothallus.

Rinodina (Acharius) Gray 1821 Important references: Purvis (1992), Wirth (1995). Key to the species: 1a Thallus K+ yellow (but sometimes weak or K-), atranorin detected by t.l.c., exciple and apothecial disc ± concolorous or with thalline exciple, exciple lacking algae or algae very few, prothallus r pure black .......... R. oxydata E Thallus always K-, no lichen substances detected by t.l.c., apothecia always with thalline exciple containing numerous algae, prothallus absent or blueEODFNDQG¿PEULDWH .................................................................................... 2 2a Apothecia immersed to slightly elevated, thallus eventually ZLWK ¿PEULDWH blue-black prothallus, ascospores of MischoblastiaW\SH¿J 8) ... 5¿PEULDWD E Apothecia slightly immersed to sessile, prothallus lacking, ascospores intermediate between Mischoblastia- and PachysporariaW\SH¿JV 8–9) ......... ........................................................ R. teichophila (Nylander) Arnold 1863

5LQRGLQD¿PEULDWD .|UEHU¿J 73) 7KDOOXV brown-black; thin, cracked; eventuallyUHVWLQJRQ“¿PEULDWHSURWKDOOXV K- (no lichen substances detected by t.l.c.); photobiont coccoid green alga (Trebouxia incrustata in own cultivated isolates). Apothecia: immersed to semi-immersed, XS WR —P LQ GLDPHWHU GLVFV ÀDW EURZQEODFN K\PHQLXP ±—P KLJK ascospores with wall thickening of Mischoblastia-type (thick walls with heartshaped lumen), 22–31 × 11–17 m. (FRORJ\ amphibious in the splash water zone of streams with well buffered water on siliceous rocks at shady to sunny sites.

Rinodina

123

Fig. 73. 5LQRGLQD¿PEULDWD: ×17. 'LVWULEXWLRQ a rare species at rather low altitudes in suboceanic areas. 6LPLODUVSHFLHV Rinodina oxydata has a variable, whitish, grey, brown to ochre thallus which turns K+ yellow (reaction sometimes weak, atranorin detected by t.l.c) SURWKDOOXV LI SUHVHQW QRW ¿PEULDWH, the ascospores are usually slightly smaller (19–25 m), and the hymenium is usually less tall (70–115 m). Rinodina teichophila is a terrestrial species with greyish to brown colour lacking a prothallus and having ascospores intermediate between Pachysporaria- and Mischoblastia-type (apical thickenings less developed than in spores of 5¿Pbriata, lumen only slightly depressed in the centre).

Rinodina oxydata $0DVVDORQJR $0DVVDORQJR¿J74) Basionym: Berengeria oxydata A. Massalongo 1853. Synonyms: Lecidea discolor Hepp 1857, Buellia discolorr (Hepp) Anzi 1860, Rinodina biatorina Körber 1859, Rinodina discolorr (Hepp) Arnold 1872, Rinodina subarenaria A.L. Smith 1918. 7KDOOXV colouration very variable, whitish, grey, brown to ochre; thin to thick, rimose to cracked, K+ yellow (rarely K-), atranorin detected by t.l.c.; prothallus, LISUHVHQWGDUNDQGHQWLUHQRW¿PEULDWHSKRWRELRQWFRFFRLGJUHHQDOJDApothecia: immersed to sessile, dark brown to black, up to 800 m in diameter; epithecium pale brown; hymenium 70–115 m high; ascospores with wall-thickening of Mischoblastia-type (lumen triangularly compressed with the longest side distinctly depressed), 19–25 × 9–14 m. (FRORJ\ at humid to damp or rarely submerged sites on solid siliceous rocks. 'LVWULEXWLRQ terrestrial populations widespread but rare, amphibious populations known from streams in mountainous regions, e.g. Vosgues and Upper Austria (F. Berger in litt.).

124

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Fig. 74. Rinodina oxydata: ×23. 6LPLODUVSHFLHV The thalli of 5LQRGLQD¿PEULDWD do not contain atranorin. They are always K- and eventually have a ¿PEULDWHSURWKDOOXV7KHDVFRVSRUHVWHQGWR be larger (22–31 m) and the exciple always contains algal cells. The terrestrial R. teichophila reacts K-, the thallus is not surrounded by a prothallus, and the ascospores have a wider lumen intermediate between Mischoblastia- and Pachysporaria-type.

Sporodictyon A. Massalongo 1852 Notes: Sporodictyon was long treated as a subgenus of Polyblastia. In a recent multigene study, however, Polyblastia was clearly shown to be polyphyletic whereas Sporodictyon IRUPHGDPRQRSK\OHWLFHQWLW\6DYLüHWDO 7KHVH authors proposed to re-establish Sporodictyon at generic rank in order to separate it from the remainder of Polyblastia which continues to be polyphyletic and is therefore still in need of further studies. Sporodictyon LQWKHVHQVHRI6DYLüHWDO   FRQVLVWV RI ¿YH VSHFLHV ZLWK S. cruentum as the only specialized freshwater representative in Central Europe. Important references:6DYLüHWDO 

Sporodictyon cruentum .|UEHU .|UEHU¿J 75) Basionym: Segestrella cruenta Körber 1853. Synonyms: Sphaeromphale cruenta (Körber) Körber 1855, Polyblastia cruenta (Körber) P. James & Swinscow 1971. Uncertain species: Sphaeromphale henscheliana Körber 1855, Sporodictyon henschelianum (Körber) Körber 1863, Verrucaria henscheliana (Körber) Leighton 1871, Polyblastia henscheliana (Körber) Lönnroth 1858.

Sporodictyon

125

Fig. 75. Sporodictyon cruentum: ×15. 7KDOOXV greenish, grey, dark brown to black; thin, continuous to rimose or cracked; photobiont coccoid green alga. Perithecia: prominent, 500–800 m in diameter; involucrellum reaching far below, lower parts covered by a thin thallus mantle; asci 8-spored; ascospores muriform, pale brown (at least when old), 40–90 × 20–40 m. (FRORJ\ amphibious or submerged in cool streams and streams on stable siliceous rocks. 'LVWULEXWLRQ scattered in high mountainous (Black Forest, Bayerischer Wald, Sudetes, Austria, Italy) to alpine regions (Alps, Tatra); widely distributed in Nordic countries. 6LPLODU VSHFLHV Thelidium methorium, 6WDXURWKHOH ¿VVD and S. clopima have VXSHU¿FLDOO\ VLPLODU WKDOOL DQG SHULWKHFLD EXW T. methorium has 2-celled ascospores and species of Staurothele have hymenial algae in the perithecia. Uncertain species: The distinction of Sporodictyon henschelianum from S. cruentum is still in neHGRIIXUWKHUVWXG\6DYLüHWDO 

6WDXURWKHOHNorman 1853 nom. cons. Notes: The genus Staurothele as currently circumscribed is a polyphyletic assemblage as shown by recent multigene studies (Gueidan et al. 2008). Staurothele s.str. comprises epilithic species only, and probably most of the freshwater speFLHVIURPVLOLFHRXVVXEVWUDWDEHORQJKHUH7KHJHQHULFDI¿OLDWLRQRIWKHVSHFLHV IURPFDOFDUHRXVVXEVWUDWDUHPDLQVWREHFODUL¿HG7KHV\QRQ\P\ RIVHYHUDOVSHcies is very complex and only a selection of older synonyms is given here. The

126

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differentiation of several species mainly relies on differences in the shape of the hymenial algae. This character is usually well applicable (average of lenght/widht ratio calculated from at least 10 different cells), but intermediate forms exist. Whether the shape of the hymenial algae is a suitable character for species delimitation has been discussed controversially (Thomson 1991). Molecular studies focussing on this problem will be published in due course (Thüs, in prep.). Only a selection of mainly terrestrial species from calcareous substrata are included in the key. Further species which have not been reported from Central European streams so far, but possibly occur in ephemeric water bodies or at sun exposed river banks are included in Poelt (1969). Important references: Clauzade & Roux (1985), Gueidan et al. (2008), Poelt (1969), Purvis et al. (1992), Thomson (1991), Vainio (1921), Wirth (1995), Zschacke (1934). Key to the species: 1a Thallus epilithic, on siliceous or calcareous substrata ............................... 2 E Thallus endolithic, on calcareous substrata ............................................... 7 2a Perithecia ± immersed in the thallus, < 20 % raised above level of surrounding plane thallus areas ............................................................................... 3 E Perithecia forming warts raised > 50 % above surrounding plane thallus areas .......................................................................................................... 5 3a 7KDOOXVFRQWLQXRXVRU¿VVXUHGQRWDUHRODWHXVXDOO\ZLWK DQ RUDQJH WLQJH tips of the perithecia often completely immersed in the thallus, restricted to upland areas ........................................................................ 6FORSLPRLGHV E Thallus areolate, usually light brown to deep black-brown, tips of the perithecia immersed or slightly raised above the thallus ................................. 4 4a Hymenial algae bacilliform, thallus brown, restricted to high-montane to alpine regions ............................................................................ 6DUHRODWD E Hymenial algae round or square, thallus light brown to greenish-grey, from lowland to high-montane regions ......................................... 6IUXVWXOHQWD 5a Mature ascospores colourless, perithecia sessile, more than 50 % naked ...... ................................................................................................. S. succedens E Mature ascospores brown, perithecia semi-immersed in thallus warts, covered by a thin thallus layer ...................................................................... 6 6a Hymenial algae bacilliform, restricted to high-montane to alpine regions .... .................................................................................................... 6FORSLPD E Hymenial algae round or square, distributed from lowland to alpine regions .......................................................................................................... 6¿VVD 7a Ascus with 1–2 spores ....... S. orbicularis (A. Massalongo) Th. Fries 1865 E Ascus with 4–8 spores .............................................................................. 8 8a Ascospores 40–82 × 18–32 Pm ................................................... 6VROYHQV E Ascospores 18–35 × 14–20 Pm .......... S. bacilligera (Arnold) Arnold 1885

Staurothele

127

6WDXURWKHOHDUHRODWD $FKDULXV /HWWDX¿J153) Basionym: Pyrenula areolata Acharius 1814. Synonyms: Staurothele clopima var. areolata (Acharius) Trevisan 1860, Pertusaria areolata (Acharius) A. Massalongo 1852. 7KDOOXV brown to black; thick, areolate; prothallus absent; photobiont StichococcusVSYHUL¿HGE\FXOWLYDWHGLVRODWHV Perithecia: immersed in thallus areoles, 200–450 m in diameter; involucrellum apical, poorly developed; hymenial algae bacilliform; asci 2-spored; ascospores muriform, brown at maturity, 30–50 × 15–25 m. Pycnidia: rare, 140 × 70 m; pycnospores 4–5 × 1 m. (FRORJ\ terrestrial on nutrient enriched, basic or slightly acidic substrata; in upland (alpine areas) at larger streams and rivers or especially at lake shores tolerating temporary inundation. 'LVWULEXWLRQ high-montane to alpine areas, widespread (also at lower altitudes) in Nordic countries. 6LPLODUVSHFLHV Staurothele frustulenta is separated by globose-cuboid hymenial algae and a usually paler thallus colour. Staurothele frustulenta occurs in similar habitats like S. areolata, however at low altitudes. The separation of the two species at species rank requires reconsideration. Staurothele clopimoides is a strictly amphibious or even aquatic upland lichen with much smaller perithecia and a continuous, rimose to areolate, but usually rather plane and often orangetinged thallus. 6WDXURWKHOH ¿VVD and S. clopima have thin thalli with perithecia which are immersed in raised thallus warts.

6WDXURWKHOHFORSLPD :DKOHQEHUJ 7K)ULHV¿JV76–77) Basionym: Verrucaria clopima Wahlenberg in Acharius 1803. Synonyms: Verrucaria cuprea var. fuscocuprea Nylander 1874, Verrucaria fuscocuprea Nylander 1881, Staurothele fuscocuprea (Nylander) Zschacke 1913, Verrucaria umbrina Wahlenberg 1812, Sphaeromphale umbrina (Wahlenberg) Mudd 1861, Staurothele umbrina (Wahlenberg) Hellbom 1884, Polyblastia umbrina (Wahlenberg) Rostrup 1871, Staurothele catalepta Malme non (Acharius) Blomberg & Forssell 1880. 7KDOOXV brown, brown-black or greenish (especially at shaded sites); thin, conWLQXRXV WR ¿QHO\ ULPRVH RU FUDFNHGDUHRODWH SKRWRELRQW Stichococcus sp. with EDFLOOLIRUPFHOOVYHUL¿HGE\FXOWLYDWHGLVRODWHV  Perithecia: immersed in prominent thallus warts, 200–600 m in diameter; involucrellum variable from apical to reaching down to the thallus base; hymenial algae bacilliform; asci 2-spored; ascospores muriform, dark brown at maturity, 25–50 × 15–25 m. (FRORJ\ amphibious in the splash water zone or at temporarily submerged sites, from shady to sun-exposed localities.

128

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Fig. 76. Staurothele clopima: specimen with almost continuous thallus; ×16.

Fig. 77. Staurothele clopima: specimen with high number of cracks in the thallus, ×16. 'LVWULEXWLRQ widespread at high-montane to alpine sites, also recorded from Nordic countries and North America. 6LPLODUVSHFLHV 6WDXURWKHOH¿VVDis very similar, but the majority of its hymenial algae are globose-cuboid. The taxonomic status of 6¿VVD requires reconsideration. Staurothele areolata has a much thicker thallus with perithecia which are immersed in thallus areoles and do not form prominent thallus warts. Staurothele succedens has colourless ascospores and naked perithecia. 6WDXURWKHOH ¿VFKHUL and S. isarina are uncertain taxa with much similarity to S. succedens, but pale brown (6¿VFKHUL) or red (S. isarina) ascospores. Both differ from S. clopima by

Staurothele

129

light grey thalli and perithecia which are semi-immersed in thalline warts or almost completely naked. The brown colour of ascospores in 6¿VFKHUL disappears when treated with potassium hydroxide (Behr 1958). For further details see notes DWWKHHQGRIWKHVSHFLHVSUR¿OHIRUS. succedens.

6WDXURWKHOHFORSLPRLGHV $Q]L -6WHLQHU¿JV 78, 151) Basionym: 6WLJPDWRPPD¿VVXP var. clopimoides Baglietto & Carestia 1865. Synonyms: Polyblastia clopimoides (Baglietto & Carestia) Anzi 1871, Sphaeromphale clopimoides (Baglietto & Carestia) Arnold 1874, Stigmatomma clopimoides (Baglietto & Carestia) Baglietto & Carestia 1881, Verrucaria clopimoides (Baglietto & Carestia) Nylander 1881. 7KDOOXV brownish to copper and often with orange tinge¿J ; thick, cracked to almost areolate but smooth; photobiont Stichococcus sp. with bacilliform cells YHUL¿HGE\FXOWLYDWHGLVRODWHV Perithecia: immersed in the thallus, visible only around the ostiole which is mantled by a black apical involucrellum; hymenial algae bacilliform, 7–18 × 3 m; ascospores muriform, brown at maturity, 38–50 × 14–21 m. (FRORJ\ amphibious in the splash water zone or submerged for several months per year on siliceous rocks in cold streams with usually permanent water supply; usually in well lit places. 'LVWULEXWLRQ subalpine to alpine areas in the Alps and at high altitudes in the Carpathians (e.g. Tatra). 6LPLODUVSHFLHV Staurothele areolata and S. frustulenta have distinctly areolate and often uneven or ± scattered thalli lacking orange or copper colour shades and preferring dryer sites.

Fig. 78. Staurothele clopimoides: ×15.

130

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6WDXURWKHOH¿VVD7D\ORU =ZDFNK¿J79) Basionym: 9HUUXFDULD¿VVD Taylor 1836. Synonyms: Thelotrema¿VVXP (Taylor) Hepp 1853, 6SKDHURPSKDOH ¿VVD (Taylor) Körber 1855, 6WLJPDWRPPD ¿VVXP (Taylor) Baglietto & Carestia 1865, 3RO\EODVWLD¿VVD (Taylor) Jatta 1900, Staurothele hazslinskyi (Körber) Blomberg & Forssell 1880, Staurothele lithina Zahlbruckner 1921, Staurothele fuliginea Zschacke 1934, Staurothele silesiaca (A. Massalongo) Zschacke 1934, Staurothele viridis Zschacke 1913. Uncertain species: Staurothele oenipontana Beschel 1954 7KDOOXV brown, brown-black or greenish (especially at shaded sites); thin, conWLQXRXV WR ¿QHO\ ULPRVH RU FUDFNHGDUHRODWH SKRWRELRQW Stichococcus sp. with URXQGRUVTXDUHFHOOVYHUL¿HGE\FXOWLYDWHGLVRODWHV Perithecia: immersed in prominent thallus warts, 200–600 m in diameter; involucrellum variable from apical to reaching down to the thallus base; hymenial algae rounded to square; asci 2-spored; ascospores muriform, dark brown at maturity, 25–50 × 15– 25 m. (FRORJ\ amphibious in the splash water zone or at temporarily submerged sites, from shady to sun-exposed localities. 'LVWULEXWLRQ from colline to alpine sites but much declined at low altitudes. 6LPLODUVSHFLHV Staurothele clopima is very similar and separated only by the bacilliform hymenial algae. Intermediate forms with mixed populations of rounded and bacilliform hymenial algae are known and the distinction of 6¿VVD from S. clopima is currently under reconsideration (Thüs in prep.). Staurothele succedens has colourless ascospores. Staurothele areolata has a much thicker thallus with perithecia which are immersed in thallus areoles and, therefore, lacks prominent thallus warts.

Fig. 79. 6WDXURWKHOH¿VVD: ×17.

Staurothele

131

Uncertain species: Staurothele oenipontana is a little known taxon, which keys out with 6¿VVD, but differs from this polymorphic taxon only by constantly 4celled instead of muriform ascospores. The 4-celled ascospores are unique among the genus Staurothele. The taxon is solely known from its type locality on dolomitic rocks at the banks of the river Inn in the city of Innsbruck together with S. ¿VVD (on the same substratum) and it is likely that it represents a genetic variety of this species rather than a distinct taxon.

6WDXURWKHOHIUXVWXOHQWD9DLQLR¿JV 80, 152) Uncertain species: Staurothele ambrosiana (A. Massalongo) Lettau 1914. 7KDOOXV brown to black; thick, areolate, occasionally with white to brownish prothallus; photobiont either Stichococcus sp. or Diplosphaera chodatiiYHUL¿HG by cultivated isolates). Perithecia: immersed in thallus areoles, 200–450 m in diameter; involucrellum apical, poorly developed; hymenial algae globose-cuboid; asci 2-spored; ascospores muriform, brown at maturity, 30–50 × 15–25 m. (FRORJ\terrestrial at nutrient enriched, basic or slightly acidic substrata, often on hard substrata along large rivers forming a typical vegetation belt which is regularly inundated – but only rarely and for short time. Tolerant to moderate silting but absent from heavily polluted river banks (Erichsen 1957, Heibel 1996, Thüs 2006). 'LVWULEXWLRQ from sea-level to high-montane regions. 6LPLODUVSHFLHV Staurothele areolata is very similar and differs only in the bacilliform hymenial algae and usually darker thallus colour. The separation of S. areolata and S. frustulenta at species rank is in need of reconsideration. Stau-

Fig. 80. Staurothele frustulenta: ×16.

132

.H\VWRWKHVSHFLHVDQGVSHFLHVSUR¿OHV

rothele areolata replaces S. frustulenta at higher altitudes at least to a certain degree. Staurothele clopimoides is a strictly amphibious or even aquatic upland lichen with much smaller perithecia and a rimose, sometimes almost continuous to areolate and often orange tinged thallus. Staurothele¿VVD and S. clopima have thin thalli with perithecia immersed in distinctly projecting thallus warts. Uncertain species: Specimens with a white prothallus (also brownish according to Pereira 1992) have been described as Staurothele ambrosiana. However, the colouration and presence of a prothallus seem to be very variable (Pereira 1992). Furthermore, S. ambrosiana is regarded as a terrestrial to riparian species but specimens of S. frustulenata with white prothallus have recently been collected in the intertidal zone of the River Elbe in and in the vicinity of Hamburg, GerPDQ\¿J &XUUHQWO\WKHVHSDUDWLRQRIS. ambrosiana and S. frustulenta is QRW VXI¿FLHQWO\ FRUUREDWHG DQG WKHUHIRUH WKH IRUPHU VSHFLHV LV WUHDWHG KHUH DV uncertain. Detailed studies are required to clarify the taxonomic status of the two species.

6WDXURWKHOHVROYHQV$Q]L =VFKDFNH¿JV81–83) Basionym: Polyblastia solvens Anzi 1864. Synonym: Verrucaria solvens (Anzi) Stizenberger 1882. Uncertain species: Staurothele meylanii de Lesdain 1931. 7KDOOXV whitish to brownish; thin, epilithic to (semi-) endolithic; photobiont coccoid green alga. Perithecia: semi-immersed, up to 700 m in diameter; involucrellum apical to reaching the thallus base; hymenial algae globose-cuboid; asci 4-spored; ascospores muriform, red-brown at maturity, 40–82 × 18–32 m. (FRORJ\ amphibious on wet calcareous rocks.

Fig. 81. Staurothele solvens (S. meylanii f. geographicum Servít): ×15.

Staurothele

133

Fig. 82. Staurothele solvens (S. meylanii f. papulare Klement): ×14. 'LVWULEXWLRQ a rare species from lowland (northern France) to montane regions in the Alps. 6LPLODUVSHFLHV The terrestrial species Staurothele bacilligera has smaller ascospores and an endolithic thallus. Uncertain species: Staurothele meylanii is mainly characterized by the small perithecia (up to 500 m in diameter) and the whitish-grey thallus colour, while in S. solvens s.str. the thallus is brownish-red. The distinction of the two species requires further study.

Fig. 83. Staurothele solvens ( Staurothele meylanii f. superiorr Klement): ×16.

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.H\VWRWKHVSHFLHVDQGVSHFLHVSUR¿OHV

6WDXURWKHOHVXFFHGHQV 5HKPH[$UQROG $UQROG¿J 84) Basionym: Polyblastia succedens Rehm ex Arnold 1870. Synonym: Verrucaria succedens (Rehm ex Arnold) Nylander 1882. Uncertain species: 6WDXURWKHOH¿VFKHUL Behr 1958, Staurothele isarina Riehmer 1959. 7KDOOXV brown to grey-black or green-black, on calcareous rocks often with whitLVKSUXLQDWKLQVPRRWKWR¿QHO\JUDQXODUSKRWRELRQWStichococcus sp. Perithecia: sessile, 300–500 × 400–900 m, the apices of the moistened, swollen periphyses are often visible as a white mass protruding out of the ostiole of the perithecia; involucrellum distinct, reaching far down; asci 8-spored, 155 × 45 m; hymenial algae bacilliform, 3–11 × 2–4.5 m; ascospores muriform, colourless, rarely pale reddish or yellowish (see notes on uncertain species), 35–45 × 15–25 m. (FRORJ\ on slightly alkaline to alkaline substrata (calcareous rocks or alkaline silicates) at very humid or frequently wetted places, especially close to waterfalls, rarely in the splash water zone or at temporarily inundated sites. 'LVWULEXWLRQ scattered in mountainous regions. 6LPLODUVSHFLHV differs from other species of Staurothele in the habitat and in the combination of epilithic thallus, large sessile perithecia and colourless ascospores. Uncertain species: Staurothele isarina is reported to contain red ascospores and mostly globose hymenial algae (Riehmer 1959). The type material (W) consists of nine pieces of rock, three of them with perithecia of Staurothele. None of the Staurothele thalli is completely in concordance with the description given by Riehmer (1959). All ascospores are colourless and the majority of the hymenial algae are bacilliform as in common forms of S. succedens. Staurothele isarina is

Fig. 84. Staurothele succedens: ×17.

Thelidium

135

reported to have a light grey thallus. This is present in most thalli of the type material, but there is also much variation in thallus colouration among the specimens with transitional forms from light grey to blackish brown as in S. succedens. Most thalli are covered by silt and partly eroded which may explain the rather unusual habit and colour compared to common forms of S. succedens. The red pigmtation of the ascospores in the type of S. isarina has either disappeared during storage or the description of red ascospores was based on one aberrant perithecium of S. succedens. 6WDXURWKHOH¿VFKHUL is similar to S. succedens, except for brownish ascospores (soon pale brownish, already in the asci) a light greyish thallus colour and perithecia, which are distinctly immersed with their basal parts in the thin to thick thallus. The brown ascospore colour disappears when treated with potassium hydroxide (Behr 1958).With the thin to thick, grey thallus and coloured ascospores 6¿VFKHUL comes close to the original description of S. isarina, mainly differing in the brownish instead of red ascospores.

7KHOLGLXPA. Massalongo 1855 Notes: Thelidium is one of the large crustose genera in the order Verrucariales which is united only formally by the type of septation of the ascospores. All species with a single or with several transversal septa but without (or only occasional) longitudinal septa are included in this genus. Recent molecular studies *XHLGDQ HW DO  6DYLü HW DO   UHYHDOHG WKDW VHSWDWH DVFRVSRUHV KDYH evolved several times independently in the Verrucariales and Thüs & Nascimbene (2008) showed that even the amphibious species of Thelidium do not form a monophyletic group. Therefore, major changes are to be expected in the genus FLUFXPVFULSWLRQDQGWKHJHQHULFDI¿OLDWLRQRIThelidium species. Important references: Clauzade & Roux (1985), Gueidan et al. (2007), Orange  6DYLüHWDO 6HUYtW 7KV 1DVFLPEHQH =VFKDFNH (1920, 1933). Key to the species: 1a Perithecia completely immersed in the substratum, visible only by the black ostiolum or the uppermost part of the involucrellum, thallus endolithic in calcareous rocks ........................................................................................ 2 E Perithecia semi-immersed (< 50 %) or epilithic, thallus semi-endolithic to completely epilithic on limestone or siliceous substrata ........................... 4 2a Perithecia 140–230 m, ascospores 32–36 × 10–14 Pm, 3-septate, amphibious species ............................................................................ T. inundatum E Perithecia larger, ascospores wider (usually terrestrial species) ................ 3 3a Ascospores 3-septate ... T. incavatum-group (several taxa of uncertain status) E Ascospores 2-septate ... T. decipiens-group (several taxa of uncertain status) 4a Ascospores 2-celled .................................................................................. 5 E Ascospores 4-celled ................................................................................ 14 5a On calcareous rocks .................................................................................. 6 E On siliceous rocks ..................................................................................... 8

136

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6a Thallus epilithic, up to 100 Pm thick, perithecia 100–300 Pm in diameter, a usually terrestrial species, tolerating only temporary submersion ... 7PLQXWXOXP E Thallus usually semi-endolithic, epilithic forms > 100 Pm thick, amphibious species, usually submerged ....................................................................... 7 7a Exciple up to 200 Pm in diameter, involucrellum apical or absent .............. ................................................................................................. 7NOHPHQWLL E Exciple 300–450 m in diameter, involucrellum absent ...... 7ULYXOLFROXP 8a Involucrellum absent or weak and usually indistinctly separated from the exciple, thallus whitish, grey, greenish or light brownish, never dark brown, photobiont cells irregularly arranged or aggregated in small groups ........ 9 E Involucrellum distinctly separated from the exciple, thallus grey, dark olive, brownish to black-brown, never green, photobiont cells irregularly arranged ................................................................................................................. 10 9a 3KRWRELRQWFHOOVLUUHJXODUO\DUUDQJHG¿J 90), ascospores 19–30 × 8–15 Pm, involucrellum absent or fused with the exciple and visible as a slight thickening in the upper part of the perithecium, on sandstone ............. T. rehmii E 3KRWRELRQWFHOOVDUUDQJHGLQVPDOODJJUHJDWHGJURXSV¿J89), ascospores 13–32 × 4–15 Pm, involucrellum absent or thin, indistinctly separated from the exciple and reaching the thallus base, on various (usually well buffered) substrata ................................................................................ 7PLQXWXOXP 10a Ascospores 16–22 Pm long .................................................. 7ODKPLDQXP E Ascospores longer .................................................................................... 11 11a More than half of the perithecia covered by a thallus mantle, involucrellum usually weak (up to 30 PPWKLFN¿J86) and often fading in lower parts ... ................................................................................................... 7SOXYLXP E Perithecia naked at least in upper two thirds, involucrellum thin to very thick (25–170 Pm), evenly coloured throughout .............................................. 12 12a Involucrellum very thick (115–170 PPLQXSSHUWKLUG¿J 87), ratio of involucrellum thickness to exciple diameter > 0.4 ......................... T. pertusatii E Involucrellum thinner (usually < 90 PPLQXSSHUWKLUG¿J88), ratio of involucrellum thickness to exciple diameter < 0.33 ................................... 13 13a Perithecia (430) 500 (1600) Pm in diameter, ascospores (26) 34 (46) Pm long ...................................................................................... T. methorium E Perithecia (250) 361 (510) Pm in diameter, ascospores (26) 28 (39) Pm long ................................................................................. 7VXEPHWKRULXP 14a Involucrellum present ............................................................................. 15 E Involucrellum absent ............................................................................... 19 15a Involucrellum thin, 10–30 (35) PP ¿J 85), ascospores (22) 30 (36) Pm long ...................................................................................... T. fontigenum E Involucrellum thick (> 40 Pm), ascospores 23–52 Pm long .................... 16 16a Thallus light coloured, whitish, grey to brownish, perithecia 300–800 Pm in diameter (usually > 500 Pm) ................................................................... 17 E Thallus dark coloured, brown to brown-black or green-grey to olive-greenblack, perithecia 200–485 Pm ................................................................. 18 17a Ascospores (13) 17 (21) m wide, base of perithecia usually leaving pits in the substratum, thallus thin and mostly endolithic to thick, with epilithic parts 10–80 Pm thick, black basal layer absent ........................ 7SDSXODUH E Ascospores (16) 19 (23) Pm wide, base of perithecia not forming pits in the substratum, thallus thick (> 100 Pm), epilithic parts 50–100 Pm thick, partly with black basal layer .................................................................................. ....... T. suzaeanum Servít (see note on uncertain species under T. papulare)

7KHOLGLXP   18a 2Q VLOLFHRXV URFNV WKDOOXV EURZQ WR EURZQEODFN SDUWO\ ZLWK WKLQ EODFN EDVDOOD\HUSURWKDOOXVEODFN ..................................................... T. nigricans 18b 2QFDOFDUHRXVURFNVWKDOOXVJUHHQLVKJUH\LVKROLYHJUHHQEODFNWREURZQ EODFNEDVDOOD\HUDEVHQWRURUDQJHWREURZQFRORXUHGSURWKDOOXVZKLWHRU DEVHQW .............................................................................. T. aethioboloides 19a Ascospores (20) 29 (36) PPORQJDVFL± PPORQJ ......... T. zwackhii 19b $VFRVSRUHV±PPORQJDVFL±PPORQJ .................................... 20 20a ([FLSOHPPLQGLDPHWHUZLWKEODFNEDVHDVFLXSWRPPORQJWKDOOXV WKLQWRVHPLHQGROLWKLFRQFDOFDUHRXVVXEVWUDWD ........ T. circumspersellum 20b ([FLSOH± PPLQGLDPHWHUZLWKFRORXUOHVVEDVHDVFL±PPORQJ WKDOOXVWKLQWRWKLFN± PP HSLOLWKLFRQVLOLFHRXVVXEVWUDWD ................. ............................................................................................... T. aquaticum

Fig. Fi g 85

Fig. 87

Fig. 89

Fig. Fi g 86

Fig. 88

Fig. 90

Plate 4. Figs. 85– 5 90. ShapeRILQYROXFUHOOXP in Thelidium. Fig. 85. T. fontigenum. Fig. 86. T. pluvium. Fig. 87. 7 T. pertusatii. Fig. 88. T. methorium. Fig. 89. T. minutulum±H[FLSOHVKDSHDQGSKRWRELRQWDUUDQJHPHQW Fig. 90. T. rehmii ±H[FLSOHVKDSHDQGSKRWRELRQWDUUDQJHPHQW

138

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Fig. 91. Thelidium aethioboloides: ×21.

7KHOLGLXPDHWKLRERORLGHV =VFKDFNH¿JV 91, 156) 7KDOOXV black-brown, olive, dark greyish-green; thin to thick (35–100 m), ± continuous, with few cracks, margins often transparent when wetted (subgelatinous); prothallus whitish, little developed or absent; cortex usually yellowish to brown; basal layer locally present, with orange-brown colour; algal layer paraplectenchymatous; photobiont green alga, cells 4–9 m, occasionally also with bacilliform algae, 8–11 × 4–5 m, dispersed in small aggregated groups. Perithecia: immersed LQWKH“SODQHWKDOOXVRUHPEHGGHGLQÀDWWKDOOXVZDUWVH[SRVLQJRQO\WKHQDNHG ostiole; exciple 145–240 m diameter; periphyses up to 21 m long; involucrellum apical to reaching the thallus base, laterally extending in the thallus; asci 73– 93 × ±—PDVFRVSRUHVDW¿UVWVRRQ±VHSWDWH± ×10–15 m. (FRORJ\ amphibious on limestone rocks, usually not permanently submerged. 'LVWULEXWLRQ only known from a few locations in the Swiss Alps (Davos), Bavaria (Allgaeu) and Italy (Trentino, Veneto, leg. J. Nascimbene); in upper montane to subalpine areas but always below the timber line. 6LPLODUVSHFLHVThelidium nigricans occurs on siliceous substrata and develops a black basal layer and eventually a dark prothallus. In other species of Thelidium the thallus is either not dark coloured and the involcurellum is absent or the ascospores have only one septum.

7KHOLGLXPDTXDWLFXP 6HUYtW¿J 92) 7KDOOXV dark olive greenish to brownish; continuous or with small cracks, thin (20–75 m); paraplectenchymatous; cortex lacking but uppermost layer of the thallus partly yellowish-brown; brownish basal layer present only in parts of the

Thelidium

139

Fig. 92. Thelidium aquaticum: ×26. thallus, up to 20 m; photobiont green alga with rounded to elongated cells, 6–9 Pm in diameter, densely packed and homogeneously distributed, only partly agglomerated. Perithecia: completely immersed in the thallus or only the upper half exposed; exciple diameter 80–115 m, exposed parts brown-black, lower parts colourless; periphyses up to 12 m; involucrellum absent or very thin and apical, up to 150 m in diameter; asci 40–60 × 10–16 m; ascospores (1) 3 (4)septate, 16–27 × 5–8 m. (FRORJ\ amphibious in a river on siliceous substrate. 'LVWULEXWLRQ only known from the type locality in Bohemia at 600 m. The potential risks for the survival of this population should be considered. 6LPLODUVSHFLHV Thelidium aethioboloides and T. suzaeanum have thicker thalli, the exciple is protected by a distinct involucrellum and the ascospores are larger.

7KHOLGLXPFLUFXPVSHUVHOOXP 1\ODQGHU =VFKDFNH¿J93) Basionym: Verrucaria circumspersella Nylander 1881. 7KDOOXV brownish; inconspicuous, very thin to vanishing; cortex and basal layer absent; photobiont cells sparse, irregularly dispersed or in small groups, coccoid green alga, 3–6 m in diameter, additional cells of Trentepohlia mostly on and near the surface, cyanobacteria scattered throughout the thallus but also completely endolithic below the lichen thallus. Perithecia: sessile; exciple 200 m in diameter; periphyses soon vanishing; involucrellum absent; asci 64–75 × 17–25 m; ascospores 3-septate, 18–27 × 7–10 m. (FRORJ\: on periodically inundated limestone.

140

.H\VWRWKHVSHFLHVDQGVSHFLHVSUR¿OHV

Fig. 93. Thelidium circumspersellum (f. lojkanum): ×25. 'LVWULEXWLRQ only known from the type locality in Hungary. 6LPLODUVSHFLHV Thelidium circumspersellum is probably only a depauperate form of T. zwackhii which has larger ascospores and asci as well as a well developed thallus.

7KHOLGLXPIRQWLJHQXP $0DVVDORQJR¿J 85, 94) Synonyms: Spermatodium fontigenum (A. Massalongo) Trevisan 1860, Involucrothele fontigena (A. Massalongo) Servít 1954, Sagedia zwackhii ff WR¿FROD Hepp 1857, Sagedia cataractarum Hepp 1857, Thelidium cataractarum (Hepp) Lönnroth 1859, Verrucaria cataractarum (Hepp) Leighton 1871, Involucrothele cataractarum (Hepp) Servít 1954, Sagedia riparia Hepp 1857, Spermatodium riparium (Hepp) Trevisan 1860, Thelidium riparium (Hepp) Zschacke 1920, Thelidium rodnense Zschacke 1920. 7KDOOXV white, whitish grey to greyish-green, occasionally with red, K+ purple patches; thin, proso- to paraplectenchymatous; basal layer absent; photobiont a coccoid green alga, cells ± clustered, 4–7 m in diameter. Perithecia: immersed to semi-immersed in the thallus; exciple 200–320 m in diameter, naked, with a thin apical involucrellum reaching 30–50 % of the height of the excipulum, laterally extending into the thallus, 200–360 m in diameter; asci 58–113 × 13–50 m; ascospores 3-septate (young ascospores simple septate), 22–36 × 10–14 m. (FRORJ\ damp to amphibious places on alkaline substrata (limestone, calciferous sandstone, silt covered wood). 'LVWULEXWLRQ widely distributed but rare all over Central Europe from low mountain ranges to (sub-) alpine altitudes.

Thelidium

141

Fig. 94. Thelidium fontigenum: ×20. 6LPLODUVSHFLHV Thelidium zwackhii lacks an involucrellum and the perithecia are usually largely exposed to almost sessile on the thallus surface.

7KHOLGLXPLQXQGDWXP =VFKDFNH¿J95) 7KDOOXV whitish to yellowish-grey; thin, almost invisible, semi-endolithic to endolithic; cortex with tightly interwoven, brownish hyphae; lower parts of the thallus with large oil cells (up to 60 m in diameter); photobiont coccoid green alga, aggregated in small groups. Perithecia: completely immersed in the sub-

Fig. 95. Thelidium inundatum: ×19.

142

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stratum; 140–230 m in diameter; exciple brown-black; without involucrellum; asci clavate; ascospores 3-septate, 32–36 × 10–14 m. (FRORJ\ amphibious in small streams on calcareous rocks. 'LVWULEXWLRQ only known from the type locality in Romania (stream Balea close to the village Vukan) and a recent collection from northern Italy but possibly overlooked. 6LPLODUVSHFLHV The terrestrial species Thelidium arnoldii and the common T. incavatum have larger ascospores and perithecia. Thelidium incavatum is occasionally found in ephemeral streams or in the splash water zone.

7KHOLGLXPNOHPHQWLL 6HUYtW¿J 96) 7KDOOXV ZKLWLVKJUH\ VHPLHQGROLWKLF WKLFN ±—P  ¿QHO\ ULPRVH WR areolate; cortex 10–25 m, colourless; basal layer absent; photobiont a coccoid green alga, cells rounded, 4–9 m, dispersed. Perithecia: semi-immersed in the thallus, naked; exciple 200 m in diameter; periphyses up to 40 m long; involucrellum absent or apical and much varying in thickness; asci 70–80 × 25–28 m; ascospores mostly uniseptate (a few unicellular ones also present), rounded at one apex, the other acute, 17–30 × 10–15 m. (FRORJ\ on temporarily inundated calcareous rocks with high content of silica. 'LVWULEXWLRQ only known from the type locality which has been destroyed after the construction of a large reservoir. Whether the species has survived at other ORFDOLWLHVQHHGVWREHFRQ¿UPHG

Fig. 96. Thelidium klementii: ×22.

Thelidium

143

6LPLODUVSHFLHV Thelidium rivulicolum has similarly thick thalli but much larger perithecia and asci, the ascospores are slightly larger, and the thallus does not penetrate into the substratum. 7D[RQRPLFDOQRWH Servít (1954) did not mention any involucrellum in his original description, but sections of the type specimen revealed that at least in some perithecia a distinct apical involucrellum is present.

7KHOLGLXPODKPLDQXP/RMNDH[=VFKDFNH¿J 97) Synonym: Involucrothele lahmiana (Lojka) J. Nowak & Tobolewski 1975. 7KDOOXV black-brown; thin, rimose; paraplectenchymatous; cortex pale; basal layer absent; photobiont a coccoid green alga, cells sparse, rounded, 4–7 Pm in diameter. Perithecia: sessile, exciple 200–350 Pm in diameter; periphyses up to 20 Pm; involucrellum adpressed to the exciple, thin and hardly discernable, extending to the thallus base; asci 66–70 × 22–25 Pm; ascospores uniseptate, 16–22 × 7–10 Pm. (FRORJ\ on inundated granite boulders. 'LVWULEXWLRQ only known from the type locality in the Koprova valley (Tatra, Slovakia). 7D[RQRPLFDOQRWHV This poorly known taxon is in need of revision. It may be FRQVSHFL¿FZLWKWKHWHUUHVWULDOThelidium tiroliense from calcareous rocks. Based on the information given by Zschacke (1920), the two taxa differ only in their different substratum and in the thallus colour.

Fig. 97. Thelidium lahmianum: ×23.

144

.H\VWRWKHVSHFLHVDQGVSHFLHVSUR¿OHV

7KHOLGLXPPHWKRULXP1\ODQGHU +HOOERP¿JV 88, 98–99) Basionym: Verrucaria methoria Nylander 1860. Selected synonyms: Involucrothele methoria (Nylander) Servít 1955, Sagedia aeneovinosa Anzi 1874, Verrucaria aeneovinosa (Anzi) Nylander 1882, Thelidium aeneovinosum (Anzi) Arnold 1868, Involucrothele aeneovinosa (Anzi) Servít 1954, Thelidium aeneovinosum var. kutakii Servít 1936, Involucrothele kutakii (Servít) Servít 1954, Polyblastia kutakii (Servít) Servít 1954, Thelidium kutakii (Servít) Servít 1946. Uncertain species: Thelidium diaboli A. Massalongo 1855, Verrucaria diaboli (A. Massalongo) Körber 1855. 7KDOOXV (black-) brown to grey; thin to moderately thick (25–80 m), continuRXV RU ¿QHO\ ULPRVH SDUDSOHFWHQFK\PDWRXV RIWHQ “ VXEJHODWLQRXV DQG ZLWK transparent margins when wet; thalli from sunny locations usually with a distinct brown cortex layer; basal layer absent; photobiont coccoid green algae (Elliptochloris bilobata, Stichococcus sp. in cultivated isolates), cells dispersed, 5–7 (12) m in diameter. Perithecia: prominent, at least basal parts often covered by a thin thallus mantle, upper part usually naked; exciple 228–465 m in diameter; periphyses up to 60 m long; involucrellum usually reaching far down, 430–1600 m in diameter; 40–100 m thick, ratio of involucrellum thickness to exciple diameter < 0.33; ascospores uniseptate, 24–46 × 13–21 m. (FRORJ\ amphibious on siliceous rocks in cool streams and rivers, but usually not submerged for more than a few months (Keller & Scheidegger 1994). Terrestrial populations exist in regions with high precipitation. Usually on well lit to semi-shaded locations in upland areas. 'LVWULEXWLRQ widely distributed at subalpine to alpine altitudes, very rare in high mountain ranges (e.g. Black Forest, Germany) and here on the verge of extinction (global warming, shading due to reforestation of old meadows).

Fig. 98. Thelidium methorium: type specimen from Karelia (Russia); ×15.

Thelidium

145

Fig. 99. Thelidium methorium: specimen from the Alps (Switzerland); ×15. 6LPLODUVSHFLHV Thelidium submethorium differs in the smaller ascospores and perithecia. Thelidium pluvium has perithecia which are covered by a thallus mantle. Thelidium pyrenophorum Mudd is a predominately terrestrial species but occasionally present at microsites that are submerged for short periods of time. It differs from T. methorium in the whitish to greyish thallus colour and smaller ascospores (19–32 m). Thelidium schibleri Zschacke is a non-amphibious species from calcareous rock with ascospores of similar size but smaller perithecia (240–480 PPSHUVFRPP$2UDQJH :KHWKHULWLVFRQVSHFL¿FZLWK T. methorium requires further studies. Uncertain species: Thelidium diaboli is treated by Zschacke (1933) as a synonym to T. aeneovonisum (=T. methorium) but no authentic material of this taxon was seen by us and the identity of T. diaboliLVLQQHHGRIFRQ¿UPDWLRQ

7KHOLGLXPPLQXWXOXP .|UEHUVO ¿JV89, 100) Synonyms: Verrucaria mesotropa Nylander 1866, Thelidium mesotropum (Nylander) A.L. Smith 1911, Arthopyrenia mesotropa (Nylander) Arnold 1870, Thelidium minutulum W. Stahl 1877, Verrucaria minutula (W. Stahl) Reinke 1895, Thelidium acrotellum Arnold 1866, Thelidium excile Arnold 1882, Thelidium hospitum Arnold 1882, Verrucaria intercedens var. aethioboloides Nylander 1861, Thelidium aethioboloides (Nylander) Vainio 1921 nom. inval. (non Zschacke 1920), Thelidium parvulum Arnold 1882, Thelidium eitneri Zahlbruckner 1921, Involucrothele eitneri (Zahlbruckner) Servít 1954, Thelidium terrestre Walt. Watson 1917, Thelidium viride Eitner 1910 non (Deakin) Zahlbruckner 1921. 7KDOOXV grey, light brownish, greenish; thin to thick (30–100 m), very variable in thallus appearance from almost continuous to tartareous-rimose or granular; hyphal arrangement proso- to paraplectenchymatous; photobiont coccoid green alga, cells 4–9 m in diameter, in aggregated small groups. Perithecia: usually

146

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Fig. 100. Thelidium minutulum: ×22. prominent and naked, occasionally semi-immersed or basal parts covered by a thin layer of algae containing thallus; exciple 100–300 m in diameter, brownish in upper part; periphyses up to 30 m; involucrellum absent, rarely thin (up to 30 m) and covering the upper exposed half of the exciple, 245–300 m in diameter, eventually spreading laterally; asci up to 90 × 23 m; ascospores 1 (2)septate, 13–32 × 4–15 m. (FRORJ\ terrestrial species, occasionally in the splash water zone of streams and rivers on limestone or basic, siliceous rocks (esp. metamorphites). 'LVWULEXWLRQ terrestrial populations widely distributed and locally abundant, amphibious populations much rarer and possibly overlooked. Known records from temporarily inundated sites cover localities from low mountainous to subalpine altitudes. 6LPLODUVSHFLHV Thelidium rehmii grows on sandstone, the photobiont cells are usually evenly distributed in the thallus (often densely packed, but not aggregated in small groups), and small ascospores (< 20 m) are generally absent. Although both species are genetically well separated (Thüs & Nascimbene 2008), some morphs of T. minutulum cannot be distinguished from T. rehmii with certainty based on morphological characters alone. The rare species T. submethorium has a dark coloured thallus (brown to olive), usually larger perithecia and ascospores. The involucrellum is always distinct, and the photobiont cells are irregularly dispersed in the thallus.

7KHOLGLXPQLJULFDQV =VFKDFNH¿J 101) 7KDOOXV black-brown; thin (30–35 m), according to Zschacke (1912) with single photobiont cells protruding up to 100 Pm below the thallus surface (chasmoendoOLWKLF ¿QHO\ULPRVHK\SKDl arrangement paraplectenchymatous; cortex absent

Thelidium

147

Fig. 101. Thelidium nigricans: ×25. but uppermost part of the thallus yellowish to brownish coloured; black basal layer only in parts of the thallus developed, partially reduced to isolated black patches; photobiont coccoid green alga, mostly arranged in small groups, 5–8 m in diameter. Perithecia: ¿UVWLPPHUVHGLQWKHWKDOOXVEXW¿QDOO\XSSHUKDOIH[posed; exciple 190–230 m in diameter; periphyses up to 30 m, involucrellum 200–485 m in diameter, reaching the base of the thallus and in some perithecia connected with the black basal layer; asci 70 × 30 m; ascospores 3-septate, 22–40 × 10–15 m. (FRORJ\ on siliceous substrata in the splash water zone. 'LVWULEXWLRQ only known from the type locality (a stream in the vicinity of the lake Bullea in the southern Carpathians) and a recent collection from a small stream in Switzerland (Rosenlaui). 6LPLODU VSHFLHV distinguished from other species by the combination of the characters 3-septate ascospores, brown thallus, well developed involucrellum reaching the base of the thallus and the occasional presence of a black basal layer.

7KHOLGLXPSDSXODUH)ULHV $UQROG¿J 102) Basionym: Verrucaria papularis Fries 1831. Synonyms: Amphoridium papulare (Fries) A. Massalongo 1853, Amphoridium hochstetteri f. papulare (Fries) Servít 1954. Uncertain species: Thelidium suzaeanum (Servít) Servít 1925, Involucrothele suzaeana Servít 1953, Polyblastia epomphala (Nylander) Zschacke 1933. 7KDOOXV grey, brown, occasionally with whitish pruina, thin and ± semi-endolithic to thick and epilithic; cortex absent, but occasionally with brownish pig-

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Fig. 102. Thelidium papulare: ×17. ment in uppermost part of the thallus; black basal layer absent; photobiont coccoid green alga, cells 6–9 m, aggregated in small groups. Perithecia: semi-immersed, with the basal part in small pits in the rock; exciple 370–485 m diameter; periphyses up to 70 m long; involucrellum apical to reaching half of the exciple height, thick (60–85 m), 300–800 m in diameter, often extending laterally in the thallus; asci 90–110 × 36–43 m; ascospores (1) 3-septate, 29–65 × 12–18 (20) m. (FRORJ\ a terrestrial species extending into temporarily submerged sites; especially in limestone areas with high changes of water level, frequently on calcareous rocks, rarely on siliceous substrata, often in shaded locations. 'LVWULEXWLRQ frequent throughout the area, amphibious populations ± restricted to localities with cool micro-climate. 6LPLODU VSHFLHV The terrestrial species Thelidium pyrenophorum (Acharius) Mudd 1861 has perithecia of similar size, but the ascospores have only one septum, their size is smaller (19–32 × 9–14 m) and the thallus is usually whitish. It occurs occasionally at temporarily inundated river banks or in ephemeral streams and springs. All other species of Thelidium from freshwater habitats are separated by the combination of 3-septate ascospores, large perithecia and the presence of a distinct involucrellum. Uncertain species: Thelidium suzaeanum was separated by Servít (1925) on account of the wider ascospores (18–22 m) and a thicker, olive to brownish thallus. The perithecia never form pits in the substratum and in some parts of the thallus a thin black basal layer is present. In the type specimens, a considerable variation in ascospore width has been observed with ascospores 16–18 m wide making up one third of all spores measured. An olive thallus colouration was not observed in the type specimen, but the olive colour possibly vanished during the

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149

long storage. Brownish thallus colours are also present in some morphs of T. papulare which, however, have narrower ascospores. Whether the correlation of the characters “predominantly wide ascospores”, “brownish to olive thallus FRORXU´DQG³WKLFNXVXDOO\HSLOLWKLFWKDOOXV´LVJHQHWLFDOO\¿[HGKDVWREHHYDOXated by molecular studies. Zschacke (1933) distinguished Thelidium epomphalum (= Polyblastia epomphala) from T. papulare E\WKHVOLJKWO\ÀDWWHQHGSHULWKHFLD with a depressed ostiole, the presence of oil-cells and the occasional formation of a single additional, transversal ascospore septum. However, the shape of the perithecia and the ostiole varies even within a single specimen – especially in amphibious populations. $OWKRXJKERWKWD[DPD\EHGLI¿FXOWWRGLVWLQJXLVKE\PRUphology, molecular data based on recently collected specimens from Skandinavia indicate that T. papulare and T. epomphalum DUHRQO\GLVWDQWO\UHODWHG6DYLüHW al. 2008).

7KHOLGLXPSHUWXVDWLL *DUURYDJOLR -DWWD¿JV 87, 103) Basionym: Verrucaria pertusatii Garrovaglio 1865. 7KDOOXV grey, brown to brown-black; thin to rather thick (30–145 m), continuous to minutely rimose; paraplectenchymatous; cortex absent but uppermost part of thallus with brownish pigment; basal layer absent; photobiont coccoid green alga, 6 m in diameter. Perithecia: prominent and naked; exciple 230–295 m in diameter; periphyses up to 45 m; involucrellum 514–570 m in diameter, very thick (115–170 m), ratio of involucrellum thickness to exciple diameter > 0.4; asci not measured; ascospores uniseptate, fresh ascospores partly with halo, 26–40 × 12–19 m. (FRORJ\ amphibious but not permanently submerged in subalpine to alpine rivers and irrigated rocks.

Fig. 103. Thelidium pertusatii: ×18.

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'LVWULEXWLRQ southeastern and southern Central Alps, endemic (?). 6LPLODUVSHFLHV Thelidium pertusatii differs from the closely related T. methorium in the the ratio of involucrellum thickness to exciple diameter which is > 0.4.

7KHOLGLXPSOXYLXP 2UDQJH¿JV86, 104) 7KDOOXV greyish, brown, greenish; 35–85 m high, continuous or with small cracks; paraplectenchymatous; basal layer absent; photobiont a coccoid green alga, up to 8 m in diameter or elongated 9 × 5.5 m. Perithecia: immersed in prominent thallus warts; exciple 100–320 m in diameter; periphyses up to 40 m long; involucrellum apical to reaching the thallus base, often ventro-laterally spreading, leaving a transparent area between exciple and involucrellum, 150–500 m wide, thin (up to 30 m), covered by a distinct thalline mantle, upper part dark coloured, lower part fading, pigment K-; asci 75–100 × 25–30 m; ascospores uniseptate, 19–36 × 8–17 m. (FRORJ\ splash zone of small rivers and streams on siliceous rocks and pebbles, preferring shaded sites. 'LVWULEXWLRQ Alps, Black Forest, to be looked for in mountain ranges; also known from the British Isles, Norway and Tasmania. 6LPLODUVSHFLHV: Among the specialized freshwater species of the genus Thelidium pluvium is unique due to the presence of rather small, uniseptate ascospores, a thin involucrellum and perithecia covered by a thallus mantle. Thelidium fumidum has a dark brown thallus and it is found only occasionally at sub-aquatic sites. It has a well developed involucrellum reaching the base of the exciple and

Fig. 104. Thelidium pluvium: ×21.

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151

a black basal layer. The involucrellum has a reddish-brown pigment which turns grey in K. The representatives of the T. methorium-group have larger perithecia and a well developed, thicker involucrellum.

7KHOLGLXPUHKPLL=VFKDFNH¿JV 90, 105) Synonym: Involucrothele rehmii (Zschacke) Servít 1954. 7KDOOXV whitish-greyish to green-grey; thin to thick (15–110 m), tartareous, ZLWK¿QHFUDFNVSURVRWRSDUDSOHFWHQFK\PDWRXVFRUWH[DQGEDVDOOD\HUDEVHQW photobiont coccoid green alga, 3–10 m in diameter, aggregated in small groups. Perithecia: semi-immersed to almost sessile; exciple 200–300 m in diameter; periphyses up to 30 m; involucrellum completely missing or indistinguishable from an occasionally slightly thickened apical part of the exciple, never laterally spreading; asci 67–93 × 26–40 m; ascospores uniseptate, 19–30 × 8–15 m. (FRORJ\on siliceous rocks (sandstone) in humid and more or less shaded places, often in the vicinity of little streams but usually restricted to micro-sites without or with only short submersion periods. 'LVWULEXWLRQonly known from a few localities in southern Germany (Franconia, Odenwald, Black Forest). 6LPLODU VSHFLHV Occasionally, Thelidium minutulum has smaller ascospores (< 19 m), but morphs with ascospores of the same size as in T. rehmii are often IRXQG7KHVHIRUPVDUHGLI¿FXOWWRGLVWLQJXLVKIURPT. rehmii by morphological characters alone although both species are well separated genetically (Thüs & Nascimbene 2008). Thelidium minutulum appears to be restricted to well buff-

Fig. 105. Thelidium rehmii: ×22.

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ered substrata (limestone, metamorphic silicates) while T. rehmii is restricted to sandstone. The thallus algae in T. minutulum are usually arranged in distinctly aggregated groups, whereas the algal cells are irregularly dispersed throughout the thallus in T. rehmii.

7KHOLGLXPULYXOLFROXP (Nylander) Migula 1931 Basionym: Verrucaria rivulicola Nylander 1875. Synonym: ?Arthopyrenia rivulicola (Nylander) Arnold. 7KDOOXV whitish to greenish (yellowish in herbarium); epilithic, but often incrusted by tuffa and ± semi-endolithic, thick (100–200 m), continuous to rimose, with powdery surface; cortex colourless or absent; basal layer absent, photobiont coccoid green alga, cells of Trentepohlia sp. sometimes also present. Perithecia: semi-immersed to sessile, naked; exciple diameter 300–450 m; periphyses not seen (soon vanishing?); involucrellum absent; asci 90 × 35 m; ascospores uniseptate, 23–42 × 7–15 m. (FRORJ\ amphibious on calcareous rocks and pebbles. 'LVWULEXWLRQ so far only recorded from Germany (Bavaria) and France but most probably overlooked and actual distribution range still uncertain. 6LPLODU VSHFLHV Thelidium klementii has much smaller perithecia which often have an apical involucrellum. Thelidium zwackhii has 3-septate ascospores.

7KHOLGLXPVXEPHWKRULXP 9DLQLR =VFKDFNH¿JV106–107) Basionym: Verrucaria submethoria Vainio 1883. 7KDOOXVolive, brown or brown-grey; thin to thick (10–130 m), continuous or ¿QHO\ULPRVHSDUDSOHFWHQFK\PDWRXV“VXEJHODWLQRXVZKHQZHWDQGZLWKWUDQVparent margins; cortex absent but upper part of thallus occasionally with brownish pigment; basal layer absent; photobiont coccoid green alga, irregularly arranged in the thallus. Perithecia: prominent; exciple 150–275 m in diameter; periphyses up to 30 m; involucrellum apical or reaching the thallus base, 250–510 m in diameter, usually well developed but < 60 m thick at medium height, ratio of involucrellum thickness to exciple diameter < 0.33; asci not measured; ascospores uniseptate, perispore lacking, 26–39 × 9–18 m. (FRORJ\ in clean streams and rivers of high mountain ranges on siliceous substrata. 'LVWULEXWLRQ beside the type locality in Finland only known from historical records from the Black Forest (Germany) and a few recent collections from northern Italy. The recent, though very scarce records from unpolluted, cold streams indicate that the species is on the verge of extinction in Central Europe.

Thelidium

153

Fig. 106. Thelidium submethorium: Russia, type specimen from Karelia; ×15.

Fig. 107. Thelidium submethorium: Italy, specimen from the Alps; ×15. 6LPLODUVSHFLHV Thelidium methorium has larger perithecia and ascospores. Thelidium pertusatii has a much thicker involucrellum and larger ascospores. Thelidium pluvium has perithecia which are covered by a thin thallus mantle and greenish or greyish colours. Thelidium minutulum is a skiophilous terrestrial species with a greyish to green thallus, a very thin or virtually lacking involucrellum ZKLFKLVPRVWO\FRQÀXHQWZLWKWKHH[FLSOHDQGDOJDHWKDWDUHXVXDOO\DUUDQJHGLQ small groups.

154

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7KHOLGLXP]ZDFNKLL+HSS $0DVVDORQJR¿J108) Basionym: Sagedia zwackhii Hepp 1853. Synonyms: Segestria zwackhii (Hepp) Trevisan 1860, Verrucaria zwackhii (Hepp) Garovaglio 1865, Pyrenula zwackhii (Hepp) Hepp 1867, Verrucaria microcarpa Davies ex Leigthon 1851, Thelidium microcarpum (Davies ex Leighton) A.L. Smith 1911, Verrucaria sparsulum Nylander 1877, Thelidium sparsulum (Nylander) A.L. Smith 1911, Verrucaria xylospila Nylander 1886, Thelidium xylospilum (Nylander) Zschacke 1920, Thelidium subgelatinosum Zschacke 1920. 7KDOOXV greyish, whitish to brownish; thin (35–65 m); proso- to paraplectenchymatous; cortex absent; occasionally with a basal layer devoid of algae; photobiont coccoid green alga, arranged in small aggregated groups. Perithecia: DW ¿UVW “ LPPHUVHG LQ WKH WKDOOXV EXW ¿QDOO\ XVXDOO\ DOPRVW VHVVLOH H[FLSOH 100–260 m in diameter; periphyses up to 20 m long; involucrellum absent but occasionally exciple swollen in uppermost third (up to 23 m thick); asci 90–120 × 29–40 m; ascospores: 3-septate, 20–36 × 9–15 m. (FRORJ\: a predominantly terrestrial species also found at amphibious sites, absent from permanently submerged sites; on a wide spectrum of substrata, often as a pioneer species on silt incrusted limestone, lime-impregnated sandstone or hard wood but usually avoiding acidic substrata. 'LVWULEXWLRQ so far rarely recorded but easily overlooked and presumably widely distributed. 6LPLODUVSHFLHV Among the species with 3-septate ascospores, Thelidium fontigenum is most similar differing by the presence of a thin involucrellum. Thelidium aethioboloides and T. suzaeanum have an olive-black to dark green-grey thallus and well developed involucrella. Thelidium circumspersellum and T. aquaticum have smaller asci and ascospores.

Fig. 108. Thelidium zwackhii: ×24.

Theligyna

155

7KHOLJQ\DA. Massalongo 1855 Important references: Henssen (1963a, 1966), Henssen & Jørgensen (1990), Jørgensen (2007b).

7KHOLJQ\DOLJQ\RWD (Wahlenberg in Acharius) P.M. Jørgensen & +HQVVHQ¿J109) Basionym: Verrucaria lignyota Wahlenberg in Acharius 1803. Selected synonyms: Arctoheppia scholanderi Lynge 1938, ?Collemopsis fuliginascens Nylander 1873, Fernaldia scholanderi Lynge 1937 nom. illeg., Porocyphus dispersus E. Dahl 1950, ?Porocyphus fuliginascens (Nylander) Couderc ex Crozals 1910, Porocyphus ocellatus (Th. Fries) Henssen 1989, ?Psorotichia fuliginascens (Nylander) Forssell 1885, Pyrenopsis ocellata Th. Fries 1866, ?Pyrenopsis umbilicata Vainio 1881, Verrucaria fuliginea (Acharius) Wahlenberg 1812. 7KDOOXV black to very dark olive-brown, usually only few millimetres in size, forming irregularly shaped minutely stalked squamules, often slightly glossy, squaPXOHV XVXDOO\ FRQÀXHQW WR VRPHWLPHV VFDWWHUHG   ±  PP ZLGH c. 0.15–0.5 mm thick, irregularly shaped, plane to uneven, surface rough, somewhat glossy or dull, attached to the substrate by a minute, short, irregular stalk or basal gelatinous holdfast; thallus anatomy ± paraplectenchymatous with hyphae forming irregular network, hyphal cells short to slightly elongated, roundish to angulate, 5–10 × ±—P SKRWRELRQW D ¿ODPHQWRXV F\DQREDFWHULXP 5LYXOD-riaceae) but ¿ODPHQWV RIWHQ PXFK FRQWRUWHG DQG VSOLW ¿ODPHQWV VRPHWLPHV EHWWHU YLVLEOH WRwards the base of the areoles. Apothecia: 1–10 per areole, small, 0.25–0.4 (0.6) mm, immersed to semi-immersed, disc black, pore-like to slightly opened and usually soon umbonate, thallus margin persisting, thin, (10) 20–35 (50) m thick, smooth, not prominent but sharply separated from the disc; proper exciple thin, 10–15 m wide, hyaline, apically thickened, 15–30 m wide, bluish green; hymenium hyaline to apically faintly bluish green, 65–100 m high, I-, soon with intrusions of agglutinated hyphae forming an umbo; epihymenium distinctly bluish green, K-, N-, paraphyses sparse, irregularly branched and anastomosing, apical cells slightly thickened; asci often irregularly narrow clavate, thin walled, I- or faintly yellowish, 8-spored; ascospores simple, hyaline, broad ellipsoid, 7.5– 12 × 5.5–9 m. Pycnidia: immersed, simple, globose, ca. 50 m wide, ostiole bluish green coloured; pycnospores cylindrical to subglobose, 2.5–3.5 × 1.5–2 m. (FRORJ\ on moist siliceous rock in cool, shady places and frequently inundated on rocks and boulders in cool mountain and glacier streams; in Fennoscandia on moist rock in shady places, also along rocky lake shores and river banks. 'LVWULEXWLRQ very rare in mountainous Central Europe and subalpine regions in the Alps, Auvergne; more common in Fennoscandia, also in Greenland and arctic-boreal North America. Records reported from warm-temperate to arid regions or from dry, exposed localities should be treated cautiously. 6LPLODU VSHFLHV A collection from the Mont Blanc region growing frequently submerged on granitic rock was reported by Crozals (1910) as Porocyphus fuligi-

156

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Fig. 109. Thelignya lignyota: thallus areolate to small-squamulose with numerous umbonate discs; ×14. nascens. Judging from the description it most certainly belongs to Thelignya lignyota. According to Santesson et al. (2004) Psorotichia fuliginascens described from Finland remains to be further studied. However, Forssell (1885) already suggested that it may belong to Thelignya.

Verrucaria Schrader 1794 nom. cons. Notes: The genus Verrucaria is a polyphyletic assemblage as shown by a recent multigene study (Gueidan et al. 2007). Because of the high amount of homoplasy LQ FUXVWRVH 9HUUXFDULDOHV D UHHPHQGDWLRQ RI WKH JHQXV UHÀHFWLQJ UHVXOWV IURP QRYHOPROHFXODUVWXGLHVLVYHU\GLI¿FXOWWRDFKLHYH$VD¿UVWVWHSZHKDYHIROlowed the separation of the new genus Hydropunctaria from the remainder of Verrucaria as proposed by Gueidan et al. (2008) since Hydropunctaria forms a well separated group at molecular level and is corrobated by a unique set of character states allowing feasible recognition based on morphological characters. Nonetheless, the phenotypic plasticity of the species of the new genus is very high and some specimens may prove indeterminable as either Hydropunctaria or Verrucaria when using the diagnostic characters adopted in the main key. For such specimens a cross reference to the species key of Hydropunctaria was included in this key for Verrucaria s.l. Important references: Clauzade & Roux (1985), Gueidan et al. (2007), Gueidan et al. (2008), Keller (1996), Krzewicka, B. & Kizka, J. (2007), Nascimbene et al. (2007), Orange (2000, 2002, 2004), Servít (1954), Thüs (2002, 2006), Zschacke (1933). Key to the species: 1a Thallus entirely endolithic, perithecia almost completely immersed in pits in the (calcareous) substratum, predominantly terrestrial species, in temporary

Verrucaria

E 2a E E 3a 4a E 5a E 6a E 7a E

8a E 9a E 10a E 11a E 12a

157

streams, at river banks with high changes in water levels, in shady localities in or close to springs in cool water or on irrigated rocks ............................. .................................................................. Verrucaria div. spp. (not treated) Thallus at least partly epilithic on siliceous or calcareous substrata, most of the perithecia not immersed in pits in the (calcareous) substratum ........... 2 Average ascospore length < 16 Pm ........................................................... 3 Average ascospore length > 16 Pm ........................................................... 5 Thallus thin to thick, perithecia immersed in the thallus or little projecting above, if projecting then tips of the perithecia not covered by a thin thallus mantle, ascospores ± ellipsoid ............. Hydropunctaria (see species key) Thallus thin (< 60 Pm), perithecia projecting above the thallus, covered by a thin thallus mantle, ascospores ± globose to ellipsoid .............................. 4 Mature asci with eight ascospores, exciple 80–150 Pm, ascospores 5–11 Pm, thallus blackish green or greenish (shade morphs), occasionally brownish .................................................................................................... 9DTXDWLOLV Mature asci with four ascospores, exciple 140–290 Pm, ascospores 9–15 Pm, thallus dark olive-greenish or dark greenish grey ...................... V. madida Involucrellum absent or thin and completely enveloping and hardly discernable as distinct from the exciple, always with black or brown exciple/involucrellum base ........................................................................................... 6 Involucrellum present, apical to reaching the thallus base, distinctly separated from the exciple, exciple base transparent or brown ........................ 8 Thallus semi-endolithic, on carbonaceous rocks, ascospores up to 18 Pm long .......................................................................................... V. austriaca Thallus epilithic, on siliceous rocks, ascospores 16–28 Pm long .............. 7 Perithecia prominent, upper half often naked, thallus thin and granular, at shaded localities, in ephemeral watercourses or on temporary wet rocks, usually rarely inundated .............................................................. V. consociata Perithecia usually immersed in the thallus for more than 50 %, thallus thick, often forming a continuous or rimose crust, in shaded to sun-exposed localities, either amphibious in the splash water zone of small streams and rivers or riparian in rarely inundated places at the banks of usually large rivers .... 29 Average ascospore length > 25 Pm ........................................................... 9 Average ascospore length < 25 Pm ......................................................... 13 Involucrellum apical to reaching the thallus base, not or little laterally extruding in the thallus, apical part of the involucrellum naked ................. 10 Involucrellum usually reaching the thallus base, laterally distinctly extruding in the thallus, apical part of the involucrellum usually covered by a thin thallus mantle (V. margacea VO) ............................................................. 11 On siliceous substrata, thallus greyish to brown, (fresh) ascospores halonate ................................................................................................... 9ODWHEURVD On calcareous or siliceous substrata, thallus whitish to yellowish grey, ascospores never halonate ......................................................... 9VXEPHUVHOOD Ascospores broadly ellipsoid to ovoid, length-width ratio 2.0, always on calcareous rocks .............................. 9PDUJDFHDDJJU9HODHRPHODHQD Ascospores ellipsoid to elongated, length-width ratio usually > 2.2, on siliceous or calcareous rocks ........................................................................ 12 Mature asci 60–80 Pm long, thallus very thin (up to 20 Pm), greyish-green to olive or brownish, exclusively on siliceous substrata, impossible to distinguish with certainty from V. margacea s.str. without ripe asci ..................... ............................................................. 9PDUJDFHDDJJU9DQGHVLDWLFD

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E Mature asci > 90 Pm long, thallus thin to moderately thick (20–70 Pm), usually brownish, in upland areas also black-brown, on siliceous or (rarely) calcareous substrata ....................................................... V. margacea s.str. 13a Wet thallus with a consistency like “solid jelly” (subgelatinous), margins often becoming ± transparent when wet, surface smooth (except for projecting perithecia), usually not cracked or with some large and wide cracks, involucrellum reaching the thallus base ..................................................... 14 E Wet thallus not with a consistency like “solid jelly” (not subgelatinous), margins never becoming transparent when wet, surface smooth or uneven, usually with numerous cracks or areolate, involucrellum reaching the thallus base or ± reduced to apical ...................................................................... 17 14a On calcareous rocks, ascospores broadly ellipsoid to ovoid ........................ ......................................................... 9PDUJDFHDDJJU9HODHRPHODHQD E On siliceous substrata, ascospores ellipsoid to elongated ......................... 15 15a Thallus black or olive-green (in the sun) to greenish-grey (shaded microsites), with blackish-green cortical pigment, greenish in shady localities and here cortical pigment inconspicuous to absent, thallus thick, ascospores from freshly collected specimens halonate, perispore swelling in K, ascospores rather short (18–20 m, rarely up to 23 m), perithecia usually immersed in the thallus or little projecting, in Central Europe only at high altitudes ....... V. pachyderma E Thallus brown, brown-black, orange-brown, brownish-grey, greenish, blackish-green or grey, cortical pigment yellowish-brown to brown black or absent, thallus thick to thin, ascospores from freshly collected specimens halonate or not, ascospores larger (usually 20–25 m), perithecia immersed in the thallus or projecting, from near sea level to alpine areas .................. 16 16a Thallus usually thin (10–60 m), with projecting perithecia, ascospores never halonate, a colourless angle of thallus tissue between lower part of the involucrellum and the base of the excipulum with ..................... 9K\GUHOD E Thallus thick (usually > 50 m), with immersed or projecting perithecia, ascospores from freshly collected samples halonate, without colourless angle of thallus tissue between lower part of the involucrellum and the base of the excipulum, cell walls brown throughout ..................................... V. funckii 17a Perithecia distinctly projecting, thallus thin, tartareous or granular, with or without cracks, thallus not areolate (except areas around the perithecia) ... 18 E Perithecia immersed or semi-immersed, thallus usually thick, a continuous crust with ± numerous cracks or areolate ................................................ 22 18a Involucrellum apical to reaching the lower half of the exciple (very rarely stretching to the thallus base), thallus whitish, scabrid, grey or brownish, on carbonaceous rocks or other substrata with alkaline surfaces, never on acidic surfaces ......................................................................... 9VXEPHUVHOOD E Involucrellum usually reaching down to the thallus base, thallus greenish, greyish or brownish, warty, granular to scabrid, on acidic (siliceous) or alkaline surfaces ............................................................................................ 19 19a Perithecia immersed in thallus warts, involucrellum covered by a thin thallus mantle (at least 50 % of the perithecium), amphibious to terrestrial species .... 20 E Perithecia naked, involucrellum not covered by a thallus mantle (at least 75 % naked), terrestrial (riparian), rarely inundated ................................ 21 20a On calcareous substrata, brown coloured cells between involucrellum surface and exciple base, thallus semi-endolithic to epilithic ......... 9FDOFDULD E On siliceous substrata, triangular area with transparent cells between the thin and sharply delimited involucrellum and the base of the exciple, thallus always epilithic .......................................................................... 9K\GUHOD

Verrucaria

159

21a Ascospores 15–28 × 8–15 Pm, at sunny to shady localities ......................... .............................................................................. V. muralis Acharius 1803 E Ascospores 11–23 × 4–8 (10) Pm, ± restricted to shady localities .......9GRORVD 22a Thallus with black basal layer ................................................................. 23 E Thallus lacking black basal layer ............................................................ 24 23a Thallus grey, whitish, pink, greenish, orange or brown, hypothallus (if present) whitish (do not mistake for damaged areas where the black basal layer may be visible!), hydrophilic species ................................ V. praetermissa E Thallus brown, hypothallus (if present) black, terrestrial species, but frequently found along well lit river banks in microsites not inundated or only submerged for short periods during the year, never in frequently wetted localities ........................................................... V. nigrescens Persoon 1795 24a Involucrellum distinct and reaching the lower half of the exciple .......... 25 E Involucrellum apical, missing or, if reaching below, the lower half of the exciple fading light brown ...................................................................... 27 25a Carbonised involucrellum cells with small lumina, involucrellum deep black throughout or “dotted” in sections, exciple in lower parts black-brown to transparent ............................................................................. 9DHWKLREROD E Carbonised involucrellum cells with distinctly enlarged lumina especially in lower parts, cell walls deep brown, involucrellum with “mesh-like” appearance in thin (!) sections, exciple in lower parts always transparent ........ 26 26a Perithecia immersed in the thallus, < 25 % naked and raised above the thallus, involucrellum dark throughout, involucrella of neighbouring perithecia often fused, hydrophilic species ........................................ V. praetermissa E Perithecia semi-immersed, usually > 25 % naked and raised above the thallus, involucrellum dark throughout to transparent in lower parts, neighbouring involucrella rarely fused, never with black basal layer, predominantly terrestrial species from shaded sites with high air humidity......... V. elaeina Borrer 1812 27a Involucrellum distinct but mostly apical, usually thick, thallus with even surface, usually strongly rimose to areolate ............................................ 28 E Involucrellum absent or weak and fading in lower parts, thallus surface usually uneven, continuous, rimose-areolate to granular, thallus margins continuous, crenulate to minutely lobate ...................................................... 29 28a Thallus whitish to light brownish, somewhat cracked or continuous, involucrellum not spreading, on calcareous or lime-incrusted siliceous substrata ............................................................................................. 9VXEPHUVHOOD E Thallus brownish to greyish, thallus rimose to areolate, involucrellum variable, adjacent to the exciple to slightly spreading ventro-laterally, exclusively on siliceous substrata ................................................... 9DHWKLREROD 29a Ascospores large 23–35 (40) Pm, thallus areolate to granular, thallus margins never crenulate to squamulose or minutely lobate; terrestrial species, occasionally on river banks at rarely submerged sites ................................. ............................................................ V. viridula (Schrader) Acharius 1803 E Ascospores small (16–20 Pm), thallus continuous to rimose, margins continuous, crenulate or squamulose to minutely lobate; amphibious species ..... 30 30a Thallus (especially at the margins or near cracks) with additional apothecia of Bacidina inundata or Ionaspis lacustris .................................................. ..................... mechanical hybrids with either 9VXEOREXODWD or 9K\GUHOD E Thallus lacking additional apothecia of Bacidina inundata or Ionaspis lacustris ....................................................................................... 9VXEOREXODWD

160

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Fig. 110

Fig. g 111

Fig. 112 2

Fig. 113

Fig. 115

Fig. 114

Fig. 116

Plate 5. Figs. 110– 0 116. ShapeRILQYROXFUHOOXP in Verrucaria I. Fig. 110. V. consociata±GDUNH[FLSOHLQYROXFUHOOXPDEVHQWW\SHVSHFLPHQ  Fig. 111. V. submersella. Fig. 112. V. sublobulata. Fig. 113. V. aethiobola±W\SHVSHFLPHQ Fig. 114. V. aethiobola. Fig. 115. V. calcaria±WKDOOXVVHPLHQGROLWKLFVXEVWUDWXPSDUWLFOHVLQJUH\ LVRW\SHVSHFLPHQ  Fig. 116. V. dolosa.

9HUUXFDULD

Fig 117 Fig.

Fig. 119

Fig 121 Fig.

Fig. g. 118

Fi 120 Fig.

Fig. 122

Plate 6. Figs. 117– 7 122. Shape RILQYROXFUHOOXP in Verrucaria II. Fig. 117. 7 V. hydrela ±W\SHVSHFLPHQ Fig. 118. V. hydrela (“V. denudata”). Fig. 119. V. funckii ±W\SHVSHFLPHQ Fig. 120. V. funckii. Fig. 121. V. pachyderma±KRORW\SHVSHFLPHQ Fig. 122. V. pachyderma.

161

162

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Fig. ig. 123

Fig. g. 124

Fig 125 Fig.

Fi 126 Fig.

Fig. 127

Fig. Fi 128

Plate 7. Figs. 123–128. Shape RILQYROXFUHOOXP in Verrucaria III: Fig. 123. V. margacea VVWU±W\SHVSHFLPHQ Fig. 124. V. margacea VO³V. applanata”). Fig. 125. V. margacea VO V. andesiatica ±W\SHVSHFLPHQ  Fig. 126. V. margacea VOV. elaeomelaena ±OHFWRW\SHVSHFLPHQ  Fig. 127. 7 V. latebrosa ±LVRW\SHVSHFLPHQ Fig. 128. V. latebrosa ³V. anziana”).

Verrucaria

163

9HUUXFDULDDHWKLREROD$FKDULXV¿JV113–114, 129) Synonyms: Verrucaria margacea var. aethiobola Nylander 1861, Verrucaria aethiobola var. primaria Vainio 1921, Verrucaria laevata Acharius 1810, Verrucaria aethiobola var. griseocinerascens Vainio 1921, Verrucaria griseocinerascens Zschacke 1927, Verrucaria czernaensis Zschacke 1927, Verrucaria delita Nylander 1876, Verrucaria hibernica Zschacke 1927. Uncertain species: Verrucaria fuscorubens (de Lesdain) Zschacke 1927, Verrucaria pseudocatalepta Servít 1952. 7KDOOXV brown, grey, orange to pinkish; thick (40) 50–120 m, rimose to areolate; surface even; para- to prosoplectenchymatous; never subgelatinous; black EDVDOOD\HUDEVHQWORZVSHFL¿city for certain photobiont taxa, but always with coccoid green algae (e.g. Diplosphaera chodatii, Elliptochloris bilobata, Trebouxia s.str.YHUL¿HGE\FXltivated isolates), size much varying, uniformly dispersed in the thallus. Perithecia: immersed to little protruding; exciple 140–220 m in diameter; periphyses up to 20 m long; involucrellum apical to almost reaching the thallus base, ± adpressed to the exciple or laterally little protruding; cell lumina in lower part of the involucrellum not much enlarged, completely carbonised or with “dotted” pigmentation; apical part often naked and exposed over the surrounding thallus; asci 8-spored; ascospores without halo or halo little developed, and not or little swelling in K, (14) 18–25 (31) × 6.5–14 m. (FRORJ\ amphibious in the upper splash water zone of streams or rivers, tolerating long periods of desiccation. Best developed at sun-exposed sites but tolerating moderate shading. On slightly acidic to alkaline, siliceous, stable rocks. Rather tolerant to silting and moderate eutrophication. In regions with acidic rocks often close to bird perches. 'LVWULEXWLRQ scattered from sea-level to high-montane regions; locally abundant in montane regions, rare in subalpine areas.

Fig. 129. Verrucaria aethiobola: ×19.

164

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6LPLODU VSHFLHV Verrucaria praetermissa LV VXSHU¿FLDOO\ YHU\ VLPLODU EXW WKH involucrellum usually reaches the thallus base and is much more laterally spreading into the thallus with distinctly enlarged cells in the lower part. In V. praetermissa, the dark coloured cell walls of the involucrellum cause a mesh-like appearance, whereas in V. aethiobola the involucrellum appears densely black or dotted and rarely mesh-like. Involucrella of neighbouring perithecia of V. praetermissa DUHRIWHQIXVHGRUFRQÀXHQWZLWKDEODFNEDVDOOD\HU7KHOHQJWKZLGWK ratio of the ascospores of V. praetermissa is higher than in V. aethiobola. Verrucaria latebrosa is an upland species with larger ascospores and perithecia. Fresh ascospores have a thick halo which is considerably swelling in K. Verrucaria submersella has a light coloured thallus and semi-circular, prominent perithecia with an involucrellum which is not laterally spreading. It has smaller perithecia and a very thin to fading or missing involucrellum. Verrucaria hydrela usually has a thin subgelatinous thallus. Morphs of V. hydrela with thicker, non subgelatinous thalli are separated from V. aethiobola by the much thinner and always spreading involucrellum which leaves a transparent angle between involucrellum and exciple base. Uncertain species: Verrucaria pseudocatalepta Servít differs from V. aethiobola s.str. in the poorly developed involucrellum, which completely envelops the exciple, including the exciple base. Verrucaria fuscorubens is distinguished from V. aethiobola by the reddish brown thallus colour and the shape of the involucrellum which is closely attached to the exciple and not laterally spreading.

9HUUXFDULDDTXDWLOLV0XGG¿JV 130, 167) Synonyms: Lithoicea aquatilis (Mudd) Stein 1879, Verrucaria vitricola Nylander 1887, Verrucaria retecta Zschacke 1927, Verrucaria schistosa Servít 1950. Uncertain species: Verrucaria rechingerii Servít 1950.

Fig. 130. Verrucaria aquatilis: ×25.

Verrucaria

165

7KDOOXV black, black-green, brown-black when dry (morphs from shady locations also green or brownish); black to green when wet, thin (10–50 m); paraSOHFWHQFK\PDWRXVDQGVXEJHODWLQRXVFRQWLQXRXVWR¿QHO\ULPRVHSKRWRELRQWD JUHHQDOJDLVRODWHVZHUHUHSHDWHGO\LGHQWL¿HGDV 'LODEL¿OXP spp.), in the thallus with rounded or elongated cells, 4–9 m in rounded cells, up to 12 m in elongated cells, occasionally arranged in short vertical columns. Perithecia: in prominent thallus warts; exciple 80–130 × 80–150 m; involucrellum usually ± reaching the exciple base and ± covered by a thin thallus layer, 140–430 m in diameter; asci 8-spored 21–26 × 14–21 m; ascospores unicellular, not halonate, 4.8–11 × 3.9–9.3 m, length/width-ratio 1.0–2.0. (FRORJ\frequently to permanently submerged, or wetted by splash water. Tolerant to a wide range of illumination. Colonizing even small pebbles in the unstable bedrock of brooks (living colonies sometimes cover complete rocks, including its ORZHUVXUIDFH DOVRIRXQGRQSODVWLFEDJVÀRDWLQJLQWKHFXUUHQW5HVWULFWHGWR watercourses and lakes with pH > 5.0. On calcareous as well as siliceous substrates. Tolerant to moderate silting in fast running watercourses, much more sensitive in slow waters and lakes. 'LVWULEXWLRQ from lowland to alpine areas, most frequent at lowland to montane altitudes. 6LPLODUVSHFLHV Verrucaria madida has 4-spored asci, larger perithecia and (in specimens from well lit sites) a characteristic olive pigment in the cortex. Hydropunctaria rheitrophila has ascospores with a larger length/width ratio, a usually ± apical involucrellum, a thicker, olive-greenish or orange to brownish thallus, and often small black patches in the thallus. Hydropunctaria scabra has a much thicker thallus, larger ascospores with a higher length/width ratio and usually a rough surface due to the presence of small black columns which are connected with a black basal layer. Uncertain species: Verrucaria rechingerii has similar globose ascospores like V. aquatilis, but the thallus is thicker and the perithecia are ± immersed in the thallus giving it a similar appearance as in Hydropunctaria rheitrophila. The involucrellum in thalli from authentic material of the Natural History Museum Vienna (Krypt. Exscicc. 1851) is variable from apical to reaching down the thallus base. A locally developed “black basal layer” (also depicted in Servít 1950) apparently consists of dead lichens which have been overgrown by new Verrucaria thalli.

Verrucaria austriaca Riedl 1990 Synonym: Verrucaria irrigua Zschacke 1927 non Taylor 1836. 7KDOOXV light grey; semi-endolithic, slightly rimose to almost areolate; photobiont coccoid green alga with rounded cells, 5–7 m in diameter. Perithecia: mostly immersed, 200–250 m in diameter; exciple black-brown in upper parts, colourless below; involucrellum lacking; asci 8-spored, 70–75 × 16–18 m; ascospores unicellular, 16–18 × 7.5–8 m.

166

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(FRORJ\ on irrigated calcareous rocks. 'LVWULEXWLRQ only known from the type locality (“Sonntagsberge bei Rosenau”) in Austria. 6LPLODU VSHFLHV In Verrucaria calcaria and V. submersella the exciple is protected by an involucrellum and the ascospores are larger. Verrucaria vindobonensis Zschacke has a dark exciple base, slightly larger ascospores (18–19 m) and an apically swollen exiple/involucrellum.

9HUUXFDULDFDOFDULD =VFKDFNH¿JV115, 131) Synonyms: Verrucaria aethiobola f. calcarea Arnold 1885, Verrucaria zschackei Riedl 1990. Uncertain species: Verrucaria kemmleri Servít 1950. 7KDOOXVgrey; semi-endolithic to epilithic, 25–100 m thick; smooth, continuous to cracked, especially around the perithecia; in some thalli partly with black basal layer (20–40 m thick); photobiont coccoid green alga, 3–4 m in diameter, uniformly dispersed in the thallus. Perithecia:LQVPDOOWKDOOXVZDUWVDSH[¿QDOO\“ naked; exciple 144–180 m in diameter; periphyses very short (10 m); involucrelOXPUHDFKLQJWKHWKDOOXVEDVHDGSUHVVHGWRWKHH[FLSOHEXWZLWKVSUHDGLQJÀDQNV DUHDEHWZHHQLQYROXFUHOOXPDQGH[FLSOHEDVH¿OOHGZLWKHQODUJHGG FHOOVZLWKEODFN brown cell walls; involucrellum covered by a thin thallus mantle; asci 8-spored, 72–75 × 22–25 m; ascospores unicellular, not halonate, 17–24 × 7–12 m. (FRORJ\amphibious on calcareous rocks at the banks of streams. 'LVWULEXWLRQ rare (Germany, North Rhine-Westphalia: Büren; Bavaria: Eichstätt and Munich), possibly overlooked and more widespread, generally (?) at rather low altitudes.

Fig. 131. Verrucaria calcaria: ×28.

Verrucaria

167

6LPLODUVSHFLHV: Verrucaria dolosa has more narrow and usually shorter ascospores. The perithecia are naked at least in the upper half. The thallus of V. dolosa often appears as an uneven or granular crust and is never semi-endolithic. Verrucaria austriaca does not have an involucrellum. Verrucaria submersella has larger perithecia with an exciple diameter of 200–320 m, an involucrellum that is mostly naked and rarely stretches to the thallus base. The thallus of Verrucaria elaeina is always epilithic and prominent parts of the perithecia are mostly naked. The involucrellum is often weakly coloured in its lower parts or leaving a transparent area close to the basal parts of the exciple. Uncertain species: Verrucaria kemmleri differs from V. calcaria only in the thicker involucrellum which reaches up to 60 m in width, while in V. calcaria a maximum of 30 m has been measured. Verrucaria kemmleri is only known from the type locality close to the village of Schneckenweiler (Germany, Baden-Württemberg) and it is uncertain whether the collection originated from an amphibious or dry, terrestrial habitat.

Verrucaria consociata6HUYtW¿JV 110, 132) 7KDOOXV greenish to olive; thin (40–60 m), forming goniocyst-like granules and patches which can become slightly rimose; cortex absent, but upper parts of thallus with yellowish-brown pigment; paraplectenchymatous but not subgelatinous; photobiont green alga, 5–10 m in diameter, in small groups or without special arrangement. Perithecia: prominent, upper half mostly exposed and naked or partly covered by a thin thallus mantle or granules; 150–180 m in diameter; exciple brownish throughout; periphyses up to 18 m long; involucrellum absent RU WKLQ FRPSOHWHO\ HQYHORSLQJ WKH H[FLSOH DQG GLI¿FXOW WR GLVWLQJXLVK XS WR 25 m thick; asci 8-spored, 65–80 × 22–27 m; ascospores unicellular, not halonate, 16–28 × 8–13 m.

Fig. 132. Verrucaria consociata: ×30.

168

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(FRORJ\on siliceous substrata at shaded places, usually rarely inundated (riparian) in or at ephemeral small streams and on temporary wet rocks. 'LVWULEXWLRQ Czech Republic 5XGRKRĜí, Suniperk, type locality), Germany (North Rhine-Westphalia, Ratingen-Hösel, leg. H. Thüs), Austria (Upper Austria, Mühl-viertel, Rannatal, leg, F. Berger), but possibly much overlooked. 6LPLODUVSHFLHV The terrestrial species of Verrucaria with hardly developed thallus are poorly known and the species delimitation between these taxa is in need of revision. Verrucaria acrotella Ach. differs in the black-brown thallus colour, shorter asci (45–55 m, up to 62 m according to Zschacke 1933) and ascospores (11–20 m). In some thalli the involucrellum is laterally sligthly protruding. Verrucaria sublobulata is an amphibious species with a thicker thallus and mostly immersed perithecia. The base of the exciple is transparent and the involucrellum is absent or poorly developed. Mechanical hybrids of V. hydrela with Bacidina inundata or Ionaspis lacustris have thicker thalli and usually at least some disciform fruit bodies (apothecia) are present or some perithecia have an involucrellum which is distinct and laterally spreading. Verrucaria viridula has larger perithecia (300–700 m) and ascospores (22–40 m long).

9HUUXFDULDGRORVD+HSS¿JV116, 133) 7KDOOXV greenish-grey, yellowish-green to brown; thin, variable from continuous to rimose or granulose, always epilithic; proso- to paraplectenchymatous, never subgelatinous; photobiont a coccoid green alga, generally dispersed or aggregated in small groups, never arranged in vertical columns. Perithecia: prominent to semi-immersed, naked or basal parts covered by algae containing thalline mantle; involucrellum reaching the thallus base, attached to the exciple or little VSUHDGLQJ DUHD EHWZHHQ EDVDO SDUW RI H[FLSOH DQG LQYROXFUHOOXP ¿OOHG E\ GDUN

Fig. 133. Verrucaria dolosa: ×24.

Verrucaria

169

coloured cells; asci 8-spored; ascospores unicellular, not halonate, 13–19 (22) × 5–8 (10) m. (FRORJ\ a terrestrial species but also amphibious in the splash water zone of small streams, also in cold springs on calcareous rocks and here tolerating longer periods of submersion (Nascimbene et al. 2007). Usually at shaded places. 'LVWULEXWLRQ Terrestrial populations are frequent and widespread, whereas amSKLELRXVSRSXODWLRQVDUHPXFKUDUHUDQGSRVVLEO\RYHUORRNHGRU PLVLGHQWL¿HGDV V. hydrela or V. elaeomelaena. 6LPLODUVSHFLHV Verrucaria calcaria has perithecia in small thalline warts (thin thallus cover at least on the lower half of the involucrellum), larger ascospores (17–24 × 7–12 m), a semi-endolithic to epilithic and smooth, rimose to cracked thallus. Verrucaria hydrela is restricted to siliceous substrata and has a laterally distinctly spreading, thin involucrellum which leaves a large transparent, triangular space between involucrellum and exciple base. The thallus is usually distinctly subgelatinous (transparent when wet). The terrestrial species V. muralis has larger ascospores with a smaller length/width-ratio and the involucrellum is usually spreading, often leaving a transparent area between exciple base and involucrellum. Verrucaria kalenskyi Servít has similar sized ascospores (16–18 m long) but a spreading involucrellum as in V. hydrela. It is unknown whether the collections made by Servít originate from amphibious habitats.

Verrucaria funckii 6SUHQJHO =DKOEUXFNQHU¿JV119– 9 120, 134, 166) Basionym: Pyrenula funckii Sprengel 1826. Synonyms: Verrucaria aethioboloides Zschacke 1933, Verrucaria bachmanniana Zschacke 1927, Verrucaria bachmannii Zschacke 1933, Verrucaria elaeomelaena f. silicola Zschacke 1927, Verru-

Fig. 134. Verrucaria funckii: ×18.

170

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caria silicicola (Zschacke) Servít 1950, Verrucaria silicea Servít 1954, Verrucaria f. silicola var. coniocarpa Zschacke 1927, Verrucaria silicea f. coniocarpa (Zschacke) Servít 1954, Verrucaria denudata var. mougeotii Zschacke 1927, Verrucaria mougeotii (Zschacke) Servít 1954. Uncertain species: Verrucaria nuda Zschacke 1933. 7KDOOXV sun-exposed morphs brown, orange-brown, also chocolate brown to black-brown at high altitudes or in fully sun-exposed situations, thalli from shady localities greenish-grey to grey; (20) 50–160 m thick, thin morphs in deep VKDGHFRQWLQXRXVWR¿QHO\ULPRVHVSHFLPHQVGDPDJHGE\UDSLGGU\LQJDOVRZLWK deep and wide cracks); paraplectenchymatous and subgelatinous; with or without black basal layer; cortex, if present, colourless or (yellowish) brown; photobiont: Heterococcus caespitsosus or 'LODEL¿OXP sp.YHUL¿Hd by own cultivated isolates), size of algal cells in the thallus 2.5–10 m, with one or more chloroplasts, occasionally cells longer than wide, usually arranged in vertical columns. Perithecia: variable corresponding to the thickness of the thallus, ± completely immersed in the thallus or in projecting thallus warts; exciple 90–350 × 100–250 m; involucrellum surrounding the exciple down to the base, laterally more or less protruding into the thallus and eventually fusing with a black basal layer; enlarged cells with distinctly brown cell walls in lower parts of the involucrellum giving a “meshlike” appearance in thin sections; asci 8-spored; ascospores unicellular, halonate or not; (15) 18–25 (27.6) × 5.4–8.7 (14) m. (FRORJ\ submerged or in the lowest part of the splash water zone on stable siliceous rocks. Characteristic element of permanently submerged communities in cool and clean (nutrient poor) springs and headwaters. At high altitudes in springs and rivers in full sunlight, at lower altitudes restricted to springs and headwaters in wooded areas. Best developed in well lit places, but tolerating moderate shading. Absent from eutrophicated watercourses with considerable growth of strong FRPSHWLWRUVIUHHOLYLQJDOJDHF\DQREDFWHULD 6HQVLWLYHWRVLOWLQJDQGDFLGL¿FDtion (absent from streams with pH permanently < 5.0). 'LVWULEXWLRQ from sea-level to (sub-) alpine altitudes but much declined at low alWLWXGHVGXHWRHXWURSKLFDWLRQDQGDFLGL¿FDWLRQDOVRHIIHFWHGE\JOREDOZDUPLQJ"  6LPLODUVSHFLHV Verrucaria pachyderma is an upland species which produces a greenish-black cortical pigment when growing in sunny sites. The colour of the dry thallus is olive to black-olive. Shade forms of V. pachyderma are more greenLVKDQGWKHFRUWLFDOSLJPHQWLVODFNLQJDQGWKHUHIRUHDUHGLI¿FXOWWRGLVWLQJXLVK I from V. funckii, but the ascospores of the former are usually shorter and have a larger length/width-ratio. Verrucaria hydrela has an involucrellum with enlarged but colourless cells below a sharply delimited outer mantle of strongly pigmented cells. Typically, a hyaline transparent triangle can be seen between the black involucrellum and the basal part of the exciple which is generally absent in V. funckii. The thallus of V. hydrela is thinner and although the ecological amplitude of the two species is overlapping, V. hydrela prefers less frequently inundated microsites. The ascospores are not halonate in V. hydrela. Verrucaria praetermissa is a splash water species with non-subgelatinous thallus, a usually cracked or almost areolate appearance, photobionts which are not arranged in vertical columns and a paler thallus colour. The thallus of morphs from shady

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171

localities can be virtually continuous, but the non-subgelatinous consistency and algae without vertical arrangement are constant characters. Verrucaria margacea s.l. has larger, distinctly projecting perithecia, larger non-halonate ascospores and a generally thin thallus. Uncertain species: Verrucaria nuda is certainly not a distinct taxon, but rather UHSUHVHQWVDPRUSKIRUPHGXQGHUFRQGLWLRQVRIH[WHQGHGSHULRGVRILQVXI¿FLHQW light supply. Similar forms can be induced in V. hydrela by wrapping well develRSHGWKDOOLZLWKEODFNSODVWLF¿OPDVZDVVKRZQLQFRUUHVSRQGLQJH[SHULPHQWVE\ 7KV 7KHWKDOOXVZDVQRWG\LQJV\QFKURQLFDOO\DVDZKROHEXW¿UVWGLVDSpeared at the margins, whereas the thallus parts surrounding the perithecia remained unchanged for several months. After three months of blocking sunlight a growth habit very similar to that of V. nuda had developed. The altered thallus was characterized by a thin margin of green thallus left around the perithecia and a complete loss of > 90 % of the algae containing thallus parts. The anatomy of the perithecia, however, did not change at all during the experiment. These experiments were conducted with thalli of V. hydrela and the characteristic transparent area between involucrellum and exciple base was always distinct in the perithecia. The type specimen of V. nudaLQVWHDGKDVDQLQYROXFUHOOXPWKDW¿WV better with V. funckii because all cells around the exciple base have distinctly black-brown cell walls. The illustration of a young perithecium by E. Bachmann in Zschacke (1933) shows homogenously carbonised areas surrounding the exciple on one side, but show a less coloured area between exciple and involucrelOXPRQWKHRWKHU7KHUHIRUHVRPHGRXEWVUHJDUGLQJWKHLGHQWL¿FDWLRQRIWKHW\SH specimen as either V. funckii or V. hydrela still remain.

9HUUXFDULDK\GUHOD $FKDULXV¿JV117–118, 157–158) Synonyms: Verrucaria denudata Zschacke 1927, Verrucaria eidorensis Erichsen 1940, Verrucaria denudata f. nuda Servít 1950, Verrucaria sublobulata f. robustiorr Servít 1951. Uncertain species: Verrucaria davosiensis Zschacke 1927, Verrucaria submauroides Zschacke 1933, Verrucaria subhydrela Servít 1951. 7KDOOXV grey, greenish to brownish; thin, 10–50 (65) m, corticate or not, cortical pigment (yellowish-) brownish; paraplectenchymatous and usually subgelatinous (but depending on growth conditions and sometimes not at all apparent); black basal layer absent or developed here and there and thin; photobiont cells usually dispersed, rarely in vertical columns, size of photobiont cells very variable in the thallus, 3.0–4.0 m (Elliptochloris bilobata, Diplosphaera chodatii, 'LODEL¿OXP VSLGHQWL¿HGE\FXOWLYDWLRQRILVRODWHV  Perithecia: prominent, ± covered by a thallus mantle (eventually much reduced by erosion or herbivore activities = V. denudata); total diameter 160–1315 m; exciple 100–255 m in diameter, upper part covered by a very thin to moderately thick (6.5–40 m) involucrellum which is usually distinct, sharply delimited and reaching the thallus base, laterally spreading and leaving a colourless area between basal part of exciple and involucrellum (hyaline triangle), diffuse light brown (never mesh-like as in V. funckii) in overaged specimens; asci 8-spored, 70–75 × 23–28 m (according to Zschacke 1933); ascospores unicellular, not halonate, (11) 20–25 (30) × 4.5–15.5 m.

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(FRORJ\ amphibious to submerged, rarely also at humid terrestrial sites, on siliceous rocks and pebbles, rapid coloniser with wide tolerance to different illumination, best developed at semi-shaded sites and one of the most shade tolerant species of Verrucaria. With wide tolerance to variable substrate pH (4.5–8.5) but restricted to siliceous substrata and absent from calcareous rocks. 'LVWULEXWLRQ one of the most frequent amphibious lichen from low altitudes to montane regions, much rarer in the subalpine belt and here ± replaced by V. margacea s.str. 6LPLODUVSHFLHV Verrucaria sublobulata has a thick, non subgelatinous thallus with semi-immersed perithecia, slightly smaller ascospores and a weakly develRSHGRIWHQ“DSLFDOLQYROXFUHOOXP7KLVVSHFLHVLVYHU\GLI¿FXOWWRGLVWLQJXLVK from mechanical hybrids of V. hydrela with Bacidina inundata or Ionaspis lacustris where the involucrellum of V. hydrela becomes similarly reduced. According to Thüs ( WKHVHPRUSKVFDQEHLGHQWL¿HGE\WKHSUHVHQFHRIGLVFLIRUPIUXLW bodies (apothecia) of Bacidina or Ionaspis in some parts of the thallus (often only very few among the much more frequent perithecia of V. hydrela), a clustered arrangement of the photobiont cells, as well as a blastidiate thallus structure and the eventual occurrence of some better developed perithecia with an anatomy typical of V. hydrela in some parts of the thallus with more prominent perithecia. The thallus of V. dolosa is generally not subgelatinous, and the ascospores are smaller. The involucrellum is variable, but usually there is no or only a small transparent area between the basal part of the exciple and the involucrellum. Verrucaria submersella is a species from alkaline, hard waters and never found in acidic waterbodies, the thallus is not subgelatinous, rimose and the perithecia are naked with an usually adpressed or only slightly spreading involucrellum. In V. praetermissa the thallus is not subgelatinous, rimose to areolate and the perithecia are ± immersed. The species of the V. margacea-group have larger ascospores and perithecia. In V. funckii the hyaline triangle between exciple base and involucrellum is absent and the thalli are usually thicker (shade forms have similar thalli, but the perithecial characters are constant). Verrucaria pachyderma is an upland species with thick thallus, a thick black basal layer, narrow ascospores and a greenish-black cortex pigment in specimens from sunny sites. The involucrellum is variable, often even within a single specimen, but usually adpressed to the exciple. Uncertain species: Verrucaria subhydrela has a smaller exciple diameter but all other characters coincide with V. hydrela. No authentic material of V. davosiensis could be traced, but based on the protologue, it seems to be similar to V. hydrela.

9HUUXFDULDODWHEURVD.|UEHU¿JV 127– 7 128, 135) Synonym: Verrucaria anziana Garovaglio 1865. Uncertain species: Verrucaria areolatodiffracta Zschacke 1927. 7KDOOXV grey (sometimes with a pink tinge) to brown, colour little changing when wet; thick (60–85 m), rimose to cracked-areolate; proso- to paraplectenchymatous, never subgelatinous; cortex colourless to yellowish-brown; basal

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Fig. 135. Verrucaria latebrosa: ×17. layer absent or colourless to yellowish; prothallus absent; photobiont various coccoid green algae (e.g. Stichococcus sp., Diplosphaera chodatii, Elliptochloris bilobata LGHQWL¿HG E\ FXOWLYDWLRQ RI LVRODWHV  JHQHUDOO\ GLVSHUVHG VL]H RI WKH photobiont cells in the thallus 4.8–8 m in diameter. Perithecia: immersed to prominent, naked; exciple 195–390 m in diameter; involucrellum usually apical but reaching down to the thallus base in damaged or regenerated specimens, cells in the lower part of involucrellum not enlarged, pigmentation “patchy” in thin sections, in thick sections often with a completely carbonised appearance; asci 8-spored; ascospores unicellular, in fresh specimens halonate, halo distinctly swelling in K, 18.1–37.0 × 8.2–14.8 m (average usually within the range of 27–30 m). (FRORJ\ amphibious, typically in the bed of non-permanent small streams and temporarily irrigated rocks, but also in the splash water zone of permanent streams and rivers on siliceous rocks and pebbles. Best developed in well lit places but also tolerating moderate shading. 'LVWULEXWLRQ (high-)montane to alpine sites; very rare at low altitudes (e.g. Harz Mountains, Germany, at 230 m). 6LPLODU VSHFLHV Verrucaria aethiobola has smaller ascospores and perithecia. Verrucaria margacea has an involucrellum which reaches the thallus base. It is distinctly laterally protruding into the thallus, and enlarged cells with brown walls in the basal part of the involucrellum give a mesh-like appearance in thin sections. Damaged specimens of V. latebrosa and V. margacea are very hard or even impossible to distinguish. Verrucaria submersella prefers alkaline (often calcareous) rocks and has a thin (on limestone occasionally semi-immersed) thallus of whitish to brownish colour, prominent perithecia and non-halonate ascospores.

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Uncertain species: Verrucaria areolatodiffracta Zschacke is very similar to V. latebrosa although the thallus is described in the protologue as being paraplectenchymatous, whereas V. latebrosa has a prosoplectenchymatous hyphal arrangement which is at least partly visible in the thallus. Type material of V. areolatodiffracta was not traced so far, but there are observations of intergrading forms between para- and prosoplectenchymatous thalli in V. latebrosa and more GHWDLOHGVWXGLHVDUHQHFHVVDU\LQRUGHUWRHYDOXDWHWKHWD[RQRPLFVLJQL¿FDQFHRI this character in V. latebrosa.

Verrucaria madida 2UDQJH¿JV 136, 155) 7KDOOXV green, olive-greyish to blackish-green; thin (40–60 m), continuous to ¿QHO\ ULPRVH SDUDSOHFWHQFK\PDWRXV DQG VXEJHODWLQRXV FRUWH[ ROLYHEODFNLVK pigmented (turning brownish in HCl) in specimens from well lit localities; prothallus absent; photobiont coccoid green alga. Perithecia: prominent, 200–420 m in diameter; covered by a thallus mantle when young, upper parts may become naked with age; involucrellum reaching down to the thallus base, laterally spreading into the thallus, with pale area between involucrellum and lower part of the exciple; asci (3) 4 (5)-spored; ascospores unicellular, not halonate, 9–15 × 5.5–7.5 m, length/width-ratio 1.4–2.6. (FRORJ\ amphibious at frequently inundated sites on siliceous rocks, often together with V. aquatilis, Hydropunctaria rheitrophila and aquatic bryophytes. 'LVWULEXWLRQ very rare in oceanic regions in northwestern Europe (Great Britain, Norway), a single collection from Central France (Cantal) and one from Germany (Baden-Württemberg: Freiburg, leg. V. Wirth).

Fig. 136. Verrucaria madida: ×17.

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6LPLODUVSHFLHV Distinguished from all other hydrophilic Verrucaria species by the 4-spored asci. Verrucaria aquatilis is most similar in outer appearance but differs in the distinctly smaller perithecia, a usually black or blackish-green thallus, 8-spored asci and smaller ascospores. Hydropunctaria rheitrophila and H. scabra have ascospores of similar size, but 8-spored asci, a thicker thallus and often black dots or columns which are either scattered in the thallus or located on the surface of the involucrellum. Other epilithic Verrucaria species have much larger ascospores.

Verrucaria margacea s.l. Notes: In the study area the V. margacea-group comprises the following three species which are treated here in detail: (1) V. margacea Wahlenberg (V. margacea s.str.), (2) V. elaeomelaena Arnold and (3) V. andesiatica Servít. Recent molecular studies by Thüs et al. (in prep.) showed that most species of the Verrucaria margacea-group form a monophyletic clade of distinct lineages with large perithecia and ascospores, a laterally spreading involucrellum and a paraplectenchymatous thallus which is usually subgelatinous when wet. The species ERXQGDULHVZLWKLQWKLVJURXSDUHGLI¿FXOWWRGUDZEDVHGRQPRUSKRORJLFDOFKDUacters, and the determination of a given specimen to one of the morphotaxa that have been described may prove utterly impossible due to the overlapping character states and the very similar ecological preferences. The molecular data, however, suggest that V. applanata Hepp 1868 may represent an independent lineage, but there seems to be no diagnostic combination of morphological characters which could serve to satisfactorily separate V. applanata from certain other genotypes within the V. margacea-complex (Thüs et al., in prep.). Therefore, the morphospecies V. applanata is provisionally included here in V. margacea, but further combined studies of molecular and morphological characters are required to solve the question of how to separate a genetically distinct lineage from its lookalikes in the V. margacea-complex.

(1) Verrucaria margacea:DKOHQEHUJ¿J 123, 137) Basionym: Thelotrema margaceum Wahlenberg in Acharius 1803. Synonyms: Pyrenula margacea (Wahlenberg) Acharius 1809, Lithocia margacea (Wahlenberg) A. Massalongo 1853, Involucrothele margacea (Wahlenberg) Servít 1954, Lithoicea elaeomelaena var. alpina Arnold 1873, Verrucaria alpicola Zschacke 1927, Verrucaria elaeomelaena f. silicicola var. platycarpa Zschacke 1927 nom. inval., Verrucaria leightonii A. Massalongo 1855. Uncertain species: Verrucaria applanata Hepp 1868, Verrucaria delita f. subcontinua Nylander 1881, Thelidium subcontinuum (Nyl.) Servít 1949, Involucrothele subcontinua (Nylander) Servít 1954, 9HUUXFDULD ¿ODUV]N\DQD Zschacke 1931, Verrucaria maas-geesterani Servít 1954, Verrucaria pachyspora Servít 1952, 9HUUXFDULDYDOOLVÀXHODH Zschacke 1927, Verrucaria zenogensis Zschacke 1933. 7KDOOXV light brown to brown-black, greyish, or even greenish at shady sites; thin to thick (15–70 m), continuous to rimose, rarely almost areolate around perithecia; proso- to paraplectenchymatous, some morphs subgelatinous, others

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Fig. 137. Verrucaria margacea s.str.: ×18. not; prothallus whitish; black basal layer absent; photobiont cells rounded or HORQJDWHG ±—P LQ GLDPHWHU LGHQWL¿HG DV (OOLSWRFKORULV ELOREDWD 'LODEL¿lum sp. in cultivated isolates), cells dispersed, usually not in vertical columns Perithecia: prominent, 225–1360 m in diameter, covered by a thin thallus mantle; exciple 150–265 × 160–335 m; covered by a distinct but thin (12–40 m) involucrellum which is reaching the thallus base and is laterally spreading; often with a triangular area of colourless cells between exciple base and or all cells in this area with black-brown cell walls, sometimes both types can be found in perithecia from the same thallus; the specimens with the colourless area between involucrellum and exciple base correspond to the type specimens of V. applanata, those with brown-black cells in this area are similar to the type specimen of V. margacea; involucrellum in lower parts with enlarged cells, deep brown cell walls give a mesh-like appearance of this area in thin sections; very rarely the involucrellum is completely colourless lacking any brownish pigments; asci 8spored, 90–100 × 36 m; ascospores unicellular, not halonate, (20.6) 25–41.2 × 8–18.3 m. (FRORJ\ amphibious in the splash water zone of cool streams and rivers on siliceous or (rarely) on hard calcareous rocks. Absent from highly acidic watercourses (pH < 5). Very tolerant towards variable illumination intensities and developing different morphs which vary in thallus colour and thickness depending on the given light intensity. 'LVWULEXWLRQ widely distributed in montane to alpine regions, very rare at low altitudes and restricted to cool springs or shaded headwaters. 6LPLODUVSHFLHV Verrucaria latebrosa has a usually thicker, more rimose to areolate thallus, usually has an apical involucrellum and the ascospores are halonate in fresh specimens. Verrucaria andesiatica differs in shorter asci (60–70 m). Verrucaria elaeomelaena is restricted to calcareous rocks and differs from V.

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margacea in the egg-shaped ascospores which are usually halonate when freshly collected. Verrucaria hydrela and V. funckii have smaller ascospores (mostly < 25 m). Uncertain species: Verrucaria applanata is separated from V. margacea s.str. by the spreading involucrellum leaving a transparent angle between involucrellum and the basal part of the exciple ¿J 124). It has a greyish to greyish-brown thallus, while V. margacea s.str. has brown-black thalli at sun-exposed sites. Verrucaria zenogensis differs from V. applanata only by the transparent exciple base, which in V. applanata is reported to be brownish in some perithecia (Zschacke 1933). 9HUUXFDULD¿ODUV]N\DQD has a similar involucrellum structure like V. applanata but differs in the whitish thallus and a thin black basal layer. It is only known from the type locality on volcanic rocks in the East-Carpathians near the village of Bad Gyertyánliget (Hungary). Verrucaria pachyspora is unique in its large and almost spherical ascospores. The type specimen, however, appears to be in poor condition and large parts are overgrown by Bacidina inundata. It is probably just an aberrant growth form of V. margacea. Verrucaria maasgeesterani differs from typical specimens of V. margacea in the presence of a black basal layer and black columns which raise from the basal layer to the thallus surface. In general, the species circumscriptions in the V. margacea-group (V. andesiatica, V. applanata, V. elaeomelaena, 9¿ODUV]N\DQD, V. maas-geesterani, V. margacea, V. pachyspora, V. tiroliensis, 9YDOOLVÀXHODH) are still unsettled and require further studies including molecular markers.

(2) 9HUUXFDULDHODHRPHODHQD $UQROG¿J126, 138) Synonyms: Verrucaria elaeomelaena f. calcicola Zschacke 1927, Verrucaria teutoburgensis Zschacke 1927. Uncertain species: Verrucaria jurana Zschacke 1927, Verrucaria tiroliensis Zschacke 1927. 7KDOOXV dry thallus greenish, grey, brownish to black-brown; usually rather thick (50–280 m), epilithic to chasmoendolithic, always with dominating epilithic part; continuous to rimose; cortical pigment (yellowish-) brown or absent; with paraplectenchymatous hyphal arrangement and subgelatinous consistency when wet; black basal layer occasionally present here and there in the thallus; photobiont green alga ('LODEL¿OXP ( sp. in cultivated isolates). Perithecia: immersed to prominent, usually with a thallus mantle, 250–700 m in diameter; exciple 200–400 m in diameter; involucrellum very variable, often varying from apical to reaching down to the thallus base and ± laterally spreading within the same thallus, upper surface of the involucrellum often rough and irregularly shaped, lower part with enlarged cells, black-brown cell walls may give a meshlike appearance; asci 8-spored, 70–90 × 25–33 m; ascospores unicellular, halonate or not, 20–30 × (10) 12–16 m. (FRORJ\ submerged or amphibious on calcareous rocks and pebbles in cool springs and streams. Poor competitor and vanishing in eutrophicated streams with fast growing algae or cyanobacteria. Wide tolerance to varying illumination.

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Fig. 138. Verrucaria margacea s.l. (V. elaeomelaena): ×19. 'LVWULEXWLRQ widespread but rare; lowland populations endangered by eutrophication and destruction of natural springs. 6LPLODU VSHFLHV Verrucaria submersella is an amphibious lichen with naked, hemispherical perithecia, and has a rimose to areolate thallus that is not subgelatinous. Verrucaria margacea has a thin thallus (15–70 m), ascospores with a higher length/width-ratio and usually lacks a black basal layer. Intermediate forms exist and V. elaeomelaena is possibly better treated as a subspecies of V. margacea s.l. Uncertain species: Verrucaria tiroliensis is a semi-endolithic lichen with a black-brown thallus and massive black basal layer. The ascospores are variable in shape with a length/width ratio from 2.0 to 2.4. Verrucaria jurana has similarly shaped ascospores but has a thin, black basal layer which is present only here and there. The thallus is brown and semi-endolithic with the fungal hyphae protruding into a depth of up to 40 m.

(3) Verrucaria andesiatica 6HUYtW¿J125) 7KDOOXV olive-brownish; thin (up to 20 m), continuous to rimose, rarely almost areolate around the perithecia; black basal layer absent; photobiont coccoid green algae, 6–9 m in diameter. Perithecia: prominent, up to 500 m in diameter, covered by a thin thallus mantle; exciple 170 m in diameter, covered by a spreading involucrellum reaching the thallus base, distinct but thin (< 35 m); asci 8-spored, 60–80 × 22–28 m; ascospores unicellular, not halonate, 26–30 × 10–14 m. (FRORJ\ amphibious in the splash water zone of streams and rivers on siliceous rocks.

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'LVWULEXWLRQRQO\NQRZQIURPWZRORFDOLWLHV8NUDLQH8åKRURG9HONH/i]\  and from Poland (High Tatra, Dolina Chochlowska) but elsewhere possibly overlooked. 6LPLODUVSHFLHV Verrucaria margacea s.str. differs in the longer asci (> 90 m) and the thicker, ± brownish thallus. Verrucaria elaeomelaena differs from V. andesiatica in the longer asci (as in V. margacea), the egg-shaped ascospores which are often halonate when freshly collected and the restriction to calcareous rocks. Verrucaria hydrela and V. funckii have smaller ascospores (mostly length < 25 m).

Verrucaria pachyderma $UQROG¿JV121–122, 139–140) Synonyms: Verrucaria pissina Nylander 1881. 7KDOOXV olive-greenish-black; thick (50–250 m); cortex well developed in thalli from well lit places, upper cortex with blackish-green or blackish-brown (not yellowish-brown) pigment, cortical pigment absent in thalli from shaded sites; black basal layer very variable, absent to building up more than 80 % of the thallus; paraplectenchymatous and subgelatinous; transparent when wet only in cases when black basal layer is lacking; photobiont coccoid green alga, arranged in ± vertical columns, rounded or elongated, 4–6 m in diameter or 10 × 4–5 m (in culture cells rounded and Stichococcus-like). Perithecia: immersed (slightly projecting in thin thalli), covered by a thallus mantle; exciple up to 200 m in diameter; involucrellum very variable, from reaching the thallus base to almost apical in the same thallus, often fused with the black basal layer and completely mantling the exciple, but also separated from the exciple base leaving a colourless area between basal part of exciple and involucrellum; asci 8-spored, 45–50 × 16–18 m; ascospores unicellular, halonate or not, 18–20 (23) × 7–8 (9) m.

Fig. 139. Verrucaria pachyderma: old herbarium specimen with wide cracks due to long storage; ×17.

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Fig. 140. Verrucaria pachyderma: freshly collected specimen with few thallus cracks; ×17. (FRORJ\ permanently submerged or amphibious in permanent, cool streams, on stable, siliceous rocks. Predominantly in well lit sites. 'LVWULEXWLRQ widespread but rare in upland areas, usually above the timer line in the Alps and Carpathians (High Tatra). So far only one locality known at rather low altitude (wooded canon of river Teigitsch in Styria, Austria; Keller 2000). More frequent in Nordic countries and NW Britain and there growing at much lower altitudes than in Central Europe. 6LPLODUVSHFLHV Verrucaria funckii differs in a yellowish-brown to black-brown, never greenish-brown cortex pigment. Dry thalli are (orange-) brownish to brown-black or grey to greenish-grey, but without olive tinge. Thalli from shaded VLWHVDUHGLI¿FXOWWRVHSDUDWHIURPV. pachyderma since the cortex pigment may be absent. The ascospores of V. funckii are usually larger, and the length/widthratio is lower. The involucrellum is always reaching down to the thallus base and is usually completely mantling the exciple. Verrucaria hydrela has usually distinctly projecting perithecia with a colourless area between the base of the exciple and the involucrellum, and the ascospores have a lower length/width-ratio. Verrucaria margacea has larger ascospores and a thinner thallus.

Verrucaria praetermissa 7UHYLVDQ $Q]L¿J 141) Basionym: Leiophloea praetermissa Trevisan 1860. Synonyms: Verrucaria laevata auct. p.p., Verrucaria annulifera Eitner 1910, Verrucaria tapetica var. ÀXvialis Eitner 1911, Verrucaria viridicana Erichsen 1940, Verrucaria javorinae Servít 1949, Verrucaria granitica Servít 1950, Verrucaria guestphalica Servít 1954.

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Fig. 141 Verrucaria praetermissa: ×6. 7KDOOXV whitish to greyish, greenish, pinkish, orange to brownish, intensive green when wet; usually thick, (40) 60–260 m, thinner in shaded localities, thallus rimose to areolate, rarely continuous (occasionally at shady locations); paraplectenchymatous but never subgelatinous; cortex distinct or absent (often damaged by herbivores), yellowish-brown to pinkish or orange, pinkish cortical pigment usually K+ reddish; prothallus white to cream-coloured; black basal layer usually present, often building up more than half of the thallus, but morphs with thin or virtually absent basal layer exist; photobiont a coccoid green alga (Diplosphaera chodatii LGHQWL¿HGLQLVRODWHGFXOWLYDWHV GLVSHUVHGLQWKHWKDOOXVGLDmeter of photobiont cells in the thallus 4.8–12.9 m, occasionally different algae (e.g. 'LODEL¿OXP 7UHERX[LD) also incorporated in the thallus though in minor quantities (endophytes?). Perithecia: immersed or slightly prominent, often only the ostiole visible, but herbivory or shading may result in much thinner thalli with more prominent, naked perithecia; exciple 90–240 m in diameter; involucrellum reaching down to the thallus base, very rarely reduced and ± apical, basal parts of involucrellum with much enlarged cells and brown cell walls and thus creating a mesh-like appearance in thin sections, involucrellum usually much protruding laterally in the thallus, 260–900 m in diameter, but often fusing with adjacent involucrella or with the black basal layer; basal parts of involucrellum and exciple fused, leaving no free space in between; asci 8-spored; ascospores unicellular, occasionally thinly halonate, but halo generally little swelling in K, (12.7) 17–25 (27.2) × 5.9–8 (13.4) m. (FRORJ\ amphibious in the splash water zone, rarely at sites that are submerged for longer periods or in the upper amphibious zone of large rivers where inundation and splash water wetting occur only for a few weeks during the year. Characteristic species for several amphibious cryptogamic plant associations on slightly acidic to basic rocks. Tolerating a wide range of illumination. Best developed on hard and stable, slightly acidic to basic siliceous or calcareous substrata. Sensitive to atmospheric as well as to ZDWHUDFLGL¿FDWLRQDQGUHVWULFWHGWRZDWHU-

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courses with a pH > 5. Rather tolerant to silting and eutrophication and able to compete even with mat-forming cyanobacteria in turbulent waters. 'LVWULEXWLRQ from sea-level to high-montane regions, very rarely reaching alpine sites. Much declined in air-polluted areas, but during the last decade rapidly recovering across large areas with reduced acidic emission and good to moderate water quality. One of the most frequent amphibious lichens in Central Europe. 6LPLODUVSHFLHV Verrucaria aethiobola has a less developed, often apical involucrellum without enlarged cells in the basal part. Pigmentation appearing patchy in thin sections. The involucrellum has no tendency to fuse with adjacent perithecia and a black basal layer is generally absent. Verrucaria hydrela usually develops a continuous thin crust of subgelatinous consistency with ± transparent margins when wet. However, non-subgelatinous morphs and mechanical hybrids for example with Bacidina inundata or Ionaspis lacustris exist. These forms can be PXFKPRUHFUDFNHGWKDQWKHFRPPRQIRUPDQGDUHEHVWLGHQWL¿HGE\WKHLQYROXcrellum structure. In V. hydrela basal parts of the involucrellum and exciple are generally separated by a transparent angle, while in V. praetermissa there is no such free space between the involucrellum and the exciple. Verrucaria latebrosa is an alpine to high-montane species with larger, halonate ascospores, a perispore that is much swelling in K and usually has an ± apical involucrellum (rarely reaching down to the thallus base) with a patchy pigmentation. Verrucaria elaeina is a terrestrial species from shady, humid localities with a thin thallus, protruding perithecia and generally absent black basal layer. It has very varely been reported from amphibions sites (GB, Orange 2006) and ephemeral brooks (Slovakia, Guttová & Palice 2002). The involucrellum is less extended when compared to most morphs of V. praetermissa. However, morphs of V. elaeina from shady ORFDOLWLHVDUHVRPHWLPHVGLI¿FXOWRULPSRVVLEOHWRVHSDUDWHIURPV. praetermissa.

9HUUXFDULDVXEOREXODWD (LWQHUH[6HUYtW¿JV 112, 142–143, 157–158) Synonym: Verrucaria lignicola Zschacke 1927. 7KDOOXV (light) olive, greenish to greenish grey, light brownish after long storage in herbarium; thick, uneven, continuous to rimose or almost areolate, margins occasionally almost lobate; proso- to paraplectenchymatous but never subgelatinous; photobiont coccoid green alga, scattered in the thallus. Perithecia: immersed to semi-immersed and then ± projecting; exciple up to 150 m in diameter; periphyses up to 10 m; involucrellum thin, apical to reaching down to the thallus base and little spreading laterally in the thallus, usually fading in lower part; asci 8spored, 40–55 × 14–20 m; ascospores unicellular, 16–20 × 7–9 m, not halonate. (FRORJ\ amphibious on siliceous rocks, often at ± shaded sites, rarely also on roots of elder (Alnus ( glutinosa). 'LVWULEXWLRQ rare (?) but widespread in mountainous regions. Due to frequent PLVLGHQWL¿FDWLRQV DQG FKDQJLQJ FRQFHSWV RI VSHFLHV GHOLPLWDWLRQ HJ 7KV 2002), literature records for this species should be used cautiously and any inter-

Verrucaria

183

Fig. 142. Verrucaria sublobulata: ×22.

Fig. 143. Verrucaria sublobulata: specimen with lobule-like thallus margin; ×14. pretation should rely on prior revision of the cited herbarium specimens. Consequently, the distribution range of V. sublobulata remains largely unknown. 6LPLODU VSHFLHV Verrucaria consociata has a thinner, often ± granular thallus, and the involucrellum is either absent or thin, completely enveloping the exciple and hard to distinguish. The base of the exciple is always dark. Verrucaria hydrela is a subgelatinous species with prominent perithecia, a distinctly dark brown, laterally spreading involucrellum and slightly larger ascospores (mostly 18–23 m). Mechanical hybrids of V. hydrela with Bacidina inundata or Ionaspis lacustris FDQEHGLI¿FXOWWRGLVWLQJXLVKDVWKHWKDOOXVLQWKHVHIRUPVLV QRWVXEJHODWLQRXV

184

.H\VWRWKHVSHFLHVDQGVSHFLHVSUR¿OHV

and the involucrellum is much reduced. Fusions with Bacidina are often characterized by a distinctly aggregated, instead of a homogenous distribution of the photobiont cells inside the thallus. If an aggregated photobiont distribution is observed or apothecia of either Bacidina or Ionaspis are found on V. sublobulatalike thalli, then parts of the thallus with the most protruding perithecia should be examined and checked for perithecia with typical V. hydrela-anatomy and intergrading (reduced) forms.

9HUUXFDULDVXEPHUVHOOD 6HUYtW¿JV 111, 144) Synonyms: Verrucaria submersa Schaerer 1850 non Borrer 1834., Verrucaria submersa b. litorea Hepp 1865, Verrucaria litorea (Hepp) Zschacke 1927, Verrucaria submersa var. frankonia Zschacke 1933, Verrucaria frankonia (Zschacke) Servít 1950, Verrucaria turicensis Zschacke 1927 non (Winter) Stizenberger 1882, Verrucaria zahlbruckneri Zschacke 1933. Uncertain species: Verrucaria erubecens Zschacke 1927, Verrucaria rivalis Zschacke 1927, Verrucaria tatrensis Servít 1950. 7KDOOXV whitish to grey, rarely greenish-grey; thin to thick (50–150 m), epilithic to semi-immersed, continuous to rimose; proso- to paraplectenchymatous; black basal layer absent; photobiont coccoid green alga with dispersed to aggregated arrangement in the thallus, rounded, 4–8 m in diameter, rarely elongated 10–13 × 4.5–7 m. Perithecia: semi-immersed, hemispherical, raised above the thallus, 250–450 m in diameter, naked; exciple 200–320 × 180–250 m; involucrellum apical to reaching to the lower half of the exiple rarely to the thallus base adpressed to the exciple or slightly laterally protruding into the thallus; asci 8-spored, 65–85 × 14–30 m; ascospores unicellular, not halonate, 20–25 (30) × 9–14 m.

Fig. 144. Verrucaria submersella: ×21.

Verrucaria

185

(FRORJ\ amphibious in the splash water zone on calcareous and (alkaline or silt covered) siliceous rocks and pebbles of streams and rivers with usually hard, well buffered water. Tolerant to limited periods of complete submersion as well as to longer periods of desiccation. Predominantly in sun-exposed sites. 'LVWULEXWLRQ montane to subalpine areas below the timber line in the Alps. 6LPLODUVSHFLHV Verrucaria aethiobola has a thicker thallus and it is restricted to siliceous rocks. Verrucaria latebrosa has a thicker thallus, larger ascospores and perithecia typically with an apical involucrellum and a halonate ascospores in freshly collected material. It prefers soft waters and appears to be restricted to siliceous rocks. Verrucaria praetermissa has ± immersed perithecia and often a black basal layer. Verrucaria hydrela is restricted to siliceous rocks and it has smaller perithecia which are coated by a thallus layer. The thallus is continuous and usually subgelatinous. The involucrellum is thin and laterally distinctly spreading. Uncertain species: Verrucaria tatrensis differs only in the stronger development of the endolithic thallus parts (penetrating up to 100 m below the rock surface). Verrucaria rivalis is another semi-endolithic taxon which is in need of revision. The epilithic parts of the thallus form very thin crusts lacking cracks. The perithecia (400–500 m in diameter) as well as the ascospores (24–30 × 12–15 m) are larger than in V. submersella s.str. Verrucaria rivalis is solely known from the type locality in Austria (North Tyrol, Waldrast). Verrucaria erubescens is an epilithic taxon from calcareous rocks which is only known from the type locality (Switzerland, Davos, Zügenstraße). It has a reddish grey-green thallus which turns blackish-green in K. The involucrellum is indistinguishable from the exciple, but the upper part of the exciple is blackish and 80 m thick, while the lower part is transparent and thin. The ascospores are 25–28 × 10–13 m. Uncertain species in Verrucaria: Verrucaria applanata Hepp 1868 – see V. margacea s.str. Verrucaria areolatodiffracta Zschacke 1926 – see V. latebrosa Verrucaria davosensis Zschacke 1927 – see V. hydrela Verrucaria erubescens Zschacke 1926 – see V. submersella 9HUUXFDULD¿ODUV]N\DQD Zschacke 1931 – see V. margacea s.str. Verrucaria fuscorubens (de Lesdain) Zschacke 1927 – see V. aethiobola Verrucaria jurana Zschacke 1927 – see V. margacea s.l. / V. elaeomelaena Verrucaria kalenskyi Servít 1951 – see V. dolosa Verrucaria kemmleri Servít 1950 – see V. calcaria Verrucaria maas-geesterani Servít 1954 – see V. margacea s.str. Verrucaria maurula Müller Argoviensis 1857 – see Hydropunctaria rheithrophila Verrucaria nuda Zschacke 1933 – see V. funckii Verrucaria pachyspora Servít 1952 – see V. margacea s.str. Verrucaria pseudocatalepta Servít 1952 – see V. aethiobola Verrucaria pulvinata Eitner 1911 – a lichenicolous fungus on Rhizocarpon disporum with small (10–11 × 9–11 Pm), globose ascospores only known from the type collection and associated with Candelariella vitellina, and thus unlikely to originate from an amphibious habitat. Verrucaria rechingerii Servít 1950 – see V. aquatilis, Hydropunctaria rheitrophila

186

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Verrucaria refugii Servít 1950 – see Hydropunctaria rheithrophila Verrucaria rivalis Zschacke 1927 – see V. margacea s.l. / V. elaeomelaena Verrucaria subhydrela Servít 1951 – see V. hydrela Verrucaria submauroides Zschacke 1933 – see V. hydrela Verrucaria tatrensis Servít 1950 – see V. submersella Verrucaria tiroliensis Zschacke 1927 – see V. margacea s.l. / V. elaeomelaena 9HUUXFDULDYDOOLVÀXHODH Zschacke 1927 – see V. margacea s.str. Verrucaria wolferi Zschacke 1927 – see Hydropunctaria rheitrophila Verrucaria zenogensis Zschacke 1933 – see V. margacea s.str. (V. applanata)

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Index of Lichen Names Normal print – valid names. Italics – synonyms and names of taxa, which need UHYLVLRQRUFRQ¿UPDWLRQXQFHUWDLQVSHFLHV6WDQGDUG print – EULHÀ\ mentioned in keys or the chapter “similar species”. Bold print – taxa with detailed descriptions. Bold italics ±UHIHUHQFHWRLOOXVWUDWLRQV

acceptanda (Aspicilia ( ) (=Koerberiella wimmeriana) 71 acceptanda (Lecanora ( ) (=Koerberiella wimmeriana) 71 acceptanda var. littoralis (Lecanora ( ) (=Koerberiella wimmeriana) 71 DFURWHOOD9HUUXFDULD   acrotellum (Thelidium  7PLQXWXOXP   adglutinata (Anziella (  3ODF\QWKLXPÀDEHOORVXP   adglutinatum (Lecothecium (  3ODF\WKLXP ÀDEHOORVXP   adglutinatum (Placynthium  3ÀDEHOORVXP  adglutinatum (Pterygium  3ODF\QWKLXP ÀDEHOORVXP   DGVFHQGHQV(QGRFDUSRQ   adscendens (Physcia) 31 aenea (Porina) 97 aeneovinosa (Involucrothele  7KHOLGLXP PHWKRULXP   aeneovinosa (Sagedia  7KHOLGLXPPHWKRULXP   aeneovinosa (Verrucaria  7KHOLGLXPPHWKRULXP   aeneovinosum (Thelidium  7PHWKRULXP  aeneovinosum var. kutakii (Thelidium) (=T. PHWKRULXP   DHWKLREROD9HUUXFDULD  160, 163, 163  aethiobola ff. calcarea (Verrucaria)  9FDOFDULD   aethiobola var. griseocinerascens (Verrucaria)  9DHWKLREROD   aethiobola var. primaria (Verrucaria)  9DHWKLREROD   aethioboloides (Nylander) Vainio non Zschacke (Thelidium  7PLQXWXOXP   DHWKLRERORLGHV7KHOLGLXP  138, 138,  217 aethioboloides (Verrucaria  9IXQFNLL  9 aggregatula (Homopsella  3RURF\SKXV coccodes) 101 ahlesiana (Porina) 95, 95, 100 ahlesiana (Segestrella  3RULQDDKOHVLDQD   ahlesii (Biatora  /HFLGHDDKOHVLL  

DKOHVLL/HFLGHD   72, 73 ahlesii (Mycobilimbia  /HFLGHDDKOHVLL   DOEHVFHQV/HFDQRUD   DOERFDHUXOHVFHQV3RUSLGLD   DOJHULFD*RQRK\PHQLD   alpicola (Verrucaria  9PDUJDFHDVVWU   alpinum (Collema  &JOHEXOHQWXP   ambrosiana (Staurothele  "6IUXVWXOHQWD   amphibia ((Abacina  5KL]RFDUSRQDPSKLELXP) 119 amphibia (Buellia  5KL]RFDUSRQDPSKLELXP 119 amphibia ((Lecidea  5KL]RFDUSRQDPSKLELXP 119 amphibia (Pyrenopsis) (=Pterygiopsis concorGDWXOD   DPSKLELXP5KL]RFDUSRQ  119, 119,  DQGHVLDWLFD9HUUXFDULD  162 177, 178 DQJHUPDQQLFXP&ROOHPRSVLGLXP   $QLVRPHULGLXP  annulifera (Verrucaria  9SUDHWHUPLVVD   anziana (Verrucaria) (=V. latebrosa) 162 anziana f. peminosa (Polyblastia) (=P. peminosa) 93 applanata (Verrucaria) (= ? V. margacea) 162,  DTXDOH/HSWRJLXP   75, 76 DTXDWLFD$VSLFLOLD  37, 37 aquatica (Lecanora (  $VSLFLOLDDTXDWLFD   aquatica (Pachyospora  $VSLFLOLDDTXDWLFD 37 aquaticum (Dermatocarpon (  'OXULGXP   DTXDWLFXP7KHOLGLXP    138, 139 aquatilis ((Lithoicea  9HUUXFDULDDTXDWLOLV   DTXDWLOLV9HUUXFDULD   164, 164 223 DUHQDULD&DORSODFD   areolata (Pertusaria  6WDXURWKHOHDUHRODWD   areolata (Pyrenula (  6WDXURWKHOHDUHRODWD   DUHRODWD6WDXURWKHOH   127 215 areolatodiffracta (Verrucaria) (= ? V. latebrosa) 

  ,QGH[RIOLFKHQQDPHV areolatus (Porocyphus) (=P. coccodes) 101 DUQROGLDQD%DFLGLQD   DUQROGLDQXP'HUPDWRFDUSRQ   50, 50,  DUQROGLL7KHOLGLXP   DVSHUHOOXP3ODF\QWKLXP   $VSLFLOLD  assimulans (Collemopsis) (= ? Pterygiopsis FRQFRUGDWXOD   assimulans (Psorotichia) (= ? Pterygiopsis FRQFRUGDWXOD   assimulans (Pyrenopsis) (= ? Pterygiopsis FRQFRUGDWXOD   DWURÀDYDYDUVXEPHUVD&DORSODFD 42, 219 atrovirens (Conferva  (SKHEHODQDWD   DXUHOOD&DQGHODULHOOD   DXVWULDFD9HUUXFDULD  165 bachmanniana (Verrucaria  9IXQFNLL   bachmannii (Verrucaria  9IXQFNLL   bachmannii var. inundatum (Dermatocarpon)  'PHLRSK\OOL]XP   %DFLGLD  %DFLGLQD  EDFLOOLJHUD6WDXURWKHOH   badioaterr (Catocarpus) (=Rhizocarpon badioaWUXP   badioatra (Buellia) (=Rhizocarpon badioaWUXP   badioatra (Lecidea ( ) (=Rhizocarpon badioaWUXP   EDGLRDWUD5LPXODULD   EDGLRDWUXP5KL]RFDUSRQ   120, 120, 121 ED\HULDQD3VHXGDUWKRS\UHQLD   biatorina (Rinodina  5R[\GDWD   byssoides (Porocyphus  3UHKPLFXV    caesia (Arthopyrenia ( ) (=Pyrenocollema FDHVLXP   caesia (Leiophloea (  3\UHQRFROOHPDFDHVLXP) 113 caesia (Physcia) 31 caesia (Pseudarthopyrenia) (=Pyrenocollema FDHVLXP   caesia (Verrucaria  3\UHQRFROOHPDFDHVLXP 113 caesiocinerea $VSLFLOLD   caesium (Leptogium  /F\DQHVFHQV   FDHVLXP3\UHQRFROOHPD  113 FDOFDULD9HUUXFDULD  160, 166 &DORSODFD  cancellata (Synalissa) (=Lempholemma polyanthes) 73 cantabria (Ephebeia  (SKHEHODQDWD   (  $QLVRPHULGLXP carinthiaca (Arthopyrenia FDULQWKLDFXP   carinthiaca (Ditremis  $QLVRPHULGLXP FDULQWKLDFXP   carinthiaca (Paraphysothele (  $QLVRPHULGLXP FDULQWKLDFXP  

FDULQWKLDFXP$QLVRPHULGLXP 35, 35 FDUQHRJODXFD%DFLGLD   FDUSDWKLFD/HFLGHOOD   cataclystum var.ÀXFWXDQV (Leptogium) (=L. plicatile) 77 catalepta 0DOPHQRQ$FKDULXV %ORPEHUJ  Forsell (Staurothele) (=6FORSLPD   cataractarum (Involucrothele  7KHOLGLXP IRQWLJHQXP   cataractarum (Porocyphus  3RURF\SKXV coccodes) 101 cataractarum (Sagedia  7KHOLGLXPIRQWLJHQXP   cataractarum (Thelidium) (=7IRQWLJHQXP   cataractarum (Verrucaria) (=7KHOLGLXP IRQWLJHQXP   chalazanellum ((Lempholemma) (=L. polyanthes) 73 chalazanodes (Lempholemma ( ) (=L. polyanthes) 73 FKDO\EHLD&DWLOODULD   FKORULQD&DORSODFD   chlorotica (Arthopyrenia ( ) (=Porina chlorotica)  FKORURWLFD3RULQD  96, 96 chlorotica (Pseudosagedia ( ) (=Porina chlorotica)  chlorotica (Pyrenula  3RULQDFKORURWLFD   chlorotica (Sagedia  3RULQDFKORURWLFD   chlorotica (Segestrella  3RULQDFKORURWLFD   chlorotica (Segestria  3RULQDFKORURWLFD   chlorotica (Verrucaria  3RULQDFKORURWLFD   FKORURWLFXOD%DFLGLQD   chloroticum (Trichothelium) (=Porina chloURWLFD   chrysophana (Aspicilia (  ,VXDYHROHQV   chrysophana (Ionaspis  ,VXDYHROHQV   chrysophana (Lecanora  ,VXDYHROHQV   cinereolutescens (Verrucaria  "+\GURSXQFWDULDUKHLWKURSKLOD   circumspersella (Verrucaria  7KHOLGLXP FLUFXPVSHUVHOOXP   FLUFXPVSHUVHOOXP7KHOLGLXP  139, 140,  circumspersellum f. lojkanum (Thelidium) (=T. FLUFXPVSHUVHOOXP 140 FOHLVWRFDUSD3\UHQRSVLV   FORSLPD6WDXURWKHOH  127, 128,  clopimaa (Verrucaria  6WDXURWKHOHFORSLPD   clopima var. areolata (Staurothele  6 DUHRODWD   clopimoides (Polyblastia  6WDXURWKHOH FORSLPRLGHV   clopimoides (Sphaeromphale  6WDXURWKHOH FORSLPRLGHV   FORSLPRLGHV6WDXURWKHOH  129, 129215 clopimoides (Stigmatomma  6WDXURWKHOH FORSLPRLGHV   clopimoides (Verrucaria  6WDXURWKHOHFORSLPRLGHV  

,QGH[RIOLFKHQQDPHV   FRDUFWDWD7UDSHOLD   coccodes (Collema  3RURF\SKXVFRFFRGHV 101 coccodes 3RURF\SKXV   101, 102,  coccodes (Psorotichia (  3RURF\SKXVFRFFRGHV) 101 cochleatum /HSWRJLXP   &ROOHPD  compactum (Lempholemma ( ) (=L. polyanthes) 73 FRQFRUGDWXOD3WHU\JLRSVLV  109, 121, 123 concordatula (Pyrenopsis) (=Pterygiopsis FRQFRUGDWXOD   conferciens (Pterygium  "3ODF\QWKLXP SDQQDULHOOXP   7 FRQVRFLDWD9HUUXFDULD  160, 167, 167 convexa var. hydrophila (Lecidea ( ) (=Porpidia K\GURSKLOD   coracodiza (Collemopsis) (=Pterygiopsis FRQFRUGDWXOD   coracodiza (Psorotichia) (=Pterygiopsis FRQFRUGDWXOD   coracodiza (Pterygiopsis  3FRQFRUGDWXOD 109 coralliferum (Collema) (=&JOHEXOHQWXP   corallinoides ssp. rosulans (Lecothecium)  3ODF\QWKLXPURVXODQV   corallinoides var. rosulans (Lecothecium)  3ODF\QWKLXPURVXODQV   corallinoides var. tantalea (Biatora)  3ODF\QWKLXPWDQWDOHXP   FRUDOORSKRUD$VSLFLOLD   creatina (Lecanora ( ) (=Koerberiella wimmeriana) 71 creatina (Lecidea ( ) (=Koerberiella wimmeriana) 71 crenatellum (Leptogium (  /ULYXODUH   cruenta (Polyblastia  6SRURGLFW\RQFUXHQWXP   cruenta (Segestrella  6SRURGLFW\RQFUXHQWXP   cruenta (Sphaeromphale  6SRURGLFW\RQ FUXHQWXP   FUXHQWXP6SRURGLFW\RQ  124, 125 FUXVWXODWD3RUSLGLD   crustulata ssp. soredizodes (Lecidea) (=PorSLGLDVRUHGL]RGHV   cuprea var. fuscocuprea (Verrucaria  6WDXURWKHOHFORSLPD   cyanescens (Collema  /HSWRJLXPF\DQHVFHQV   F\DQHVFHQV/HSWRJLXP  76, 77 cyanocarpa (Aspicilia ( ) (=Hymenelia cyanoFDUSD   cyanocarpa +\PHQHOLD   68213 cyanocarpa (Ionaspis) (=Hymenelia cyanoFDUSD   cyanocarpa (Lecanora) (=Hymenelia cyanoFDUSD   cyanothalama (Lecidea) ( (=Porpidia hydrophila) cylindrica (Umbilicaria) 31 czernaensis (Verrucaria) 9DHWKLREROD  

davosiensis (Verrucaria) (= ? V. hydrela) 171,  GHFLSLHQV7KHOLGLXP   delita (Verrucaria  9DHWKLREROD   delita f. subcontinua (Verrucaria) (= V. PDUJDFHD9DSSODQDWD   GHOXWXOD$EVFRQGLWHOOD   GHPDQJHRQLL3K\OOLVFXP   GHPLQXHQV'HUPDWRFDUSRQ   denudata (Verrucaria) (=V. hydrela) 161, 171 denudata f. nuda (Verrucaria) (=V. hydrela) 171 denudata var. mougeotii (Verrucaria) (=V. IXQFNLL   'HUPDWRFDUSRQ  GHXVWD8PELOLFDULD   diaboli (Thelidium  "7PHWKRULXP   diaboli (Verrucaria) "7KHOLGLXPPHWKRULXP   diaphana (Biatora  *\DOLGHDGLDSKDQD   diaphana (Biatorina (  *\DOLGHDGLDSKDQD   diaphana (Catillaria  *\DOLGHDGLDSKDQD  1 diaphana (Gyalidea) 61, 61 diaphana (Lecidea  *\DOLGHDGLDSKDQD   diblastum (Placynthium  3WDQWDOHXP   dichotoma (Tremella  &ROOHPDGLFKRWRPXP   GLFKRWRPXP&ROOHPD   44, 44, 45 discolor (Lecidea  5LQRGLQDR[\GDWD   discolor (Rinodina  5R[\GDWD   GLVSHUVD/HFDQRUD   dispersus (Porocyphus) (=Thelignya lignyota)   GLVSRUXP5KL]RFDUSRQ   GLVWDQV$QLVRPHULGLXP   GLVWLQFWXP5KL]RFDUSRQ   dolichotera (Pannaria  3ODF\QWKLXPGROLFKRWHUXP GLVSRUXP5KL]RFDUSRQ   dolichoterum (Lecothecium  3ODF\QWKLXP GROLFKRWHUXP GLVSRUXP5KL]RFDUSRQ   GROLFKRWHUXP3ODF\QWKLXP   85, 85 dolichoterum f. melanterum (Placynthium)  3GROLFKRWHUXP   dolichoterum var. pluriseptatum (Placynthium  3GROLFKRWHUXP   GRORVD9HUUXFDULD  160 168, 168,  HJHQXOD%DFLGLQD   eidorensis (Verrucaria) (=V. hydrela) 171 eitneri (Involucrothele (  7KHOLGLXPPLQXWXOXP   eitneri (Thelidium  7PLQXWXOXP   HODHLQD9HUUXFDULD   HODHRPHODHQD9HUUXFDULD  162  177, 178 elaeomelaena f. calcicola (Verrucaria) (=V. elaeomelaena) 177 elaeomelaena f. silicicola var. coniocarpa (Verrucaria  9IXQFNLL   elaeomelaena f. silicicola var. platycarpa (Verrucaria  9PDUJDFHDVVWU  

  ,QGH[RIOLFKHQQDPHV elaeomelaena f. silicola (Verrucaria)  9IXQFNLL   elaeomelaena var. alpina (Verrucaria)  9PDUJDFHDVVWU   (QGRFDUSRQ  endococcina (Parmelia) (=Phaeophyscia HQGRFRFFLQD   HQGRFRFFLQD3KDHRSK\VFLD  83, 83 (SKHEH  epomphala (Polyblastia)  epomphalum (Thelidium) (=Polyblastia epomSKDOD   HSXORWLFD+\PHQHOLD   erubescens (Verrucaria  "9VXEPHUVHOOD   erysiboda (Verrucaria) (=Porina lectissima) 99 excile (Thelidium  7PLQXWXOXP   expallens (Lecanora)  ¿ODUV]N\DQD(Verrucaria) (= ? V. margacea)  ¿PEULDWD5LQRGLQD 122, 123 ¿VFKHUi (Staurothele  "6VXFFHGHQV    ¿VVD (Polyblastia  6WDXURWKHOH¿VVD   ¿VVD (Sphaeromphale  6WDXURWKHOH¿VVD   ¿VVD6WDXURWKHOH   130, 130 ¿VVD (Verrucaria  6WDXURWKHOH¿VVD   ¿VVXP (Stigmatomma  6WDXURWKHOH¿VVD   ¿VVXP (Thelotrema  6WDXURWKHOH¿VVD   ¿VVXPvar. clopimoides (Stigmatomma)  6WDXURWKHOHFORSLPRLGHV   ÀDEHOORVD (Pannaria  3ODF\QWKLXPÀDEHOORVXP   ÀDEHOORVXP3ODF\QWKLXP  86, 86 ÀDFFLGXP&ROOHPD  45, 46 ÀDFFLGXV (Lichen  &ROOHPDÀDFFLGXP   ÀDYLGD(LJOHUD   ÀDYRFDHUXOHVFHQV3RUSLGLD   ÀRWRZLDQD (Biatora) (=Pyrenocarpon ÀRWRZLDQXP   ÀRWRZLDQD (Montinia) (=Pyrenocarpon ÀRWRZLDQXP   ÀRWRZLDQD (Psorotichia) (=Pyrenocarpon ÀRWRZLDQXP   ÀRWRZLDQD (Pyrenopsis) (=Pyrenocarpon ÀRWRZLDQXP   ÀRWRZLDQD (Thelochroa) (=Pyrenocarpon ÀRWRZLDQXP   ÀRWRZLDQD (Verrucaria) (=Pyrenocarpon ÀRWRZLDQXP   ÀRWRZLDQXP3\UHQRFDUSRQ  111, 112 ÀRWRZLDQXV (Collemopsis) (=Pyrenocarpon ÀRWRZLDQXP   ÀRWRZLDQXV (Porocyphus) (=Pyrenocarpon ÀRWRZLDQXP   ÀRWRZLDQXV (Pyrenocarpus) (=Pyrenocarpon ÀRWRZLDQXP   ÀXYLDWLOH (Collema  &GLFKRWRPXP   ÀXYLDWLOH'HUPDWRFDUSRQ  'OXULGXP  

ÀXYLDWLOLV (Lichen (  &ROOHPDGLFKRWRPXP   fontigena (Involucrothele  7KHOLGLXP IRQWLJHQXP   fontigenum (Spermatodium  7KHOLGLXP IRQWLJHQXP   7 140, 141 IRQWLJHQXP7KHOLGLXP  137, frankonia (Verrucaria  9VXEPHUVHOOD   fritzei (Gyalecta  *\DOLGHDIULW]HL   IULW]HL*\DOLGHD   62, 213 IUXVWXOHQWD6WDXURWKHOH    131, 131, 215 IUXVWXORVD3VRURWLFKLD 111 fuligenea (Staurothele  6¿VVD   fuliginascens (Collemopsis) (= ? Thelignya OLJQ\RWD   fuliginascens (Porocyphus) (= ? Thelignya OLJQ\RWD   fuliginascens (Psorotichia) (= ? Thelignya OLJQ\RWD   fuliginea (Verrucaria) (=Thelignya lignyota)   fuliginea var. subareolata (Pyrenopsis) (=P. VXEDUHRODWD   IXPLGXP7KHOLGLXP) P   funckii (Pyrenula  9HUUXFDULDIXQFNLL   IXQFNLL9HUUXFDULD  161, 169, 169,  223 furfurellus (Porocyphus  3RURF\SKXV coccodes) 101 furvellum (Collema  &JOHEXOHQWXP   fuscocuprea (Staurothele  6FORSLPD   fuscocuprea (Verrucaria  6FORSLPD   fuscorubens (Verrucaria) (?=V. aethiobola)   IXVFRYLUHQV&ROOHPD  47, 47 fuscovirens ((Lichen  &ROOHPDIXVFRYLUHQV  7 JHRJUDSKLFXP5KL]RFDUSRQ   geophila (Psorotichia (  3RURF\SKXVUHKPLFXV   JLEERVD5LPXODULD   gibbosa var. lacustris (Lecanora (  ,RQDVSLV ODFXVWULV   gibbosa var. supertegens (Lecanora) (= AspiFLODVXSHUWHJHQV   JODXFRSKDHD3RUSLGLD   JOHEXOHQWXP&ROOHPD  48, 48 glebulentum (Collemodium) (=Collema glebulentum   ( ) (=Collema glebulenglebulentum (Leptogium tum   globulosa (Psorotichia  3RURF\SKXV UHKPLFXV   globulosus (Porocyphus  3UHKPLFXV   JORPHUXODQV$VSLFLOLD   *RQRK\PHQLD  JUDQGLV3RULQD  97, 97, 99 grandis (Pseudosagedia) (=Porina grandis) 97 grandis (Sagedia) (=Porina grandis) 97 granitica (Verrucaria  9SUDHWHUPLVVD  griseocinerascens (Verrucaria) (=V. aethiobola) 

,QGH[RIOLFKHQQDPHV   JXHQWKHUL3RULQD  98, 98 guentheri (Pseudosagedia (  3RULQDJXHQWKHUL  guentheri (Trichothelium  3RULQDJXHQWKHUL  guentheri (Verrucaria  3RULQDJXHQWKHUL   guentheri var. grandis (Porina ( ) (=P. grandis) 97 guestphalica (Verrucaria) (=V. praetermissa)  *\DOLGHD  haematopsisVHQVXDXFWEULWPyrenopsis)  3VXEDUHRODWD   hazslinskyi (Staurothele)  6¿VVD   henscheliana (Polyblastia  6SRURGLFW\RQ FUXHQWXP   henscheliana (Sphaeromphale  6SRURGLFW\RQ FUXHQWXP   henscheliana (Verrucaria  6SRURGLFW\RQ FUXHQWXP   henschelianum (Sporodictyon  6FUXHQWXP  KHSSLL*RQRK\PHQLD   59, 60 heppii (Omphalaria) (=Gonohymenia heppii)  heppii (Thyrea  *RQRK\PHQLDKHSSLL   KHWHURPRUSKD+\PHQHOLD   hibernica (Verrucaria  9DHWKLREROD   KLVSLGXOD(SKHEH   hochstetteri f. papulare (Amphoridium ( )  7KHOLGLXPSDSXODUH   KRORFDUSD&DORSODFD   hospitum (Thelidium  7PLQXWXOXP   hyalinescens (Bacidia) (=Gyalidea hyalineVFHQV   hyalinescens (Bilimbia) (=Gyalidea hyalineVFHQV   hyalinescens (Gyalecta) (=Gyalidea hyalineVFHQV   hyalinescens (Gyalidea) 62, 213 hyalinescens (Lecidea ( ) (=Gyalidea hyalinescens)  K\GUHOD9HUUXFDULD   161 171  219 hydrocharum (Collema)  /HSWRJLXPSOLFDWLOH 77 hydrocharum (Leptogium) (=L. plicatile) 77 hydrophila ((Huilia  3RUSLGLDK\GURSKLOD   hydrophila a (Lecidea  3RUSLGLDK\GURSKLOD  K\GURSKLOD3RUSLGLD   105, 105 hydrophilum (Haplocarpon) (=Porpidia K\GURSKLOD   +\GURSXQFWDULD  +\PHQHOLD  LQFDYDWXP7KHOLGLXP   LQFRQVSLFXOD$UWKRS\UHQLD   inornata (Aspicilia (  "$VXSHUWHJHQV   intercedens var. aethioboloides (Verrucaria)  7KHOLGLXPPLQXWXOXP   LQWHUMXQJHQV3RULQD   99

interjungens (Pseudosagedia) (=Porina interMXQJHQV   interjungens (Verrucaria) (=Porina interMXQJHQV   interseptula (Porina  3LQWHUMXQJHQV   interseptula (Verrucaria) (=Porina interMXQJHQV   intricata (Ephebe  (ODQDWD   LQWULFDWD/HFDQRUD   intricata (Usnea  (SKHEHODQDWD   intricatus (Lichen  (SKHEHODQDWD   inundata (Bacidia  %DFLGLQDLQXQGDWD   LQXQGDWD%DFLGLQD   41, 41  inundata (Biatora  %DFLGLQDLQXQGDWD   LQXQGDWD/HFDQLD   inundata (Woessia  %DFLGLQDLQXQGDWD   LQXQGDWXP7KHOLGLXP   141, 141 LQYROXWD7UDSHOLD   LRGRSXOFKUD*RQRK\PHQLD   ,RQDVSLV  irrigua (Verrucaria) (=Porina lectissima) 99 irrigua Zschacke non Taylor (Verrucaria) (=V. DXVWULDFD   isarina (Staurothele  "6VXFFHGHQV    LVLGLLJHUD&DORSODFD   43, 111, 219 LVLGLRLGHV3HUWXVDULD   japonicum (Anisomeridium (  $FDULQWKLDFXP  javorinae (Verrucaria  9SUDHWHUPLVVD   MHPWODQGLFXP5KL]RFDUSRQ   MHQHQVLV*\DOHFWD   josiae (Lecanorella) (=Koerberiella wimmeriana) 71 jurana (Verrucaria) (=V. elaeomelaena) 177,  NDOHQVNêL9HUUXFDULD   kalenskýi (Paraphysothele  $QLVRPHULGLXP FDULQWKLDFXP   kemmleri (Verrucaria  "9FDOFDULD    NHQPRUHQVLV3RURF\SKXV   kernstockii (Verrucaria  +\GURSXQFWDULD UKHLWKURSKLOD   kernstockii ff. minutipuncta (Verrucaria)  +\GURSXQFWDULDUKHLWKURSKLOD   NOHPHQWLL7KHOLGLXP   142, 142 koerberi (Amphoridium (  3RULQDJXHQWKHUL   koerberi (Sagedia  3RULQDJXHQWKHUL   koerberi (Segestria  3RULQDJXHQWKHUL   koerberi (Spermatodium  3RULQDJXHQWKHUL  k koerberi (Verrucaria  3RULQDJXHQWKHUL   Koerberiella 71 kutakii (Involucrothele  7KHOLGLXPPHWKRULXP   kutakii (Polyblastia  7KHOLGLXPPHWKRULXP  kutakii (Thelidium  7KHOLGLXPPHWKRULXP 

  ,QGH[RIOLFKHQQDPHV lacustris (Aspicilia (  ,RQDVSLVODFXVWULV   lacustris (Hymenelia  ,RQDVSLVODFXVWULV   ODFXVWULV,RQDVSLV  69 213 lacustris (Lecanora  ,RQDVSLVODFXVWULV   lacustris (Lichen  ,RQDVSLVODFXVWULV   ODFXVWULV3WHU\JLRSVLV   laevata $VSLFLOLD   38, 38 laevata (Lecanora (  $VSLFLOLDODHYDWD   laevata (Sagedia  $VSLFLOLDODHYDWD   laevata (Urceolaria  $VSLFLOLDODHYDWD   laevata (Verrucaria) 9DHWKLREROD   laevata DXFWQRQ$FKDULXVVerrucaria) SS 9K\GUHOD9SUDHWHUPLVVD   ODHYLJDWXP$QLVRPHULGLXP   lahmiana (Involucrothele (  7KHOLGLXPODKPLDQXP   ODKPLDQXP7KHOLGLXP  143, 143 ODQDWD(SKHEH   57, 57 lanatus (Lichen (  (SKHEHODQDWD   lapponica (Ephebe  (ODQDWD   ODWHEURVD9HUUXFDULD   162 172, 176 latzelianum (Endocarpon  "(SVRURGHXP  lavata (Amphoroblastia ( ) (=Polyblastia lavata)  lavata (Buellia  5KL]RFDUSRQODYDWXP   lavata (Lecidea (  5KL]RFDUSRQODYDWXP   ODYDWD3RO\EODVWLD   ODYDWXP5KL]RFDUSRQ  121, 121 lecanopsoides (Collemposis  3RURF\SKXV coccodes) 101 lecanopsoides (Psorotichia  3RURF\SKXV coccodes) 101 lecanopsoides (Pyrenopsis  3RURF\SKXV coccodes) 101 /HFDQRUD  /HFLGHD  /HFLGHOOD  Lecideopsis *\DOLGHD   lecideopsis var. fritzei (Gyalecta) (=Gyalidea IULW]HL   lectissima ((Arthopyrenia) (=Porina lectissima) 99 OHFWLVVLPD3RULQD  99, 100 lectissima (Sagedia) (=Porina lectissima) 99 lectissima (Segestria) (=Porina lectissima) 99 lectissima (Sphaeromphale) (=Porina lectissima) 99 lectissima (Verrucaria) (=Porina lectissima) 99 leightonii (Verrucaria  9PDUJDFHDVVWU   Lemmopsis 111 Lempholemma 73 OHQWLFXODULV&DWLOODULD   OHSWRJLHOOD3RURF\SKXV   /HSWRJLXP  leptophyllodes 'HUPDWRFDUSRQ   51, 51,  leptophyllodes (Endocarpon) (=DermatocarSRQOHSWRSK\OORGHV  

leucophyma (Aspicilia ( ) (=Koerberiella wimmeriana) 71 leucophyma (Lecanora) (=Koerberiella wimmeriana) 71 leucophyma (Perspicinora) (=Koerberiella wimmeriana) 71 leucophyma var. littoralis (Aspicilia ( ) (=Koerberiella wimmeriana) 71 OLFKHQRLGHV/HSWRJLXP   /LFKLQHOOD  lignicola (Verrucaria  9VXEOREXODWD   lignyota (Thelignya) 33, 155, 156 lignyota (Verrucaria) (=Thelignya lignyota)  Lithina (Staurothele)  6¿VVD   litorea (Verrucaria  9VXEPHUVHOOD   littoralis (Aspicilia ( ) (=Koerberiella wimmeriana) 71 littoralis (Lecanora ( ) (=Koerberiella wimmeriana) 71 littoralis (Pertusaria) (=Koerberiella wimmeriana) 71 /RERWKDOOLD  lorenzianum (Dermatocarpon) (=D. leptophylORGHV   OXULGXP'HUPDWRFDUSRQ   52 219 luridus (Lichen  'HUPDWRFDUSRQOXULGXP  lusca (Aspicilia (  $ODHYDWD   lusca (Lecanora  $VSLFLOLDODHYDWD   maas-geesterani (Verrucaria) (?=V. margacea)  PDFURFDUSD3RUSLGLD   macrocarpa var. hydrophila (Lecidea ( ) (=PorSLGLDK\GURSKLOD   PDGLGD9HUUXFDULD  174, 174, 217 PDJQXVVRQLL/HSWRJLXP   margacea (Involucrothele  9HUUXFDULD PDUJDFHDVVWU   margacea (Lithocia (  9HUUXFDULDPDUJDFHDVVWU  margacea (Pyrenula  9HUUXFDULDPDUJDFHD VVWU   PDUJDFHDVO9HUUXFDULD   162 178 PDUJDFHDVVWU9HUUXFDULD   162, 175,  margacea var. aethiobola (Verrucaria)  9DHWKLREROD   margaceum (Thelotrema  9HUUXFDULDPDUJDFHDVVWU   martindalei (Ephebeia  (SKHEHODQDWD   massalongiana (Naetrocymbe) (= ? PyrenocolOHPDVD[LFROD   massalongiana (Sagedia) (= ? Pyrenocollema VD[LFROD   massalongiana (Segestrella) (= ? PyrenocolOHPDVD[LFROD   massalongiana (Segestria) (= ? Pyrenocollema VD[LFROD  

,QGH[RIOLFKHQQDPHV   massalongiana (Spermatodium) (= ? PyrenoFROOHPDVD[LFROD   PDVVLOLHQVH/HSWRJLXP   PDVWUXFDWD$VSLFLOLD   maurula (Verrucaria  "+\GURSXQFWDULD UKHLWKURSKLOD   meiophylliza (Placidiopsis) (=Dermatocarpon PHLRSK\OOL]XP   PHLRSK\OOL]XP'HUPDWRFDUSRQ  52, 53 melanaspis (Aspicilia ( ) (=Lobothallia PHODQDVSLV   melanaspis (Lecanora ( ) (=Lobothallia melanaspis)  ( ) (=Lobothallia melanaspis (Lichen melanaspis)  PHODQDVSLV/RERWKDOOLD  82, 82 melanaspis (Parmelia) (=Lobothallia PHODQDVSLV   melanaspis (Parmularia ( ) (=Lobothallia melanaspis   melanaspis (Placodium) (=Lobothallia PHODQDVSLV   melanaspis (Placolecanora) (=Lobothallia PHODQDVSLV   melanaspis (Squamarina) (=Lobothallia PHODQDVSLV   PHODQRFDUSD+\PHQHOLD   melantera (Pannaria  3ODF\QWKLXPGROLFKRWHUXP   melantera (Pannularia  3ODF\QWKLXPGROLFKRWHUXP   melantera (Parmeliella (  3ODF\QWKLXPGROLFKRWHUXP   mesotropa (Arthopyrenia (  7KHOLGLXPPLQXWXOXP   mesotropa (Verrucaria  7KHOLGLXPPLQXWXOXP   mesotropum (Thelidium  7PLQXWXOXP   methoria (Involucrothele  7KHOLGLXPPHWKRULXP   methoria (Verrucaria  7KHOLGLXPPHWKRULXP  PHWKRULXP7KHOLGLXP   137, 144, 144, 145 meylanii (Staurothele  "6VROYHQV   0LFDUHD  microcarpa (Verrucaria  7KHOLGLXP]ZDFNKLL  microcarpum (Thelidium  7]ZDFNKLL   microphylla var. radiata (Lecidea) (=PlacynWKLXPURVXODQV   PLQLDWXPDXFW'HUPDWRFDUSRQ   miniatum var. miniatum (Dermatocarpon) " 'DUQROGLDQXP   minutipuncta (Verrucaria  +\GURSXQFWDULD UKHLWKURSKLOD   minutipuncta f. laevior (Verrucaria) (=HydroSXQFWDULDUKHLWKURSKLOD   PLQXWXOXP7KHOLGLXP   137, 145, 146,  montinii (=Pyrenocarpon) 111 mougeotii (Verrucaria  9IXQFNLL  

muelleri (Lempholemma) (=L. polyanthes) 73,  PXOWLVSRUD(SKHEH  58, 58 multispora (Spilonematopsis) (=Ephebe PXOWLVSRUD   PXUDOLV/HFDQRUD   PXUDOLV9HUUXFDULD   myriococcum (Lempholemma ( ) (=L. polyanthes) 73 neglecta (Forssellia) (=Pterygiopsis neglecta) 110 QHJOHFWD3WHU\JLRSVLV   110, 110 nigra f. tantalea (Pannularia  3ODF\QWKLXP WDQWDOHXP   QLJUHVFHQV9HUUXFDULD   QLJULFDQV3KDHRSK\VFLD   QLJULFDQV7KHOLGLXP   146, 147 QLJUXP3ODF\QWKLXP   nigrum var. tantaleum (Placynthium) (= P. WDQWDOHXP   nuda (Verrucaria  "9IXQFNLL   obscurata (Buellia (  5KL]RFDUSRQODYDWXP   obscuratum (Diplotomma) (= Rhizocarpon ODYDWXP   obscuratum (Phalodictyum) (= Rhizocarpon ODYDWXP   obscuratum (Rhizocarpon  5ODYDWXP   obscuratum f. amphibium (Rhizocarpon)  5DPSKLELXP   obscuratum f. lavatum (Rhizocarpon)  5ODYDWXP   ocellata (Pyrenopsis) (=Thelignya lignyota)  ocellatus (Porocyphus) (=Thelignya lignyota)  odora ((Aspicila  ,RQDVSLVRGRUD   odora (Gyalecta  ,RQDVSLVRGRUD   RGRUD,RQDVSLV  70, 213, 223 odora ((Lecanora  ,RQDVSLVRGRUD   odora (Patellaria  ,RQDVSLVRGRUD   odora (Pinascisca  ,RQDVSLVRGRUD   oenipontana (Staurothele) 130, 131 RUELFXODULV3KDHRSK\VFLD   RUELFXODULV6WDXURWKHOH   oxydata (Berengeria (  5LQRGLQDR[\GDWD   R[\GDWD5LQRGLQD   123, 124 SDFK\GHUPD9HUUXFDULD   161, 170, 179, 179, 180 pachyspora (Verrucaria) (= ? V. margacea)  SDFL¿FD$QLVRPHULGLXP   pallida * psorodea (Verrucaria) (=Endocarpon SVRURGHXP   pannariellum (Lecothecium  3ODF\QWKLXP SDQQDULHOOXP   SDQQDULHOOXP3ODF\QWKLXP  88, 89, 91 pannariellum (Pterygium  3ODF\QWKLXP SDQQDULHOOXP  

  ,QGH[RIOLFKHQQDPHV pannariellum f. conferciens (Placynthium)  "3SDQQDULHOOXP   pannariellum f. sparsum (Placynthium)  3ÀDEHOORVXP   pannariellum var. rosulans (Placynthium)  3URVXODQV   pannariellum var. squamulosum (Placynthium)  3URVXODQV   papulare (Amphoridium (  7KHOLGLXPSDSXODUH  SDSXODUH7KHOLGLXP  147, 148 papularis (Verrucaria  7KHOLGLXPSDSXODUH  parvulum (Thelidium  7PLQXWXOXP   SHPLQRVD3RO\EODVWLD  93, 93 peminosa (Verrucaria) (=Polyblastia peminosa) 93 SHUVSLQXORVD(SKHEH   SHUWXVDWLL7KHOLGLXP   137, 149, 149 pertusatii (Verrucaria  7KHOLGLXPSHUWXVDWLL  petraeum var. lavatum (Rhizocarpon ( ) (=RhizoFDUSRQODYDWXP   petraeus var. obscuratus (Lichen ( ) (=RhizoFDUSRQODYDWXP   3KDHRSK\VFLD  phonolithicum (Dermatocarpon) (=D. leptoSK\OORGHV   3K\VFLD  pissina (Verrucaria) (=V. pachyderma) 179 placodioides (Trapelia) 30 3ODF\QWKLXP  plicatile (Collema  /HSWRJLXPSOLFDWLOH 77 SOLFDWLOH/HSWRJLXP   77, 78 plicatile var. hydrocharum ((Leptogium) (=L. plicatile) 77 plicatilis (Lichen  /HSWRJLXPSOLFDWLOH   pluriseptatum (Lecothecium  3ODF\QWKLXP GROLFKRWHUXP   pluriseptatum ((Placynthium  3GROLFKRWHUXP  SOXYLXP7KHOLGLXP   137 149, 149,  polyanthes (Lempholemma) 7, 33, 73, 74 polyanthes (Lichen) (=Lempholemma polyanthes) 73 3RO\EODVWLD  SRO\SRUL$QLVRPHULGLXP   polytropa (Lecanora)  3RULQD  3RURF\SKXV  3RUSLGLD  praetermissa (Leiophloea  9HUUXFDULD SUDHWHUPLVVD   SUDHWHUPLVVD9HUUXFDULD    3VHXGDUWKRS\UHQLD  pseudocatalepta (Verrucaria) (= ? V. aethiobola)  psorodeum (Dermatocarpon) (=Endocarpon SVRURGHXP   SVRURGHXP(QGRFDUSRQ 54, 54, 55

psorodeum (Verrucaria) (=Endocarpon psoroGHXP   3VRURWLFKLD  3WHU\JLRSVLV  pubescens DXFWEphebe  (ODQDWD   pubescens var. complicata (Ephebe) (=E. ODQDWD   pulvinata (Verrucaria   SXVLOOXP(QGRFDUSRQ   pusillum var. aquale (Leptogium (  /DTXDOH  pyracea f. submersa (Caloplaca) (=Caloplaca DWURÀDYDYDUVXEPHUVD   pyracea f. submersa (Lecanora) (=Caloplaca DWURÀDYDYDUVXEPHUVD   pyraceum f. submersum (Placodium) (=CalopODFDDWURÀDYDYDUVXEPHUVD   3\UHQRFDUSRQ  3\UHQRFROOHPD  S\UHQRSKRUXP7KHOLGLXP   3\UHQRSVLV  pyrenopsoides (Collema  3RURF\SKXV coccodes) 101 pyrenopsoides (Gabura  3RURF\SKXV coccodes) 101 pyrenopsoides (Lecanora (  3RURF\SKXV coccodes) 101 pyrenopsoides (Psorotichia  3RURF\SKXV coccodes) 101 UDGLRVD/RERWKDOOLD   rechingerii (Verrucaria  "+\GURSXQFWDULD UKHLWKURSKLOD"9DTXDWLOLV   UHGXFWXP5KL]RFDUSRQ   refugii (Verrucaria  "+\GURSXQFWDULD UKHLWKURSKLOD   rehmica (Psoropsis  3RURF\SKXVUHKPLFXV  rehmica (Psorotichia  3RURF\SKXVUHKPLFXV  UHKPLFXV3RURF\SKXV   102, 103 rehmii (Involucrothele  7KHOLGLXPUHKPLL  UHKPLL7KHOLGLXP   137151, 151 retecta (Verrucaria  9DTXDWLOLV   UKHLWKURSKLOD+\GURSXQFWDULD   64, 64, 65 5KL]RFDUSRQ  UKRGRVWLFWD&U\SWRWKHOH   rhodostictaDXFWPHGLHXUPyrenopsis) SS 3VDQJXLQHD3VXEDUHRODWD   Rinodina 23 riparia (Collemopsis  3RURF\SKXVUHKPLFXV  riparia (Psorotichia  3RURF\SKXVUHKPLFXV  riparia (Sagedia)  7KHOLGLXPIRQWLJHQXP   riparium (Spermatodium  7KHOLGLXPIRQWLJHQXP   riparium (Thelidium  7IRQWLJHQXP   riparius (Porocyphus  3UHKPLFXV   

,QGH[RIOLFKHQQDPHV   riparius (Pyrenopsis  3RURF\SKXVUHKPLFXV  ULYDOH/HSWRJLXP   rivale (Polychidium  /HSWRJLXPULYDOH  rivale (Thelidium  3RO\EODVWLDULYDOLV   ULYDOLV3RO\EODVWLD  94, 94 rivalis (Verrucaria  "9VXEPHUVHOOD    rivulare (Leptogiopsis (  /HSWRJLXPULYXODUH 79 ULYXODUH/HSWRJLXP   rivulare (Leptogium ( DXFWEULW &ROOHPD GLFKRWRPXP   rivularis (Gabura  /HSWRJLXPULYXODUH   rivularis (Lichen  /HSWRJLXPULYXODUH   ULYXODULV3VHXGDUWKRS\UHQLD   108 rivulicola (Arthopyrenia ( ) (= ? Thelidium rivulicolum   rivulicola (Verrucaria) (=Thelidium rivulicolum   ULYXOLFROXP7KHOLGLXP   ULYXORUXP 'HUPDWRFDUSRQ  53, 219 rivulorum (Endocarpon) (=Dermatocarpon ULYXORUXP   rivulorum (Placidiopsis) (=Dermatocarpon ULYXORUXP   rodnense (Thelidium  7IRQWLJHQXP   roseola (Gyalecta  *\DOLGHDURVHROD   URVHROD*\DOLGHD   URVXODQV3ODF\QWKLXP  89, 90 rubiginosa (Verrucaria) (=Porina lectissima) 99 rufescens (Psorotichia (  3RURF\SKXVUHKPLFXV  UXSHVWULV3URWREODVWHQLD   rusticula (Biatora  /DKOHVLL   rusticula (Lecidea  /DKOHVLL   sagedioides (Verrucaria  +\GURSXQFWDULD UKHLWKURSKLOD   sanguinea (Psorotichia) (=Pyrenopsis VDQJXLQHD   VDQJXLQHD3\UHQRSVLV  117, 223 saxicola (Arthopyrenia ( ) (=Pyrenocollema VD[LFROD   saxicola (Leiophloea) (=Pyrenocollema VD[LFROD   saxicola (Naetrocymbe) (=Pyrenocollema VD[LFROD   VD[LFROD3\UHQRFROOHPD   114, 114,  saxicola (Sagedia) (=Pyrenocollema saxicola)  saxicola (Verrucaria) (=Pyrenocollema saxicola)  saxicolum (Spermatodium) (= Pyrenocollema VD[LFROD   scabra +\GURSXQFWDULD  64 67 217 scabra (Verrucaria  +\GURSXQFWDULDVFDEUD 

VFKLEOHUL7KHOLGLXP   schistosa (Verrucaria  9DTXDWLOLV   scholanderi (Arctoheppia ( ) (= Thelignya OLJQ\RWD   scholanderi (Fernaldia) (= Thelignya OLJQ\RWD   VFLDVWUD3KDHRSK\VFLD   scotinodes (Verrucaria  +\GURSXQFWDULD UKHLWKURSKLOD   septemseptata (Porina  3DKOHVLDQD   septemseptata (Segestria) (=Porina ahlesiana)  sernanderi (Leptogium (  /ULYXODUH   silesiaca (Staurothele  6¿VVD   silicea (Verrucaria  9IXQFNLL   silicicola (Verrucaria  9IXQFNLL   solvens (Polyblastia (  6WDXURWKHOHVROYHQV  VROYHQV6WDXURWKHOH  132, 132, 133 solvens (Verrucaria  6WDXURWKHOHVROYHQV   soredizodes (Haplocarpon) (=Porpidia VRUHGL]RGHV   soredizodes (Huilia) (=Porpidia soredizodes)  VRUHGL]RGHV3RUSLGLD  106, 106 sparsulum (Thelidium  7]ZDFNKLL   sparsulum (Verrucaria  7KHOLGLXP]ZDFNKLL  6SRURGLFW\RQ  6WDXURWKHOH  VWLJPDWHD/HFLGHOOD   suaveolens (Gyalecta  ,RQDVSLVVXDYHROHQV 70 VXDYHROHQV,RQDVSLV   70, 213, 223 suaveolens (Patellaria  ,RQDVSLVVXDYHROHQV 70 suaveolens (Urceolaria  ,RQDVSLVVXDYHROHQV 70 suaveolens f. odora (Ionapsis ( )  ,RGRUD 70 subarenaria (Rinodina  5R[\GDWD   subareolata (Collemopsis) (=Pyrenopsis VXEDUHRODWD   VXEDUHRODWD3\UHQRSVLV 117, 118, 221 subcontinua (Involucrothele) (=V. margacea / 9DSSODQDWD   subcontinuum (Thelidium) (=V. margacea / 9DSSODQDWD   subgelatinosum (Verrucaria  7KHOLGLXP ]ZDFNKLL   subhydrela (Verrucaria) (=V. hydrela) 171,  sublacteum ((Paraphysothele)  $QLVRPHULGLXP FDULQWKLDFXP   sublacteum (Thelidium  $QLVRPHULGLXP FDULQWKLDFXP   VXEOREXODWD9HUUXFDULD   160 182, 183, 219 sublobulata f. robustior (Verrucaria) (=V. hydrela) 171 submauroides (Verrucaria) (= ? V. hydrela)  submersa 6FKDHUHUQRQ%RUUHU (Verrucaria)  9VXEPHUVHOOD  

  ,QGH[RIOLFKHQQDPHV submersa var. franconia (Verrucaria)  9VXEPHUVHOOD   submersa ȕlitorea (Verrucaria  9VXEPHUVHOOD   VXEPHUVHOOD9HUUXFDULD   160,  184, 184 submethoria (Verrucaria  7KHOLGLXPVXE PHWKRULXP   VXEPHWKRULXP7KHOLGLXP   152, 153 VXESDOOLGD&DORSODFD   subtorulosum (Collema  /HSWRJLXPVXE WRUXORVXP   VXEWRUXORVXP/HSWRJLXP  80, 81,  succedens (Polyblastia (  6WDXURWKHOHVXFFHdens)  VXFFHGHQV6WDXURWKHOH   134, 134 succedens (Verrucaria  6WDXURWKHOHVXFFHGHQV  VXSHUWHJHQV$VSLFLOLD  39, 39 supertegens (Lecanora ( ) 39 suzaeanum (Involucrothele  "7KHOLGLXP SDSXODUH   suzaeanum (Thelidium  "7SDSXODUH    sylvatica (Aspicilia (  $ODHYDWD   sylvatica (Lecanora  $VSLFLOLDODHYDWD   tantalea (Racoblenna (  3ODF\QWKLXPWDQWDOHXP 91 WDQWDOHXP 3ODF\QWKLXP  91,  221 tapetica var. ÀXYLDOLV (Verrucaria) (=V. praeWHUPLVVD   tatrensis (Verrucaria  "9VXEPHUVHOOD  WHLFKRSKLOD5LQRGLQD   tenella (Physcia) 31 WHUHWLXVFXOXP/HSWRJLXP   terrestre (Thelidium  7PLQXWXOXP   teutoburgensis (Verrucaria) (=V. elaeomelaena) 177 7KDOOLQRFDUSRQ  7KHOLGLXP  7KHOLJQ\D  thelostomum (Thrombium) (= ? Pyrenocarpon ÀRWRZLDQXP   tichothecioides (Arthopyrenia ( ) (=PyrenocolOHPDWLFKRWKHFLRLGHV   WLFKRWKHFLRLGHV3\UHQRFROOHPD   tichothecioides (Xanthopyrenia ( ) (=PyrenocolOHPDWLFKRWKHFLRLGHV   WLUROLHQVH7KHOLGLXP   tiroliensis (Verrucaria) (= ? V. elaeomelaena)  WUDFKRQD%DFLGLD   tremelloides DXFWLeptogium  ) (=L. cyanescens)  tremelloides var. caesium (Leptogium ( ) (= L. F\DQHVFHQV  

WXEHUFXORVDPorpidia   tunaeforme (Collema  &IXVFRYLUHQV   tuniforme (Collema  &IXVFRYLUHQV   turicensis =VFKDFNHQRQ:LQWHU 6WL]HQEHUJHU (Verrucaria  9VXEPHUVHOOD   Umbilicaria 31 Umbilicata (Pyrenopsis  "7KHOLJX\D OLJQ\RWD   XPERQDWD0HWDPHODQHD   umbonata (Pyrenula) (= ? Pyrenocarpon ÀRWRZLDQXP   umbrina (Polyblastia  6WDXURWKHOHFORSLPD  umbrina (Sphaeromphale  6WDXURWKHOH FORSLPD   umbrina (Staurothele  6FORSLPD   umbrina (Verrucaria  6WDXURWKHOHFORSLPD  valentior (Biatora  /HFLGHDDKOHVLL   valentior (Lecidea  /DKOHVLL   YDOOLVÀXHODH (Verrucaria) (= ? V. margacea)  9HUUXFDULD  YLQGRERQHQVLV9HUUXFDULD   viride Eitner non Deakin (Thelidium) (=T. PLQXWXOXP   viridicana (Verrucaria) (=V. praetermissa)  viridis (Staurothele  6¿VVD   YLULGXOD9HUUXFDULD   YLWHOOLQD&DQGHODULHOOD   vitricola (Verrucaria  9DTXDWLOLV   vivariensis (Porocyphus) (= P. coccodes) 101 ZHEHUL(Dermatocarpon  'OXULGXP   ZLPPHULDQD.RHUEHULHOOD  71, 71 ZLPPHULDQD (Lecanora) (=Koerberiella wimmeriana) 71 ZLPPHULDQD (Zeora) (=Koerberiella wimmeriana) 71 ZROIHUL(Verrucaria  "+\GURSXQFWDULD UKHLWKURSKLOD   xylospila (Verrucaria  7KHOLGLXP]ZDFNKLL  xylospilum (Thelidium  7]ZDFNKLL   zahlbruckneri (Verrucaria  9VXEPHUVHOOD  zenogensis (Verrucaria) (=V. margacea / V. DSSODQDWD   zschackei (Verrucaria) (=V. calcaria)  ]ZDFNKLLL (Pyrenula  7KHOLGLXP]ZDFNKLL   ]ZDFNKLL (Sagedia  7KHOLGLXP]ZDFNKLL   ]ZDFNKLLL (Segestria  7KHOLGLXP]ZDFNKLL   ]ZDFNKLL7KHOLGLXP  154, 154 ]ZDFNKLL (Verrucaria  7KHOLGLXP]ZDFNKLL  ]ZDFNKLLȕWR¿FROD(Sagedia  7KHOLGLXP IRQWLJHQXP  

Index to Figures (alphabetically arranged by lichen species name) Figure numbers in bold standard: photos. Figure numbers in bold italics: line drawings of involucrellum outlines.

Anisomeridium carinthiacum: Germany, Hesse, Vogelsberg, in streamlet Graswiesenbach, 20. Mar. 2003, H. Thüs W0816 )5 ¿J 10. Aspicilia aquatica: Russia, Karelia, distr. Kandalakcha, in stream Tchornaja Retchka, 16. Aug. 1997, M. Schultz 13100b (herb. 6FKXOW] ¿J11. Aspicilia laevata: Slovakia, Vysoké Tatry, Bielovodska Dolina, Sep. 2005, H. Thüs 7)5 ¿J12. Aspicilia supertegens: Austria, Tyrol, Ötztaler Alpen, Pitztal, Seebach unterhalb des Riffelsees, 25. Aug. 1961, J. Poeltt & M. Steiner0 ¿J r 13. Bacidina inundata: Austria, Carinthia, Karnische Alpen, S of Kötschach-Mauthen, Angerbach, 31. Aug. 2007, M. Schultz KHUE6FKXOW] ¿J 14. &DORSODFDDWURÀDYD var. submersa: Germany, Bavaria, Oberpfalz, between Süssenbach and Kleinanglhof, rocks in river Regen, 30. Aug. 2000, O. Dürhammerr 2039 (herb. 'UKDPPHU ¿J162 (colour plate 4). Caloplaca isidiigera: Germany, SchleswigHolstein, Wedel, Willkomm-Höft, slack boulders at bank of river Elbe, 21. Apr. 2008, M. Schultz 07143a (herb. Schultz); ¿J 161 (colour plate 4). Collema dichotomum: Scotland, Forfar, River Isla, 12. Aug. 1968, P.W. James %0 ¿J 15. Collema dichotomum: Austria, Upper Austria, Mühlviertel, Waldaistal, Guttenbrunner Leiten, 2. Oct. 1994, F. Bergerr 8012 (herb. %HUJHU ¿J16. &ROOHPDÀDFFLGXP: Austria, Carinthia, Karnische Alpen, Angerbach valley, mossy boulder along streamlet, 31. Aug. 2007, M. Schultz KHUE6FKXOW] ¿J17. Collema fuscovirens: Germany, Baden-Württemberg, Neckarsteinach, amphibious along ULYHU1HFNDUSLFWXUHWDNHQLQWKH¿HOGE\ R.Duque-Thüs6HS¿J18. Collema glebulentum: Czech Republic, eastern Sudety, Jeseníky, 3. Sept. 1969, $9Č]GD, /LFKVHOH[V0 ¿J 19. Dermatocarpon arnoldianum: Austria, Schladminger Tauern, Preberkessel, in streamlet, 25. Oct. 2006, H. Thüs & R. Türk :%0 ¿J 20.

Dermatocarpon leptophyllodes: Czech 5HSXEOLF%RKHPLD.Otþ$XJ J. Anders (PRM-704352, lectotype of D. lorenzianum $QGHUV ¿J 21. Dermatocarpon luridum: Germany, Bavaria, Spessart, Lauberbach, 12. April 1989, LPDJHWDNHQLQWKH¿HOGE\B. Büdel; ¿J159 (colour plate 4). Dermatocarpon meiophyllizum: Germany, Baden-Württemberg, Südschwarzwald, river Alb, 8. May 2004, H. Thüs W543b %0 ¿J22. Dermatocarpon rivulorum: Austria, Schladminger Tauern, stream in Preberkessel, 25. Oct. 2006, H. Thüs & R. Türkk W1826 %0 ¿J160 (colour plate 4). Endocarpon psorodeum: Germany, RhinelandPalatinate, St. Goar, rocky banks of the river Rhine, May 2006, H. Thüs 1492 %0 ¿J23. Endocarpon psorodeum: Germany, BadenWürttemberg, Eberbach, rocky banks of the river Neckar, June 2006, H. Thüs 1285 %0 ¿J24. Endocarpon psorodeum: Endocarpon psorodeum: Germany, Rhineland-Palatinate, Mainz, bank reinforcement of the river Rhine, Aug. 2007, H. Thüs 1678 (BM); ¿J25. Ephebe lanata: Austria, eastern Tyrol, along margin of mountain streamlet, 3. Sept. 1998, H. Sipman 43604 (B; accomp. E. multispora ¿J26. Ephebe multispora: Austria, eastern Tyrol, margin of mountain streamlet, 3. Sept. 1998, H. Sipman % ¿J27. Gonohymenia heppii: Switzerland, Wallis, Boverniche, river Dranse, 18. July 1872, J. Müllerr $UJDX* ¿J 28. Gyalidea diaphana: Czech Republic, Krkonoše/Giant Mts., Blaugrund, 1933, V. Kut`ák/LFK%RKHP+%* ¿J 29. Gyalidea fritzei: Czech Rep., Krkonoše/Giant Mts., Blaugrund, 1934, V. Kut`ákk (in: 9Č]GD/LFKVHO([V 0 ¿J 149 (colour plate 1). Gyalidea hyalinescens: Germany, Baden, Geroldsauer Wasserfall, Sept. 1858, F. Arnold d (M, holotype of Secoliga carnea); ¿J150 (colour plate 1).

206

Index to Figures

Hydropunctaria rheitrophila: Germany, Erzgebirge, in river Flöha, 2003, H. Thüs ()5 ¿J 30. Hydropunctaria rheitrophila: Germany, Odenwald, between Hebstahle and Gaimühle, Sensbach, submersed in stream, 01. June 1988, M. Weißbeckerr 1430 (herb. (LFKOHU &H]DQQH ¿J31. Hydropunctaria rheitrophila: Germany, Mecklenburg-Vorpommern, Ludwigslust, Schlosspark, Wasserspiele „24 Sprünge“, 24. May 2008, U. Schiefelbein (herb. 6FKLHIHOEHLQ ¿JV 34–36. Hydropunctaria scabra: Slovakia, Tatra Minor, Košariská mountains, upper part of the valley Biela voda, alt. 1300 m, 7. Oct 1966, $9Č]GD, Lich. Sel. 876 (W–isoW\SH ¿J 32. Hydropunctaria scabra: Austria, Schladminger Tauern, Preberkessel, cascades, H. Thüs & R. Türk:%0 ¿JV k 33, 154 (colour plate 3). Hymenelia cyanocarpa: Italy, Trentino-Alto Adige, Conca della Levade, 2006, J. Nascimbene KHUE1DVFLPEHQH ¿J 148 (colour plate 1). Ionaspis lacustris: Germany, Baden-Württemberg, Black Forest, near Altglashütten, in stream, 19. Sept. 1997, M. Schultz 03041e KHUE6FKXOW] ¿J 145 (colour plate 1). Ionaspis odora: Austria, Carinthia, Karnische Alpen, S of Tröplach, streamlet below Gartnerkofel towards Watschiger Alm, 1. Sept. 2007, M. Schultz 08333 (herb. 6FKXOW] ¿J 146 (colour plate 1). Ionaspis odora (Trentepohlia-photobiont, isolated culativate from lichen thallus): Switzerland, Grisons, Fluela valley, in a small streamlet, 26. Aug. 2003, H. Thüs & C. Kellerr :%0 ¿J 169 (colour plate 6). Ionaspis suaveolens: Austria, Carinthia, Karnische Alpen, S of Tröplach, streamlet below Gartnerkofel towards Watschiger Alm, 1. Sept. 2007, M. Schultz 08331 KHUE6FKXOW] ¿J 147 (colour plate 1). Ionaspis suaveolens (Trentepohlia-photobiont in lichen thallus): Switzerland, Berner Oberland, Sustenpass, 26. Aug. 2006, V. Wirth .5 ¿J168 (colour plate 6). Koerberiella wimmeriana: Switzerland, Uri, Oberalppass, stream, 19. Sept. 1992, G. Ernstt +%* ¿J 37. Lecidea ahlesii: Germany, Hesse, Vogelsberg, Schwarzer Fluss, Feb. 2004, H. Thüs :%0 ¿J 38. Lempholemma polyanthes: England, northwest York, on mosses at river bank, 31. July 1970, B. Coppins %0 ¿J39. Leptogium aquale: Austria, Tyrol, Waldrast, d Exs. Matrai, 8. Aug. 1872, F.C.G. Arnold, 0±KRORW\SH ¿J 40.

Leptogium cyanescens: Georgia, western Caucasus, valley of river Zapadnaia Gumista, 13. July 1980, V. Vašák, Iter &DXFDV6HF0 ¿J41. Leptogium plicatile: Austria, Upper Austria, Danube valley, Schlögener Schlinge, PLJPDWLWHERXOGHUVÀXVKHGE\ZDYHV 0DULPDJHWDNHQLQWKH¿HOGE\ F. Berger¿J 42. Leptogium rivale: Czech Rep., Krkonoše, Horni úpský vodopád, 27. July 2001, J. Halda & Z. Palice KHUE3DOLFH ¿J 43. Leptogium rivulare: Sweden, E.M. Fries, Lich. 6XHF0 ¿J 44. Leptogium subtorulosum: France, HauteSavoie, Villette, river Arve, 1879, J. Rome * ¿J 45. Lobothallia melanaspis: Slovakia, Magás Tatra, Malompatak, July 1914, J. Szurák & G. Timkó, Fl. Hung. Exs. 317 (HBG); ¿J 46. Phaeophyscia endococcina: Austria, Schladminger Tauern, 25. Oct. 2006, H. Thüs & R. Türkk :%0 ¿J 47. Placynthium dolichoterum: Scotland, Ben Lawers, 17. Aug. 1880, J. Stirton (BM, type of Pannaria melantera6WLUWRQ ¿J48. 3ODF\QWKLXPÀDEHOORVXP: France, Vosgues, Le Tanet/Tanneck, Col de la Schlucht, 9. Sept. 1971, V. Wirth678 ¿J49. Placynthium nigrum f. crustaceum: Switzerland, Graubünden, Davos, Fluela stream, 22. July 2003, H. Thüs & C. Kellerr 1136 %0 ¿J50. Placynthium pannariellum: Russia, Karelia, distr. Kandalkscha, shore of Nilma lake, 11. Aug. 1996, M. Schultz 13006d (herb. 6FKXOW] ¿J 51. Placynthium rosulans: Sweden, G.E. Du Rietz D0 ¿J 52. Placynthium tantaleum: Germany, Bavaria, Roßhaupten, Lech ravin, 1951, A. Schröppel 0 ¿J 163 (colour plate 5). Polyblastia peminosa: Romania, Hunyad, Lake Zenoga, 9. Aug. 1873, H. Lojka 2475 0 ¿J53 Polyblastia rivalis: Austria, Tyrol, Waldrast, F. Arnold([V0±KRORW\SH ¿J d 54. Porina ahlesiana: Romania, 12. Aug. 1872, H. Lojka 0 ¿J55. Porina chlorotica: Germany, Hesse, Vogelsberg, Schwarzer Fluss, Feb. 2004, H. Thüs :%0 ¿J56. Porina grandis: Germany, Black Forest, in river Präg, 23.11.1955, A. Riedd 138/I V9 )5 ¿J57. Porina guentheri: Germany, Hesse, Vogelsberg, Schwarzer Fluss, Feb. 2004, H.Thüs :)5 ¿J58. Porina lectissima: Germany, Hesse, Vogelsberg, Schwarzer Fluss, Feb. 2004, H. Thüs :%0 ¿J59.

,QGH[WR¿JXUHV   Porocyphus coccodes: Poland, Karkonozse/ Giant Mts., Lomnitz Fall, G.W. Körber, Lich. sel. Germ. 29 (M, isotype of P. cataractarum.|UEHU ¿J60. Porocyphus rehmicus: Austria, Wachau, Spitz, WHPSRUDULO\ÀXVKHGEDQNRI'DQXEH Aug. 1969, V. Wirth 678 ¿J 61. Porpidia hydrophila: Germany, Lower Saxonia, Harz Mts., Braunlage, Odertal, 9. Oct. 2004, V. Wirth .5 ¿J62. Porpidia soredizodes: Germany, Hesse, Vogelsberg, Schwarzer Fluss, Feb. 2004, H. Thüs )5 ¿J 63. Pterygiopsis concordatula: Norway, Trondheim, Ilsvigen, C. Kindtt 29 (UPS-349845); ¿J 164 (colour plate 5). Pterygiopsis concordatula (single-celled cyanobacterial photobiont): Czech Rep., Krkonoše/Giant Mts., Velkýkotel, 31. Aug. 2002, Z. Palice 10520 (herb. Palice); ¿J 171 (colour plate 6). Pterygiopsis neglecta: Germany, SchleswigHolstein, Wedel, Willkomm-Höft, slack boulders at bank of River Elbe, 21. Apr. 2008, M. Schultz 07143b (herb. Schultz); ¿J 64. 3\UHQRFDUSRQÀRWRZLDQXP: Switzerland, Lake Zurich, P. Hepp, Fl. Eur. 92 (M–isotype); ¿J65. Pyrenocollema saxicola: Germany, Northrhine-Westfalia, river Anger, June 1999, H. Thüs :%0 ¿J 66. Pyrenopsis sanguinea: Germany, BadenWürttemberg, Schönau, ”Haideck”, 30. Sept. 1971, V. Wirth 678 ¿J67. Pyrenopsis sanguinea (Gloeocapsa-like photobiont): Germany, Baden, V. Wirth 26037 678 ¿J 170 (colour plate 6). Pyrenopsis subareolata: Germany, BadenWürttemberg, Menzenschwand, 2. Oct. 1969, V. Wirth 678 ¿J 68. Pyrenopsis subareolata: France, Vosgues, Le Tanet/Tanneck, Col de la Schlucht, 9. Sept. 1971, V. Wirth 2951 (STU); ¿J165 (colour plate 5). Rhizocarpon amphibium: Sweden, Värmland, Norra Finnskoga, ravin of River Tåsan, 8. Aug. 1988, L.-E. Muhrr 11283, Lich. Exs. 8SVDO0 ¿J 69. Rhizocarpon badioatrum: Switzerland, Graubünden, Davos, Fluela valley, 22. July 2003, H. Thüs & C. Kellerr W1150 (BM); ¿JV70–71. Rhizocarpon lavatum: Germany, Bavaria, Allgäuer Alpen, Oberstdorf, wet rock, 23. Sept. 2006, M. Schultz 08227 (herb. 6FKXOW] ¿J72. 5LQRGLQD¿PEULDWD: Germany, Hesse, Odenwald, in Moldau river, 11. Mar. 2000, H. Thüs 2GZ)5 ¿J 73. Rinodina oxydata: France, Vosgues, J. Harmand/LFK*DOO)5 ¿J d 74.

Sporodictyon cruentum: Slovakia, Vysoké Tatry, Litvorová Kotlina, Sept. 2005, H. Thüs )5 ¿J75. Staurothele areolata: Switzerland, Berner Oberland, Sustenpass, 26. Aug. 2006, H. Thüs:%0 ¿J 153 (colour plate 2). Staurothele clopima: Austria, Schadminger Tauern, stream in Preberkessel, 25. Oct. 2006, H. Thüs & R. Türkk W1815 (BM); ¿J76. Staurothele clopima: Austria, Schadminger Tauern, stream in Preberkessel, 25. Oct. 2006, H. Thüs & R. Türk:%0 ¿J k 77. Staurothele clopimoides: Austria, Schladminger Tauern, 25. Oct. 2006, H. Thüs & R. Türk :%0 ¿J151 (colour plate 2). Staurothele clopimoides: Austria, Schadminger Tauern, stream in Preberkessel, 25. Oct. 2006, H. Thüs & R. Türkk W1826 (BM); ¿J 78. 6WDXURWKHOH¿VVD: Slovakia, Vysoké Tátry, Bielovodská Dolina, Sept. 2005, H. Thüs )5 ¿J79. Staurothele frustulenta: Germany, RhinelandPalatinate, St. Goar, rocky banks of the river Rhine, May 2006, H. Thüs W1999 %0 ¿J 80. Staurothele frustulenta: Germany, Hamburg, Övelgönne, slack boulders at bank of river Elbe, 2007, M. Schultzz 07148 (herb. Schultz); ¿J 152 (colour plate 2). Staurothele solvens: Germany, southern Bavaria, Roßhaupten, Lech ravin, 1952, M. Servítt (M, type of Staurothele meylanii f. geographicum 6HUYtW ¿J81. Staurothele solvens: Germany, southern Bavaria, Roßhaupten, Lech ravin, O. Klementt (M, type of Staurothele meylanii f. papulare.OHPHQW ¿J82. Staurothele solvens: Germany, southern Bavaria, Roßhaupten, Lech ravin, O. Klement (M, type of Staurothele meylanii f. superiorr .OHPHQW ¿J83. Staurothele succedens: Germany, Bavaria, Allgäuer Alpen, Hinanger Wasserfall, Aug. 2005, H. Thüs :%0 ¿J 84. Thelidium aethioboloides: Germany, Bavaria, Allgäuer Alpen, Faltenbach in Oberstdorf, cataract near ski-jump, July 2005, H. Thüs :%0 ¿J91. Thelidium aethioboloides: Germany, Bavaria, Ammergauer Alpen, Lindertal, on limestone pebbles in stream, 5. Sept. 2004, M. Schultz FKHUE6FKXOW] ¿J156 (colour plate 3). Thelidium aquaticum: Czech Rep., Bohemia, RudohoĜí, 1950, M. Servítt (PRM-842845– W\SH ¿J92. Thelidium circumspersellum: Romania, Hunyadin, Ponor-Ohaba, 1874, H. Lojka 3384 (W, type off T. circumspersellum f. lojkanum 6]DWDOD ¿J93.

208

Index to Figures

Thelidium fontigenum: Germany, Bavaria, Franconia, Diedenhofen, on irrigated wood, H. Rehm, ex Herb. Mass., Anzi Lich Rar. 9HQHW:±LVRW\SH ¿J85. Thelidium fontigenum: Switzerland, Berner Oberland, streamlet in Rosenlauital, 27. Aug. 2006, H. Thüs :%0 ¿J94. Thelidium inundatum: Germany, Bavaria, Allgäuer Alpen, Oberstdorf, Faltenbachtobel, G. Lettau % ¿J 95. Thelidium klementii: Gemany, Bavaria, Roßhaupten, Illasberg, Lech ravine, on temporarily inundated rocks, 1953, O. Klement0±KRORW\SH ¿J t 96. Thelidium lahmianum: Slovakia, Vysoké Tátry, Koprova valley, 21. Sept. 1868, H. Lojka: ¿J97. Thelidium methorium: Norway: Sør-Trøndelag, Dronthjem, W.P. Schimperr (H-NYL 2121– KRORW\SH ¿J 98. Thelidium methorium: Switzerland, Grisons, Fluela valley, in a small streamlet, H. Thüs )5 ¿JV88, 99. Thelidium minutulum: Germany, BadenWürttemberg, Neckar valley, Wolfsschlucht close to Zwingenberg, 28. Mar. 2006, H. Thüs:%0 ¿JV 89, 100. Thelidium nigricans: Switzerland, Berner Oberland, streamlet in Rosenlauital, 27. Aug. 2006, H. Thüs & R. Duque-Thüs :%0 ¿J101. Thelidium papulare: Germany, Bavaria, Allgäuer Alpen, in river Breitach, July 2005, H. Thüs:%0 ¿J102. Thelidium pertusatii: Italy, Trentino-Alto Adige, Val de la Mare, J. Nascimbene 1541 KHUE1DVFLPEHQH ¿J87, 103. Thelidium pluvium: Austria, Styria, in river Teigetsch, 25. May 2004, H. Thüs W1383 %0 ¿J 86, 104. Thelidium rehmii: Germany, Baden-Württemberg, Odenwald, stream in canyon close to Reichenbach, 28. Mar. 2006, H. Thüs :)5 ¿J 90, 105. Thelidium submethorium: Russia, Karelian ASSR, ad saxa gneissacea in rivulo Kiekkipuro, 1877, E.A. Vainio (H–isotype); ¿J106. Thelidium submethorium: Italy, Trentino-Alto Adige, Paneveggio Nat. Park, Valbona, subalpine spruce forest, periodically submerged, Oct. 2005, J. Nascimbene JN1991 KHUE1DVFLPEHQH ¿J107. Thelidium zwackhii: Austria, Carinthia, Karnische Alpen, streamlet contributing to Valentinbach near Eder SW of KötschachMauthen, 29. Aug. 2007, M. Schultz EKHUE6FKXOW] ¿J 108. Thelignya lignyota: Sweden, H. Magnusson, /LFK+%* ¿J 109. Verrucaria aethiobola: type specimen +±$&+ ¿J 113.

Verrucaria aethiobola: Germany, Black Forest, in river Wiese, Sep. 1998; H. Thüs :)5 ¿J114. Verrucaria aethiobola: Russia, Karelia, distr. Kandalakscha, Kindra Peninsula, in stream, 19. Aug. 1996, M. Schultz 13071e (herb. 6FKXOW] ¿J129. Verrucaria aquatilis: Germany, NorthrhineWestfalia, streamlet contributing to river Alme, Oct. 2006, H. Thüs W1628 (BM); ¿J 130. Verrucaria calcaria: Germany, Bavaria, Frankonia, Eichstätt, F. Arnoldd (M–isoW\SH ¿JV 115, 131. Verrucaria consociata: Czech Republic, RudoKRĜt6XQLSHUN)OHFNHOPKOHLQVWUHDP 350±W\SHVSHFLPHQ ¿J110. Verrucaria consociata: Austria, Mühlviertel, Rannatal, on pebbles in stream, 26. Feb. 2008, F. Bergerr 22600 (herb. Berger); ¿J 132. Verrucaria dolosa: Germany, Baden-Württemberg, streamlet in Heppenheimer Klinge, Nov. 2004, H. Thüs :%0 ¿JV 116, 133. Verrucaria funckii: Germany, Bavaria, Fichtelgebirge, C.P.J. Sprengell in: Funck, Krypt. *HZ1U)5±OHFWRW\SH ¿J 119. Verrucaria funckii: Austria, Schadminger Tauern, stream in Preberkessel, 25. Oct. 2006, H. Thüs & R. Türk W1815 (BM); ¿J 120, 134. Verrucaria hydrela: Sweden, E. Acharius +±$&+KRORW\SH ¿J 117. Verrucaria hydrela: Germany, Lower Saxony, Oldenburg, Hasbruch, H. Sandstede, Arnold Lich. Exsic.1712b (B, isotype of V. denudata =VFKDFNH ¿J 118. Verrucaria hydrela: Austria, Oberösterreich, Kobernaußerwald, in streamlet contributing to Schwarzmoosbach, F. Bergerr 22520 (herb. Berger, accomp. V. sublobulata); ¿JV 157–158 (colour plate 4). Verrucaria latebrosa: Poland, Karkonozse/ Giant Mts., Kleine Schneegrube, 27. July r 127. 1853, G.W. Körber)5±LVRW\SH ¿J Verrucaria latebrosa: Switzerland, Graubünden, Davos, in Fluela stream, 22. July 2003, H. Thüs & C. Keller:%0 ¿JV r 128, 135. Verrucaria madida: Germany, Baden-Württemberg, Schwarzwald, river Dreisam, 8. Aug. 2003, V. Wirth.5 ¿J 136. Verrucaria madida: Wales, Breconshire, in streamlet, A. Orange 10: ¿J155 (colour plate 3). Verrucaria margacea s.str.: Lapland, G. Wahlenberg836±KRORW\SH ¿J g 123. Verrucaria margacea s.str.: Switzerland, Berner Oberland, Sustenpass, on rocks in a stream near Steingletscher, 26. Aug. 2006, H. Thüs:%0 ¿J137.

,QGH[WR¿JXUHV   Verrucaria margacea s.l. (V. andesiatica): Ukraine, Uzhorod, Velké Lázy, J. Nádvorník (PRM-791756–holotype) Fig. 125. Verrucaria margacea s.l. (V. applanata): Germany, Baden-Württemberg, Heidelberg, W. Zwackh-Holzhausen Exs. 212 (FR– V\QW\SH ¿J124. Verrucaria margacea s.l. (V. elaeomelaena): Germany, Northrhine-Westfalia, spring of river Alme, Oct. 2006, H. Thüs W1631 %0 ¿JV126, 138. Verrucaria pachyderma: Austria, Tyrol, Oetztal, Umhauser Wasserfall, July 1870, F. Arnoldd 0±KRORW\SH ¿JV121, 139. Verrucaria pachyderma: Switzerland, Graubünden, Davos, in Fluela stream, 22. July 2003, H. Thüs & C. Kellerr W1750 %0 ¿JV122, 140.

Verrucaria praetermissa: Germany, Northrhine-Westfalia, Neandertal, Düssel, upstream of Thunisbrücke, Mar. 2003, H. Thüs:%0 ¿J 141. Verrucaria sublobulata: Czech Republic, .UNRQRãH*LDQW0WV-iQVNpOi]QČV. Kut`ákk 350±V\QW\SH ¿J142. Verrucaria sublobulata: Austria, Upper Austria, Kobernaußerwald, in streamlet contributing to Schwarzmoosbach, F. Bergerr 22520 (herb. Berger, accomp. V. hydrela ¿JV 112, 143, 157–158 (colour plate 4). Verrucaria submersella: Germany, Bavaria, Allgäuer Alpen, Kanzelwand close to Oberstdorf, in streamlet, July 2005, H. Thüs :%0 ¿JV 111, 144.

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Colour Plate 1. Fig. 145. Ionaspis lacustris ±DSRWKHFLDOGLVFVSDOHZKLWLVKî Fig. 146. I. odora –DSRWKHFLDOGLVFVSDOHZKLWLVKWRSDOHEURZQLVKî – Fig. 147. I. suaveolens±DSRWKHFLDZLWKEODFNLVKGLVFVî Fig. 148. Hymenelia cyanocarpa±DSRWKHFLDLPPHUVHGZLWKEOXHEODFNGLVFVî Fig. 149. Gyalidea fritzei±DSRWKHFLDZLWKGDUNEURZQSURSHUPDUJLQî Fig. 150. G. hyalinescens±DSRWKHFLDZLWKSDOHSURSHUPDUJLQî

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Colour Plate 2. Fig. 151. Staurothele clopimoides±7KDOOXVFRSSHUEURZQZLWKRUDQJHWLQJHî Fig. 152. S. frustulenta±7KDOOXVSDOHJUH\LVKEURZQZLWKGLVWLQFWSURWKDOOXVî Fig. 153. S. areolata±7KDOOXVGDUNEURZQî

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Colour Plate 3. Fig. 154. Hydropunctaria scabra ±WKDOOXVGDUNROLYHJUHHQî Fig. 155. Verrucaria madida ±WKDOOXVEODFNLVKROLYHî Fig. 156. Thelidium aethioboloides±WKDOOXVEURZQLVKROLYHî

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Colour Plate 4. Figs. 157–158. Verrucaria hydrela±VXEJHODWLQRXVZKHQZHWDQGV. sublobulata ± QRWVXEJHODWLRXVî±157:WKDOOLLQGU\VWDWH±158:WKDOOLLQZHWVWDWH Fig. 159. Dermatocarpon luridum±WKDOOXVIUHVKJUHHQZKHQZHWî Fig. 160. D. rivulorum ±WKDOOXVFRORXUQRWWXUQLQJIUHVKJUHHQZKHQZHWî Fig. 161. Caloplaca isidiigera±EOXLVKWLQJHLQLVLGLDWHSDUWî Fig. 162. C. DWURÀDYD YDU submersa ±RUDQJHDSRWKHFLDZLWKJUH\WKDOOLQHH[FLSOH DUURZ î

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Colour Plate 5. &RORXUVLQF\DQREDFWHULDOOLFKHQV Fig. 163. Placynthium tantaleum ±WKDOOXVGDUNROLYHJUHHQZLWKEOXLVKSURWKDOOXV DUURZ DQGEODFNDSRWKHFLDî Fig. 164. Pterygiopsis concordatula ±WKDOOXVEODFNLVKZLWKVHPLLPPHUVHGEODFN DSRWKHFLDî Fig. 165. Pyrenopsis subareolata±WKDOOXVGDUNUHGGLVKEURZQZLWKVRPHLPPHUVHG DSRWKHFLDZLWKSRUHOLNHGLVFVDUURZ î

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Colour Plate 6. Fig. 166. Heterococcus VSLQ Verrucaria funckii±YHUWLFDODUUDQJHPHQWRIWKH SKRWRELRQWFHOOVFKDUDFWHULVWLFIHDWXUHLQVSHFLDOLzHGDTXDWLFVerrucaria VSHFLHVî Fig. 167. 'LODEL¿OXPVS LQWKDOOXVRIVerrucaria aquatilis±SXUHJUHHQFHOOV î Fig. 168. Trentepohlia VS±SDOH\HOORZLVKFRORXUHGSKRWRELRQWFHOOVLQKHUEDULXP VSHFLPHQRIIonaspis suaveolensî Fig. 169. TrentepohliaVS±GLVWLQFWO\RUDQJHFRORXUHGOLYLQJFHOOVRILVRODWHG FXOWLYDWHIURPDWKDOOXVRIIonaspis odora î Fig. 170. GloeocapsaOLNHSKRWRELRQWLQ Pyrenopsis sanguinea±FHOOVVLQJOH ODUJHJHODWLQRXVVKHDWKUHGGLVKî Fig. 171. 6LQJOHFHOOHGF\DQREDFWHULDOSKRWRELRQWLQPterygiopsis concordatula ± FHOOVVLQJOHVPDOOJHODWLQRXVVKHDWK\HOORZLVKEURZQî

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