Ephemeroptera of South America (Aquatic Biodiversity of Latin America)

ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America

Aquatic Biodiversity in Latin America Biodiversidad Acuática en América Latina

Volume 2

Ephemeroptera of South America Volumen 2

Ephemeroptera de América del Sur

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ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur

BIODIVERSIDAD ACUÁTICA

EN

AMÉRICA LATINA

Volumen 2

EPHEMEROPTERA DE AMÉRICA DEL SUR por

Eduardo Domínguez, Carlos Molineri, Manuel L. Pescador, Michael D. Hubbard & Carolina Nieto Lector Científico a los Editores: Michel Sartori Editores de la Serie: Joachim Adis, Jorge R. Arias, Guillermo Rueda-Delgado & Karl Matthias Wantzen

Sofia–Moscow 2006

ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America

AQUATIC BIODIVERSITY IN LATIN AMERICA Volume 2

EPHEMEROPTERA OF SOUTH AMERICA by

Eduardo Domínguez, Carlos Molineri, Manuel L. Pescador, Michael D. Hubbard & Carolina Nieto Scientific Reader to the Editors: Michel Sartori Series Editors: Joachim Adis, Jorge R. Arias, Guillermo Rueda-Delgado & Karl Matthias Wantzen

Sofia–Moscow 2006

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ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur

AQUATIC BIODIVERSITY BIODIVERSIDAD ACUÁTICA

IN

LATIN AMERICA AMÉRICA LATINA

EN

Series Editors/Editores de la Serie: Joachim Adis, Jorge R. Arias, Guillermo Rueda-Delgado & Karl Matthias Wantzen VOLUME 2. EPHEMEROPTERA OF SOUTH AMERICA by Eduardo Domínguez, Carlos Molineri, Manuel L. Pescador, Michael D. Hubbard & Carolina Nieto Scientific Reader to the Editors: Michel Sartori

VOLUMEN 2. EPHEMEROPTERA DE AMÉRICA DEL SUR por Eduardo Domínguez, Carlos Molineri, Manuel L. Pescador, Michael D. Hubbard & Carolina Nieto Lector Científico a los Editores: Michel Sartori

Front cover: Nymph of Chiloporter eatoni (LESTAGE) (Ameletopsidae) and Amazon River near Manaus, Brazil (photos: E. Domínguez, W.J. Junk; design: Zheko Aleksiev & Elke Bustorf) Aquatic Biodiversity in Latin America Vol. 2 ISSN 1312-7276 First published 2006 ISBN-10: 954-642-259-2 ISBN-13: 978-954-642-259-0 The book should be cited as follows: Domínguez, E., Molineri, C., Pescador, M.L., Hubbard, M.D. & C. Nieto. 2006. Ephemeroptera of South America. In: Adis, J., Arias, J.R., Rueda-Delgado, G. & K.M. Wantzen (Eds.): Aquatic Biodiversity in Latin America (ABLA). Vol. 2. Pensoft, Sofia-Moscow, 646 pp. © PENSOFT Publishers All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form by any means, electronic, mechanical, photocopying, recording or otherwise, without the prior written permission of the copyright owner. Pensoft Publishers Geo Milev Str. 13a, Sofia 1111, Bulgaria [email protected] www.pensoft.net Printed in Bulgaria, June 2006

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To Bill and Jan Peters whose constant support and encouragement inspired us to write this book.

CyanMagentaYellowBlack Adis Vol.2 Page 6

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ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur

Undescribed male imago of Miroculis sp. nov. (Leptophlebiidae), from the Uruzú River, Parque Provincial Urugua-I, Misiones Province, Argentina; collected by E. Domínguez on 26-XI-2001. Digital drawing (Adobe-Photoshop) by Victoria Saxe (www.victoriasaxe.com) © 2006

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AQUATIC BIODIVERSITY IN LATIN AMERICA BIODIVERSIDAD ACUÁTICA

EN

AMÉRICA LATINA

Series Editors: Joachim Adis, Jorge R. Arias, Guillermo Rueda-Delgado & Karl Matthias Wantzen Biodiversity is a key word in science and global management schemes, however very few people are able to identify the species and their ecology that make up “biodiversity”. For many years, researchers and students from numerous countries have complained about the lack of “tools” to identify aquatic invertebrates from Latin America. Keys found in accepted entomological textbooks are mostly limited, superficial and rarely cover Neotropical biota in sufficient detail. On the other hand, specialized information on taxonomy or ecology is scattered throughout the literature in many single publications. An international team of editors have combined their efforts with Pensoft Publishers to launch a new major series on the Aquatic Biodiversity in Latin America (ABLA). Their goal was to find experts who combine the current state of knowledge in taxonomy and ecology, in order to produce a concise and affordable handbook for each group. About 15 separate monographs, written by reference scientists from various countries will offer a new, unrivalled view of the aquatic fauna of South America. Information on the ecology and status of the taxa (written in English) is combined with illustrated identification keys to families and genera, in both English and Spanish. The series is aimed at zoologists, ecologists, hydrobiologists, biogeographers, conservationists and students interested in aquatic biodiversity. The series will be an essential tool for any biological library. Volume 1: Amazon Fish Parasites (Second edition) by Vernon E. Thatcher. Pensoft Publishers, Sofia-Moscow, ISBN-10: 954-642-258-4, ISBN-13: 978-954-642258-3, 165x240 mm, 508 pp., including 194 plates of figures in line drawings and photos (15 plates in color); publication date: March 2006. Volume 2: Ephemeroptera of South America by Eduardo Domínguez, Carlos Molineri, Manuel L. Pescador, Michael D. Hubbard & Carolina Nieto. Pensoft Publishers, Sofia-Moscow, ISBN-10: 954-642-259-2, ISBN-13: 978-954-642259-0, 165x240 mm, 646 pp., including 234 plates of figures in line drawings and photos (16 plates in color); publication date: May 2006. Mayflies represent a large part of the freshwater benthic biomass and are of central importance for the functioning of aquatic ecosystems. The nymphs of many taxa are sensitive to changes in habitat and water quality and most bioindicator systems include their

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ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur

occurrence. This book contains the currently known reference data on each taxon, considering its status, characters for identification, biology and distribution. This book covers all known 14 families, 103 genera and 466 species of the insect order Ephemeroptera from continental South America. It includes updated keys (in English and Spanish) to separate families, genera and species (when possible). The book is addressed to specialists in systematics, ecologists, limnologists and biologists in general.

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ADDRESSES Authors: PROF. DR. EDUARDO DOMÍNGUEZ CONICET-INSUE Facultad de Ciencias Naturales Universidad Nacional de Tucumán Miguel Lillo 205 4.000 Tucumán Argentina e-mail: [email protected] DR. CARLOS MOLINERI CONICET-INSUE Facultad de Ciencias Naturales Universidad Nacional de Tucumán Miguel Lillo 205 4.000 Tucumán Argentina e-mail: [email protected] PROF. DR. MANUEL L. PESCADOR Laboratory of Aquatic Entomology Florida A&M University Tallahassee, Florida 32307-4100, USA e-mail:[email protected] PROF. DR. MICHAEL D. HUBBARD Laboratory of Aquatic Entomology Florida A&M University Tallahassee, Florida, 32307-4100 USA e-mail: [email protected].

DR. CAROLINA NIETO CONICET-INSUE Facultad de Ciencias Naturales Universidad Nacional de Tucumán Miguel Lillo 205 4.000 Tucumán Argentina e-mail: [email protected] Series Editors: PROF. DR. JOACHIM ADIS Max-Planck-Institute for Limnology Tropical Ecology Working Group Postfach 165 24302 Plön, Germany e-mail: [email protected] DR. JORGE R. ARIAS 5870 Colfax Avenue Alexandria, Virginia 22311, USA e-mail: [email protected] M.Sc. GUILLERMO RUEDA-DELGADO Grupo de Investigación en Cuencas y Humedales Tropicales GICHT UNIMAGADALENA Universidad de Bogotá Jorge Tadeo Lozano Laboratorio de Limnología Carrera 4 No. 22-61 Bogotá, D.C. Colombia e-mail: [email protected]

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ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur

DR. KARL MATTHIAS WANTZEN Institute of Limnology University of Konstanz Postfach M 659 78457 Konstanz, Germany e-mail: [email protected]

Scientific Reader to the Editors: DR. MICHEL SARTORI Musée cantonal de zoologie Palais de Rumine Place de la Riponne 6 CH-1014 Lausanne, Switzerland e-mail: [email protected]

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FOREWORD Ephemeroptera of South America by Eduardo Domínguez, Carlos Molineri, Manuel L. Pescador, Michael D. Hubbard & Carolina Nieto More than 100 works have been published on the systematics of South American Ephemeroptera (mayflies) in the last decade, including descriptions of 25% of the known species. Sixty-one species were described for the single family Baetidae, representing half of the presently known species. These statistics are indicative of the great effort recently produced to improve our knowledge of this fragment of the South American aquatic fauna. With about 14 families, 100 genera and more than 450 species, we begin to further understand and appreciate the diversity of the South America mayflies. Based on present knowledge and distribution, this group of insects is not only well diversified, but it also possesses a high degree of endemism. Two families, 65% of the genera and almost all species are endemic to South America. Only a few widespread species extend their ranges as far north as Central America or southern Mexico. The history of the first species described from South America is quite enigmatic and reflects, to some extent, the difficulties encountered by present ephemeropterists. Two centuries ago, WEBER described the first South American mayfly under the name of Ephemera astrostoma; it was subsequently transferred to the genus Palingenia. This was problematic, as the Palingeniidae do not occur in South America. Because the type specimen has been lost, no one knows what Ephemera astrostoma should be and the species is considered as nomen dubium. The first three valid species were described fifty years later by the Swiss entomologist PICTET in his famous book “Histoire naturelle générale et particulière des insectes névroptères. Famille des éphémérines”. Since then, over 300 publications have been devoted to the taxonomy and systematics of the South American mayflies. While some taxa were well described and illustrated, others have rather brief and incomplete descriptions, rendering many identifications only tentative. This is the case in the approximately 50 papers dedicated by NAVÁS to South American mayflies; the most symptomatic, or should I say absurd, case is a species of Callibaetis (Baetidae) described under eight different names by the same author. To the dismay of most taxonomists, no less than 250 synonymies or changes in generic attribution have been necessary to obtain the present comprehensive account of South American mayflies. Further revisions in some problematic genera (i.e., the 40 species of Campsurus in the family Polymitarcyidae or the provisional attribution of some species of Baetidae only known at imaginal stage) will likely increase the number of changes. If it is not enough to discourage the most motivated workers, only 10% of the species are known at both imaginal and nymphal stages, and stages are sometimes described under different specific or even generic names. Nevertheless, identification of species or genera remains of primordial importance for any kind of ecological study. Aside from the difficulties mentioned above, access to the

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ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur

relevant literature can be difficult and time-consuming, making more evident and urgent the necessity of taxonomic keys and a comprehensive summary of the present knowledge. Writing an encyclopedic book on the Ephemeroptera of South America in these conditions represents more than a great challenge that the small size of the order may not have necessarily predicted. A first attempt was made by HECKMAN (2002) to gather the present knowledge in an encyclopedia of South American mayflies. Unfortunately, the result is more similar to a general key that depends on the quality of the original descriptions. It is obvious that only a group of specialists with a wide background and closely cooperating can overcome such a monumental task. Five scientists from two institutions have come together to write this second volume of the Aquatic Biodiversity of Latin America devoted to Ephemeroptera: Eduardo DOMÍNGUEZ, Carlos MOLINERI and Carolina NIETO from the National University of Tucumán (UNT), Argentina as well as Michael HUBBARD and Manuel PESCADOR from Florida A&M University (FAMU), Tallahassee, Florida, USA. The UNT team has played, and continues to play, a major role in increasing our knowledge of the systematics of the South American mayflies, having published more than 50 quality taxonomic papers, having edited an important synthesis (Trends in Research in Ephemeroptera and Plecoptera (DOMÍNGUEZ 2001), and having organized the IX International Conference on Ephemeroptera in Argentina. The UNT also constitutes one of the few places in the world where the study of the systematics is still recognized as relevant and new taxonomists are continuously being trained. The FAMU team is also led by the same spirit of work, collaboration and exchange of knowledge. It is impossible to evoke the FAMU without reference to the late Bill PETERS, the first Professor and head of the Entomology Department at FAMU and the first chairman of the permanent committee on Ephemeroptera. Among others, Michael HUBBARD and Manuel PESCADOR benefited from his encouragement and guidance as graduate students and to some extent can be considered as his heirs. Michael has spent a lot of his energy in compiling the different faunas and checklists. Probably, the most evident result is the publication of the catalogue of the family and genus groups of the mayflies of the world (HUBBARD 1990) as well a first catalogue of the South American mayflies (HUBBARD 1982). Manuel summarized status of taxonomy of the South American mayflies and put in advance the problems and priorities (PESCADOR et al. 2001). The scientists of both teams constantly work together toward the diffusion and exchange of knowledge in mayflies. One of the first results of their collaboration was the compilation of the first comprehensive modern checklist of South American mayflies (Ephemeroptera Galactica: http://www.famu.org/mayfly). My real hope is that this book will be the tool guide for scientists who want (or need!) to work with the essential component of the aquatic ecosystem that are mayflies. It will provide all the necessary keys and information for identification of this complex group of insects. Certainly, it will be regarded as a masterpiece, contributing to our knowledge of the systematics, ecology and distribution of South American mayflies. DR. JEAN-LUC GATTOLLIAT Museum of Zoology Lausanne, Switzerland

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REFERENCES DOMÍNGUEZ, E. (2001): Trends in research in Ephemeroptera and Plecoptera. New York, Kluwer Academic/Plenum. HECKMAN, C.W. (2002): Encyclopedia of South American aquatic insects: Ephemeroptera. Dordrecht, Kluwer Academic. HUBBARD, M.D. (1982): Catálogo abréviado de Ephemeroptera da America do Sul. Papeis Avulsos de Zoologia 34:257-282. HUBBARD, M.D. (1990): Mayflies of the world. A catalog of the Family and Genus Group Taxa (Insecta: Ephemeroptera). - Flora and Fauna Handbook 8: 1-119. Sandhill Crane Press, Gainesville. PESCADOR, M.L., HUBBARD, M.D. & M.D.C. ZUÑIGA (2001): The status of the taxonomy of the mayfly (Ephemeroptera) fauna of South America. - In: DOMÍNGUEZ, E. (ed.): Trends in research in Ephemeroptera and Plecoptera: 37-42. New York, Kluwer Academic/Plenum.

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ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur

PREFACE Mayflies are one of the dominant and diverse groups of aquatic insects in South America. Our knowledge of the systematics of the fauna in the region, however, is still limited and fragmentary. Inadequate taxonomic keys to identify the various taxa remain a major problem. Moreover, the literature dealing with the taxonomy of the group are scattered in various publications, and it is a time-consuming exercise to do a literature search. This trend, however, has slowly changed with a growing number of published taxonomic papers on the group in the last decade or so. This book represents the first authoritative attempt to comprehensively consolidate the available taxonomic information on the mayfly fauna of South America. It includes taxonomic keys to families, genera, and species (when possible), as well as taxonomic accounts, diagnosis, geographic distribution and biology. Mayflies are excellent indicators of water quality and to fully appreciate the utility of the order as a biomonitoring tool requires a good taxonomic knowledge of the group. We hope that this book will immensely foster the much needed incentives for aquatic biologists to conduct more in-depth studies of the mayfly fauna in the region. This book was initiated in 1986-87, when the senior author was a postdoctoral research fellow in the Laboratory of Aquatic Entomology at Florida A&M University, Tallahassee, Florida. Since then, taxonomic knowledge of the group has enormously improved, and herein, is a comprehensive documentation of information on the fauna in the region. The taxonomic keys presented in this book were used and tested by students and aquatic biologists who enrolled in courses, and participated in workshops that were offered at universities in various countries in the region such as Argentina, Bolivia, Chile, Colombia, Ecuador, and Venezuela. Through these efforts, constructive comments by the students have immensely improved the contents and utility of the taxonomic keys. We sincerely thank our colleagues and friends, G. Cuezzo, S. Elliott, R. W. Flowers, J. L. Gattolliat, P. Malzacher, M. Mercado, W. L. Peters (deceased), B. Richard, G. Wibmer and M. del C. Zuñiga for their encouragement and invaluable contributions to the successful completion of the book. Janice G. Peters made a special contribution correcting important portions of several chapters. The Scientific Reader M. Sartori and the Series Editors: J. Adis, J. R. Arias, G. Rueda-Delgado and K.M. Wantzen went patiently through the different versions, making valuable suggestions, improving the content and the final shape of the book. Alvaro, Daniela and Rodrigo Domínguez helped in the last editorial corrections. Finally, our recognition to our families, for their loving support and understanding during the long and arduous process of writing this book. THE AUTHORS Tucumán, September 2005

ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America

Table of Contents 1I 2 M Nymph

NTRODUCTION

17

ORPHOLOGY

18 18

Adult Eggs

20

IOLOGY

23

3B 4W 5H 6D 7 GL

22

ORKING WITH

IGHER

SPECIMENS

CLASSIFICATION

25 27

ISTRIBUTION

28

FAMILIES, SUBFAMILIES, ENERA AND SPECIES

29

IST OF

8S

YSTEMATICS

Keys to South American families Claves para las familias sudamericanas Family Ameletopsidae Family Baetidae Family Caenidae Family Coloburiscidae Family Coryphoridae

43 43 48 52 57 194 224 227

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ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur

Family Family Family Family Family Family Family Family Family

9C 10 GS

OLOR

Ephemeridae Euthyplociidae Leptohyphidae Leptophlebiidae Melanemerellidae Nesameletidae Oligoneuriidae Oniscigastridae Polymitarcyidae

PLATES

230 234 244 331 533 536 538 555 557 589

UIDE TO CHANGES IN

CIENTIFIC

11 R 12 S 13 A

NAMES

607

EFERENCES

611

UBJECT INDEX

635

DDENDUM

645

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1 INTRODUCTION The Order Ephemeroptera is a small group of winged insects, with approximately 375 genera and 3,000 species grouped in 37 families worldwide. They have two-winged stages, the subimago and imago, which are unique among the insects. Mayflies are the most primitive of the extant winged insects, with fossil records dating back to the Carboniferous (more than 280 m.y.b.p.) and Permian (230 m.y.b.p.). The non-feeding adults are short lived and depend largely on nymphal reserves; the main functions of the adult are mating, oviposition and dispersal. Most of the mayfly’s life is spent in the aquatic immature stages (see discussion below). Mayflies are a diverse and vital biotic component of freshwater ecosystems. They are found in nearly every type of aquatic habitat and a few have ventured into brackish environments. The nymphs are members of various functional feeding groups (MERRITT & CUMMINS, 1996), such as collectors/gatherers, filterers, scrapers/grazers, shredders and predators. Some species have the ability to shift from one functional feeding group to another as the nymphs mature, thus increasing the flexibility of utilizing a variety of food resources for growth and development (BRITTAIN, 1982). The nymphs play a significant role in nutrients cycling and are involved in the processing of enormous amount of organic matter in streams, rivers and lakes. They serve as vital food resource for fish, amphibians, and other predaceous aquatic organisms including insects. Many aerial predators (insects, birds, bats, etc.) consume great numbers of mayfly adults during the emergence, swarm or oviposition periods (GRANT, 2001). Moreover, mass emergences of mayflies transfer large amount of phosphates and nitrates from aquatic to terrestrial environments thus helping in removing and cleansing pollutants in freshwater systems (EDMUNDS et al., 1976). Their ubiquity and abundance in freshwater ecosystems makes them excellent indicators of water quality. The use of mayflies as biological indicators of water quality has been well documented (RESH & JACKSON, 1993; BARBOUR et al., 1996; MORSE et al., 1997).

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2 MORPHOLOGY To identify the various mayfly taxa with certain degree of confidence and reliability, it is fundamentally important to have good understanding and knowledge of the external morphology of both the nymph and adults. NYMPH (Figs. 1A, B) Head: The head is heavily sclerotized, variable in shape, and usually with a variety of armature, projections, and processes. The compound eyes are located either dorsally or near the posterolateral corners of the head. There are three simple eyes or ocelli, the median ocellus usually smaller than the lateral ones. The antennae vary in length, and are attached anterior to or between the compound eyes. The fully functional mouthparts are directed either ventrally or anteriorly, depending on the orientation of the head, which is either hypognathous (downward) or prognathous (forward). In the hypognathous head the mouthparts are ventrally directed, compared to anteriorly in the prognathous head. The mouthparts consist of the flap-like labrum, a pair of heavily sclerotized mandibles, a pair of maxillae, membranous tongue-like hypopharynx, and the labium. Differences in shape, structures, and setations of the mouthparts between taxa are mostly functionally driven, and are excellent sources of taxonomic characters. The basic structures of nymphal mouthparts are illustrated in Fig. 1B. Thorax: The thorax has three distinct segments: the pro-, meso-, and the metathorax. Like the adult, the nymphal meso- and metathorax are fused. The thoracic nota are heavily sclerotized, notably the mesothorax. Wingpads are located on the meso- and metathorax (often referred to as pterothorax or wing bearing segments), although wingpads may be absent in the metathorax of some species. The thoracic segments each have a pair of legs: prothoracic legs (fore legs), mesothoracic (middle) legs and metathoracic (hind) legs. The segments of the legs include the coxa, trochanter, femur, tibia and tarsus (Fig. 1A). Most segments usually have thin and/or thick setae, and fringes of hairs, and each tarsus has a single claw, which is either denticulate or non-denticulate. Sometimes the thick setae are referred as “spines”. Like the mouthparts, structural differences and morphological modifications of the legs are functionally related such as for clinging, burrowing, filtering food, and gill protection. Abdomen: The abdomen consists of ten distinct segments, some may be concealed beneath a thoracic shield or carapace (e.g., Baetiscidae, Prosopistomatidae), or in mature nymphs of families such as Leptohyphidae and Caenidae, just the first abdominal segment may be

ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America

Hind Leg

A

Fore Leg

Middle Leg

Terminal Filament

Abdomen

Mesonotum Fore Wing Pads

Cerci

Ocelli

Pronotum Head

Labrum

Gills Femur Tibia

Tarsus

Antenna

Claw

Labrum

B Incisor

Incisor

Prostheca Molar

Prostheca Molar Clypeus Lingua

Superlingua

Left Mandible

Right Mandible

Hypopharynx

Palpi

Galea-lacinia Glossa

Paraglossa Palpi

Stipes

Left Maxilla

Right Maxilla Labium

Plate 1. Figs. 1A-B, nymph. 1A, habitus, dorsal view; 1B, mouth parts.

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covered by the mesothoracic wingpads. The abdominal terga may possess spines and/or tubercles or both, and some segments may have posterolateral spines. The abdominal sterna are relatively smooth, and patterns of pigmentation of the terga and sterna are sometimes species specific. All mayfly nymphs have gills except for the mayfly genus Murphyella (Coloburiscidae, Figs. 79A, 227C). The gills vary in shape and structure, and provide an enormous source of taxonomic characters. Gills are mostly found on the abdomen, except for a few taxa which have the gills at the base of the coxae (e.g., some species of Baetodes, Fig. 23E, and Camelobaetidius, Figs. 32H, 33D) or at the base of the maxillae (e.g., Lachlania, Oligoneuriidae). Most mayflies have the gills dorsally or laterally attached to the abdomen, others however, have the gills attached ventrally (e.g., some Baetodes). In some species, the first pair of abdominal gills is ventral and the rest of the gills are dorsal or lateral as in some Oligoneuriidae (Figs. 204J, 207A). Most nymphs have gills on abdominal segments I-VII, although some taxa have the gills on segments I-IV (Siphlonella), I-VI (Massartella), II-V (Leptohyphodes, Coryphorus), II-VI (Melanemerella, Fig. 201A), III-VII or IV-VII (Palearctic species). Most taxa have three caudal filaments consisting of a pair of cerci and a median (terminal) filament (Figs. 219A, 227A-B, 228A, 230A). Some species have the median filament greatly reduced or absent (Figs. 25B, 227C). ADULT (Figs. 2A, 224A-B, 225A-B, 226A-B, 228C, 229B, 230B, 231B, 232A-B) In this book, we will use “adults” when we refer to subimaginal and/or imaginal stages indistinctly. When referring specifically to one of these stages, we will call them “imago” or “subimago”. Head: The males mostly have distinctly larger compound eyes, and are closer together than those of the females. In some groups (e.g., Caenidae, Leptohyphidae, Polymitarcyidae) both sexes have similar compound eyes, smaller and widely separated. In the Baetidae and some Leptophlebiidae male compound eyes are turbinate with the upper facets enlarged, radiantly colored, and stalked (Figs. 40A, 165L-M, 218A-C). Thorax: The thorax has three distinct segments, the prothorax with the fore legs, the mesothorax with the middle pair of legs and the fore wings, and the metathorax with the hind legs and hind wings (which may be absent in some species). In most mayflies the adult fore legs are sexually dimorphic and distinctly longer in males than in females. In some species (Polymitarcyidae), however, the middle and hind legs of the female are greatly reduced, and in Campsurus, these legs are broken off when the species molts to the adult. Most mayflies have two pairs of wings; the fore wings and the much smaller hind wings. The hind wings are greatly reduced or completely lost in one or both sexes of the family Caenidae, Baetidae, Leptohyphidae and some species in the Leptophlebiidae. The basic wing venation of the fore and hind wings of mayfly adults is shown in Figs. 2B-C. The abbreviations of the veins used in the text are as follows: C= Costa Sc= Subcosta

ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America

R1 R2

Fore Wing C Sc

R3 R4+5 MA 1

Bullae

MA2

Costal Projection Eye Ocelli

MA MP

CuA

Antenna 2

Frons Coxa Trochanter

Tibia Tarsi

1

Fore Leg

Middle Leg

C Rs CuP

A

B

Cu

A

Sc

R1 R3

MP1 IMP

Thorax

Femur

3

MP2

10 g in 9 dW Hin 8 7 4 5 6

Caudal Filaments Genitalia

Abdomen Hind Leg

A

R2

M M M A1 R4+5 P M 1 A 2 P 2

C

21

C Sc R Rs M A C P C MA uP uA

Forceps

Forceps Socket

Penes Subgenital Plate

D

Plate 2. Figs. 2A-D, adult. 2A, lateral view; 2B, fore wing; 2C, hind wing; 2D, genitalia, ventral view.

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R1, R2, R3, R4+5= Radius 1, Radius 2, etc. In general, R2 to R5 are called Radial sector. MA= Medius anterior MA1, MA2= Medius anterior 1, Medius anterior 2 IMA= Intercalary Medius anterior MP= Medius posterior MP1, MP2= Medius posterior 1, Medius posterior 2 IMP= Intercalary Medius posterior CuA= Cubitus anterior ICu1, ICu2= Cubital Intercalary 1, Cubital Intercalary 2 CuP= Cubitus posterior A1, A2= Anal 1, Anal 2, etc. Normally only A1 used, and called simply “A”. Abdomen: The abdomen is composed of ten segments. A pronounced lateral longitudinal membrane or conjunctiva (or pleural folds) on both sides of the abdomen delineates the tergum and sternum. The posterior portion of the sternum IX of the female and male is referred to as the subanal plate, or subgenital plate; in males, the distal portion is called the styliger plate. The basic structures of the male genitalia are shown in Fig. 2D. A pair of slender and usually segmented forceps or claspers is attached to the posterior margin of the subgenital plate. Dorsal to the subgenital plate are the paired penes, which are either separated or partially to entirely fused. The shape and processes of the penes provide excellent source of taxonomic characters. The caudal filaments are attached to the abdominal tergum X. Many species have two caudal filaments, the cerci; others have three with a median filament either well developed, greatly reduced or atrophied. EGG (Figs. 221A-E, 222A-F, 223A-D) The abdomen and most of the thorax of the female adults are usually filled with eggs, which are generally ovoid to rectangular, and have different sculptures (Figs. 221A-E, 222A-F, 223A-D) and sizes ranging from 150 to 200 µm (in largest species 250-300 µm). Due to the short life span of the adults, the eggs are already developed in the nymph before its emergence, and can be used to associate the last nymphal instars with the female adults. The eggs have different attachment structures to enhance secure deposition to the substratum. Some groups have one (Figs. 221B, D-E) or two polar caps (Fig. 221C), composed of several adhesive filaments that separate when deposited in the water. Other groups have from few to many Ktc (Knob-terminated coiled threads) that are distributed in part or over the whole surface (Figs. 222A-E). When these Ktc come in contact with the substratum, they are triggered to attach the eggs (Figs. 222F, 223A-D). In other cases, the whole surface is adhesive, without any individual attachment structure as those mentioned above (Fig. 221A). The micropyle (Fig. 222C) is the structure that allows sperm to enter the egg. Sometimes the micropyle has accessory structures such as a sperm guide and/or a micropylar canal, collectively constituting the micropylar area of the egg chorion (Figs. 221C, E).

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3 BIOLOGY Mating is a visually mediated process in mayflies. Thus, the enlarged male eyes provide better acuity and sensitivity to locate the females particularly among non-swarming species and those with asynchronous emergence. The greatly elongated fore legs of the male adults are used for grasping and holding the females while in copula. Unique among mayflies is the presence of two-winged stages, the subimago and the imago (although females of some Paleartic groups such as Palingeniidae remain in the subimaginal stage). The subimagos are easily recognized from the imagos by the generally opaque or dull color (Figs. 220E, 224A-B, 226A, 231B), ciliated wings and caudal filaments. Possible reasons for the retention of the subimaginal stage in mayflies include the facilitation of aquatic-aerial shift by the hydrophobic subimaginal skin, or the need for two molts to complete the elongation of various body appendages such as the fore legs, caudal filaments and external genitalia. Swarming often referred to as “nuptial flight” in mayflies is basically a male initiated activity to attract the females. Swarming facilitates mating, and apparently serves, in addition to attracting the opposite sex, as a mechanism to protect the population from predation by minimizing its effect through saturation or predator satiation. The swarm occurs in a variety of places such as above the water surface, shore lines or some markers like rock boulders, tree stumps, clearings between trees and shrubs alongside a river or streams, etc. The time of day of swarming is species-specific, and is closely connected to light intensity. Swarming mostly takes place at dusk or early in the morning, although some species swarm between mid-morning and noon. Facultative parthenogenesis has been reported for a few mayfly species. Most mayflies lay 400 to 3000 eggs, although extremes quantities ranging from species with 100 eggs to others with as many as 12000 eggs have been recorded. Intraspecific variations on fecundity is a common occurrence among bivoltine species (two summer emergence periods) whereby early emerging females which, on the average, are larger in body size and mostly have more eggs than the later emerging populations. Mayflies exhibit different ways of ovipositing. Most species lay the eggs by flying over the surface of the water and intermittently dip the tip of the abdomen in the water, releasing a few eggs at a time until the oviposition process is completed; a few species simply explode or release the eggs all at once. Some species cling onto the surface of wood snags, tree stumps or stones above the water surface and dip the tip of the abdomen in the water to lay the eggs. A few species have the females swim underwater and directly deposit the eggs under stones or in rock crevices.

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Incubation period in mayflies varies from a few days to almost a year. A few species of Baetidae are ovoviviparous (e.g., Callibaetis radiatus, C. sellacki, C. willineri). Egg development in mayflies is very much a temperature regulated process. Temperature, in combination with the modifying influences of other environmental factors (e.g., food quality, current velocity) very much regulates nymphal growth and development in mayflies (BRITTAIN, 1982). Likewise, the same environmental factors had been reported to control the number of postembryonic molts, which vary from between 10 and 50. The number of molts in mayflies is higher than in most other insect orders. Three life cycle patterns or voltinisms are well-recognized in mayflies: univoltinism, multivoltinism and semivoltinism (BRITTAIN, 1982). Univoltine species have one generation a year and occur commonly in temperate areas. Multivoltine species usually have two to three generations in a year in temperate areas, and very likely more generations in the tropics. A relatively uncommon life cycle in mayflies, the semivoltine species, requires more than a year and up to three years to complete a generation. The shedding of the nymphal exuviae prior to emergence is a critical period for mayflies as they are exposed to and become vulnerable to both aquatic and terrestrial predators. In most mayflies, notably the river dwelling species, shedding of nymphal skin takes place on the surface of the water. In some species such as Siphlonella ventilans (Oniscigastridae), the nymphs crawl out of the water and grasp on to solid objects such as tree stumps, snags or rocks (Figs. 219B, 220A-E). The duration of subimaginal stage varies from a few minutes to a few hours depending on a combination of environmental conditions such as temperature and humidity, and species traits. The ciliated wings and caudal filaments of the subimagos, especially the large river dwelling mayflies that remain in this stage, apparently protect them from drowning while in copula, and oviposition (RUFFIEUX et al., 1998).

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4 WORKING WITH SPECIMENS Collecting: The various techniques of effectively collecting, and rearing mayflies are comprehensively discussed by EDMUNDS et al. (1976) and for aquatic insects in general by MERRITT & CUMMINS (1996). We strongly recommend to anyone seriously interested in sampling aquatic insects to refer to these two references because the different methods to successfully collect aquatic insects both qualitatively and quantitatively, from various types of aquatic habitats, are extensively documented. Thus, we felt that it would be expedient to exclude collecting techniques in this book, instead, focus on the rearing methods to associate the nymphal and adult stages of mayflies, highlighting the techniques that we have successfully used over the years. Rearing: For many mayfly taxa, identification of nymphs is possible only to the genus level. By rearing the nymphs to the adult stage, the nymph can be associated with the adult and then both the shed nymphal exuviae and the adult can be used for identification. Association of the nymph and adult mayflies is a reasonably easy process. The two methods that we have commonly used with reasonable success include a simple aquarium set-up and on site flotation method. In both cases, mayfly nymphs collected in the field are placed in plastic drink cups, which have side sections cut away and covered with plastic mesh glued to the sides of the cup. This allows for fresh water to flow freely through the rearing cup. Each rearing cup is provisioned with stream substrates, and a small stick mainly for food and foothold for the nymphs, respectively. For the on site rearing, the cups are placed in holes cut out of a piece of Styrofoam that is anchored to the riverbank. In this way, the cups will not be carried away and the water level inside the cup will remain constant even if the river changes it level during the process. The cups are covered with a lid or a piece of cloth to keep the emerging adult. The cups must be checked after dark, and the emerged subimagos transferred to dry rearing cages. For the aquarium technique, is necessary to transport the living nymphs back to the laboratory. The cups with the nymph are placed in a Styrofoam cooler partially filled with stream water. To hold cups in place and keep above water level, they are placed in holes cut out of a piece of Styrofoam, which fit snugly in the cooler. The water in the cooler is oxygenated with a batteryoperated aerator while transporting the nymphs. The rearing cups are covered with a cloth mesh to prevent the emerging adult from escaping and placed in an aerated aquarium filled with stream water. Water temperature is a critical factor in rearing success. As much as possible, the water temperature in the aquarium must be maintained close to the prevailing stream temperature where the nymphs were collected. In general, if the water

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in the aquarium is above 24ºC, nymphs will either cease development or die. Mature nymphs, notably individuals with black wing pads, are checked on a daily basis for emergence. In both techniques, the newly emerged subimago is allowed time to complete the process of molting to the imago before preserving the imago, its nymphal, and subimaginal exuviae in a vial of ethyl alcohol. Dissection: Nymphs coated with silt or minute debris should be cleaned by ultrasonic cleaner for 3-6 minutes for better and more accurate viewing of structures and color patterns. In addition, glycerin is sometimes used to clean silted nymphs. Jewelers’ forceps, microdissecting scissors, dissecting dishes, and dissecting needles work very well in preparing specimens and associated structures for examination. Dissected specimens and body parts not mounted on slides should be placed in microvials and separated from the rest of the specimens in the main container. Mounting of Specimens on Slides: It is often necessary to make slide mounts of various parts of specimens in order to accurately examine taxonomic characters. Temporary slide mounts made with glycerin or glycerin-jelly are often all that is necessary to see many characters. Sometimes it is preferable to make permanent slide mounts using CMC10 or Canada balsam. CMC10 is convenient in that it is water soluble and parts can be mounted directly from alcohol or water and if necessary remounted after soaking a slide in water to loosen the cover slip, which can then be removed to allow for repositioning of parts. Mounting media also provide some clearing of parts, which aid in viewing certain structures. Microscopes: Most of the external morphological characters that are involved in identifying mayflies can be seen under a dissecting microscope that is equipped with 4050X magnification and an illuminated base. Most taxonomists also employ fiber optic lighting for additional illumination from above or to the side of specimens. A compound microscope with 100-400X magnification is needed for viewing certain mayfly structures. Preservation and Storage: The external morphological structures of the nymphs that are used to identify mayflies preserve well as long as proper preservation procedures are followed. To maintain and preserve the integrity of the insect structures and tissues, newly collected benthic samples should be preserved in 85-95% ethyl alcohol and replaced with the same concentration within 24 hours if the samples are not processed or sorted immediately. Once the specimens are sorted, they should be stored in 80% ethyl alcohol and appropriately labeled. For molecular work, specimens should be collected and preserved in 95% alcohol.

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5 HIGHER CLASSIFICATION The higher classification of Ephemeroptera is currently in a state of flux and there is no consensus about the phylogenetic relationships of the group. For this reason, the families presented herein are listed in alphabetical order. At present, there are approximately 15 families, 100 genera and 450 species reported for South America. The single citation of Epeorus (Heptageniidae) by ROJAS et al. (1993) must be considered a mistake. Despite the rapidly increasing number of taxonomic papers and evident improvement in the knowledge of the systematics of mayflies in the region, the fauna of several areas remain uncollected; at best, little collection has been undertaken. The number of described species per country is evidently the result of collection efforts, rather than actual species richness. Most of these problems, as well as the status of the knowledge of the taxonomy of the mayfly fauna of South America have recently been reviewed by PESCADOR et al. (2001). As previously indicated, the South American Ephemeroptera literature is widely scattered and often difficult to access. There is a Web Site that includes a periodically updated catalog of the South American species (http://www.famu.org/mayfly/cats.html), and links to other regional catalogs and Pdf files of papers on Ephemeroptera. The Mayfly Newsletter annually provides a bibliography of recently published papers related to Ephemeroptera, and can be obtained free of charge from the editor: Peter Grant at South Western Oklahoma State University, Weatherford, Oklahoma, 73096-3098, USA, [email protected].

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6 DISTRIBUTION South America has a unique biota composed of numerous extant and extinct endemic lineages of plants and animals. In Ephemeroptera this situation is also true, counting with a high number of endemism at different taxonomic levels: two families (14%), 67 genera (65%) and 434 species (97%). Some families show a high generic and specific diversity in South America, reaching Central and North America due to one or few widely distributed genera or species (e.g., Leptohyphidae, Polymitarcyidae, Euthyplociidae, Oligoneuriidae). Only two families widely distributed in the Northern Hemisphere and with some representatives in Central America, do not reach South America: Heptageniidae and Isonychiidae. The other world-wide distributed families such as Caenidae, Baetidae and Leptophlebiidae show a high generic and specific endemicity in South America. Two distinct biogeographic areas are recognized in South America (CABRERA & WILLINK, 1973; MORRONE, 2001): the Andean-Patagonic (approximately from 40ºS southward) and the Neotropical (approximately from 40ºS northward). The Andean-Patagonic area includes species that are restricted to the mountain cool, oxygenated water bodies from Central and Southern Argentina and Chile. Their closest relatives are located mainly in New Zealand and Australia (EDMUNDS, 1975). Representatives of this group are the families Ameletopsidae, Coloburiscidae, Nesameletidae, and Oniscigastridae. The Neotropical area includes species that are widely distributed in subtropical and tropical South America. The relatives of these groups occur mainly in Africa, Madagascar, Central and North America. Representatives of this group include members of the families Caenidae, Euthyplociidae, Leptohyphidae, Oligoneuriidae, Polymitarcyidae, among others. In the case of Leptophlebiidae, there are representative lineages in both biogeographical regions, and interestingly, the phyletic lineages show the same biogeographic pattern mentioned above. In Baetidae, a family with a distribution similar to Leptophlebiidae, the geographic patterns can not be ascertained, due to lack of knowledge of the phylogenetic relationships of the genera. Many subdivisions were proposed to describe the geographic patterns of the biota in both areas (the Andean-Patagonic and the Neotropical), CABRERA & WILLINK (1973) and MORRONE (2001) treat as many as 40 biogeographic provinces. Distributional records of some mayfly taxa seem to fit well in some of these areas, but much remains to be done on the biogeography of Neotropical mayflies. Focus on distributional data needs to be improved, especially in the subtropical and tropical components.

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7 LIST OF FAMILIES, SUBFAMILIES, GENERA AND SPECIES FAMILY AMELETOPSIDAE SUBFAMILY AMELETOPSINAE • Genus Chaquihua • Chaquihua bullocki (NAVÁS, 1930) • Chaquihua penai DEMOULIN, 1955 SUBFAMILY CHILOPORTERINAE • Genus Chiloporter • Chiloporter eatoni (LESTAGE, 1931) FAMILY BAETIDAE • Genus Adebrotus • Adebrotus amazonicus LUGO-ORTIZ & MCCAFFERTY, 1995 • Genus Americabaetis • Americabaetis alphus LUGO-ORTIZ & MCCAFFERTY, 1996 • Americabaetis bridarolli (NAVÁS, 1933) • Americabaetis jorgenseni (ESBEN-PETERSEN, 1909) • Americabaetis labiosus LUGO-ORTIZ & MCCAFFERTY, 1996 • Americabaetis longetron LUGO-ORTIZ & MCCAFFERTY, 1996 • Americabaetis maxifolium LUGO-ORTIZ & MCCAFFERTY, 1996 • Americabaetis oldendorffi (WEYENBERGH, 1883) • Americabaetis peterseni (HUBBARD, 1974) • Americabaetis robacki (LUGO-ORTIZ & MCCAFFERTY, 1994) • Americabaetis titthion LUGO-ORTIZ & MCCAFFERTY, 1996 • Americabaetis weiseri (NAVÁS, 1926) • Genus Andesiops • Andesiops angolinus (NAVÁS, 1933) • Andesiops ardua (LUGO-ORTIZ & MCCAFFERTY, 1999) • Andesiops peruvianus (ULMER, 1920) • Andesiops torrens (LUGO-ORTIZ & MCCAFFERTY, 1999) • Genus Apobaetis • Apobaetis fiuzai SALLES & LUGO-ORTIZ, 2002 • Apobaetis signifer LUGO-ORTIZ & MCCAFFERTY, 1997

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• Genus Aturbina • Aturbina beatrixae GILLIES, 2001 • Aturbina georgei LUGO-ORTIZ & MCCAFFERTY, 1996 • Genus Baetodes • Baetodes andamagensis MAYO, 1972 • Baetodes arawak (TRAVER, 1943) • Baetodes chilloni MAYO, 1972 • Baetodes cochunaensis NIETO, 2004 • Baetodes copiosus NIETO, 2004 • Baetodes gibbus NIETO, 2004 • Baetodes huaico NIETO, 2004 • Baetodes itatiayanus DEMOULIN, 1955 • Baetodes levis MAYO, 1968 • Baetodes pehuenche NIETO, 2004 • Baetodes peniculus MAYO, 1973 • Baetodes proiectus MAYO, 1973 • Baetodes pseudogibbus NIETO, 2004 • Baetodes rutilus NIETO, 2004 • Baetodes sancticatarinae MAYO, 1972 • Baetodes serratus NEEDHAM & MURPHY, 1924 • Baetodes solus MAYO, 1972 • Baetodes spinae MAYO, 1968 • Baetodes spinifer TRAVER, 1943 • Baetodes traverae MAYO, 1972 • Baetodes uruguai NIETO, 2004 • Baetodes yuracare NIETO, 2004 • Genus Bernerius • Bernerius incus WALTZ & MCCAFFERTY, 1987 • Genus Callibaetis • Callibaetis camposi NAVÁS, 1930 • Callibaetis dominguezi GILLIES, 1990 • Callibaetis fasciatus (PICTET, 1843) • Callibaetis gonzalezi (NAVÁS, 1934) • Callibaetis gregarius NAVÁS, 1930 • Callibaetis guttatus NAVÁS, 1915 • Callibaetis jocosus NAVÁS, 1912 • Callibaetis nigrivenosus BANKS, 1918 • Callibaetis pollens NEEDHAM & MURPHY, 1924 • Callibaetis radiatus NAVÁS, 1920 • Callibaetis sellacki (WEYENBERGH, 1883) • Callibaetis viviparus NEEDHAM & MURPHY, 1924 • Callibaetis willineri NAVÁS, 1932 • Callibaetis zonalis NAVÁS, 1915

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• Genus Camelobaetidius • Camelobaetidius alcyoneus (TRAVER, 1943) • Camelobaetidius anubis (TRAVER & EDMUNDS, 1968) • Camelobaetidius apis NIETO, 2003 • Camelobaetidius billi DOMINIQUE, THOMAS, ORTH & DAUTA, 2000 • Camelobaetidius cayumba (TRAVER & EDMUNDS, 1968) • Camelobaetidius coveloae (TRAVER, 1971) • Camelobaetidius dryops (NEEDHAM & MURPHY, 1924) • Camelobaetidius edmundsi DOMINIQUE, MATHURIAU & THOMAS, 2001 • Camelobaetidius huarpe NIETO, 2003 • Camelobaetidius ipaye NIETO, 2003 • Camelobaetidius janae DOMINIQUE, THOMAS, ORTH & DAUTA, 2000 • Camelobaetidius leentvaari DEMOULIN, 1966 • Camelobaetidius mantis TRAVER & EDMUNDS, 1968 • Camelobaetidius mathuriae DOMINIQUE & THOMAS, 2001 • Camelobaetidius matilei THOMAS, PERU & HOREAU, 2001 • Camelobaetidius ortizi DOMINIQUE & THOMAS, 2001 • Camelobaetidius patricki DOMINIQUE, MATHURIAU & THOMAS, 2001 • Camelobaetidius penai (TRAVER & EDMUNDS, 1968) • Camelobaetidius phaedrus (TRAVER & EDMUNDS, 1968) • Camelobaetidius serapis (TRAVER & EDMUNDS, 1968) • Camelobaetidius suapi NIETO, 2002 • Camelobaetidius tantillus (NEEDHAM & MURPHY, 1924) • Camelobaetidius tuberosus LUGO-ORTIZ & MCCAFFERTY, 1999 • Camelobaetidius yacutinga NIETO, 2003 • Genus Chane • Chane baure NIETO, 2003 • Genus Cloeodes • Cloeodes anduzei (TRAVER, 1943) • Cloeodes auwe SALLES & BATISTA, 2004 • Cloeodes aymara (TRAVER, 1971) • Cloeodes binocularis (NEEDHAM & MURPHY, 1924) • Cloeodes hydation MCCAFFERTY & LUGO-ORTIZ, 1995 • Cloeodes irvingi WALTZ & MCCAFFERTY, 1987 • Cloeodes jaragua SALLES & LUGO-ORTIZ, 2003 • Cloeodes nocturnus (NAVÁS, 1922) • Cloeodes penai (MORIHARA & EDMUNDS, 1980) • Cloeodes redactus WALTZ & MCCAFFERTY, 1987 • Cloeodes stelzneri (WEYENBERGH, 1883) • Cloeodes turbinops (NEEDHAM & MURPHY, 1924) • Cloeodes venezuelensis (TRAVER, 1943) • Genus Cryptonympha • Cryptonympha copiosa LUGO-ORTIZ & MCCAFFERTY, 1998 • Cryptonympha dasilvai SALLES & FRANCISCHETTI, 2004

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• Genus Fallceon • Fallceon murphyae (HUBBARD, 1974) • Fallceon yaro (TRAVER, 1971) • Genus Guajirolus • Guajirolus ektrapeloglossa FLOWERS, 1985 • Guajirolus queremba NIETO, 2003 • Genus Harpagobaetis • Harpagobaetis gulosus MOL, 1986 • Genus Iguaira • Iguaira poranga SALLES & LUGO-ORTIZ, 2003 • Genus Mayobaetis • Mayobaetis ellenae (MAYO, 1973) • Genus Moribaetis • Moribaetis aneto (TRAVER, 1971) • Moribaetis comes (NAVÁS, 1912) • Genus Nanomis • Nanomis galera LUGO-ORTIZ & MCCAFFERTY, 1999 • Genus Paracloeodes • Paracloeodes binodulus LUGO-ORTIZ & MCCAFFERTY, 1996 • Paracloeodes eurybranchus LUGO-ORTIZ & MCCAFFERTY, 1996 • Paracloeodes ibicui LUGO-ORTIZ & MCCAFFERTY, 1996 • Paracloeodes leptobranchus LUGO-ORTIZ & MCCAFFERTY, 1996 • Genus Prebaetodes • Prebaetodes sitesi LUGO-ORTIZ & MCCAFFERTY, 1996 • Genus Rivudiva • Rivudiva minantenna LUGO-ORTIZ & MCCAFFERTY, 1998 • Rivudiva trichobasis LUGO-ORTIZ & MCCAFFERTY, 1998 • Genus Spiritiops • Spiritiops silvudus LUGO-ORTIZ & MCCAFFERTY, 1998 • Genus Tomedontus • Tomedontus primus LUGO-ORTIZ & MCCAFFERTY, 1995 • Genus Tupiara • Tupiara ibirapitanga SALLES, LUGO-ORTIZ, DA-SILVA & FRANCISCHETTI, 2003 • Genus Varipes • Varipes cajuato NIETO, 2004 • Varipes helenae SALLES & BATISTA, 2004 • Varipes lasiobrachius LUGO-ORTIZ & MCCAFFERTY, 1998 • Varipes minutus NIETO, 2004 • Varipes singuil NIETO, 2004 • Genus Waltzoyphius • Waltzoyphius fasciatus LUGO-ORTIZ & MCCAFFERTY, 1995 • Waltzoyphius roberti THOMAS & PERU, 2002 • Genus Zelusia • Zelusia principalis LUGO-ORTIZ & MCCAFFERTY, 1998

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FAMILY CAENIDAE • Genus Brachycercus • Brachycercus sp. ORTH et al. (2000) • Genus Brasilocaenis • Brasilocaenis intermedia MALZACHER, 1986 • Brasilocaenis irmleri PUTHZ, 1975 • Brasilocaenis mendesi MALZACHER, 1998 • Brasilocaenis puthzi MALZACHER, 1986 • Brasilocaenis renata MALZACHER, 1986 • Brasilocaenis septentrionalis MALZACHER, 1990 • Genus Caenis • Caenis argentina NAVÁS, 1915 • Caenis burmeisteri MALZACHER, 1990 • Caenis candelata MALZACHER, 1986 • Caenis chamie ALBA-TERCEDOR & MOSQUERA, 1999 • Caenis cigana PEREIRA & DA-SILVA, 1990 • Caenis cuniana FROEHLICH, 1969 • Caenis dominguezi MALZACHER, 2001 • Caenis fittkaui MALZACHER, 1986 • Caenis gonseri MALZACHER, 2001 • Caenis grimi MALZACHER, 2001 • Caenis ludicra NAVÁS, 1920 • Caenis nemoralis NAVÁS, 1922 • Caenis pflugfelderi MALZACHER, 1990 • Caenis plaumanni MALZACHER, 2001 • Caenis pseudamica MALZACHER, 1990 • Caenis quatipuruica MALZACHER, 1986 • Caenis reissi MALZACHER, 1986 • Caenis sigillata MALZACHER, 1986 • Genus Cercobrachys • Cercobrachys colombianus SOLDÁN, 1986 • Cercobrachys peruanicus SOLDÁN, 1986 FAMILY COLOBURISCIDAE • Genus Murphyella • Murphyella needhami LESTAGE, 1930 FAMILY CORYPHORIDAE • Genus Coryphorus • Coryphorus aquilus PETERS, 1981 FAMILY EPHEMERIDAE • Genus Hexagenia • Hexagenia (Pseudeatonica) albivitta (WALKER, 1853) • Hexagenia (Pseudeatonica) callineura BANKS, 1914 • Hexagenia (Pseudeatonica) mexicana EATON, 1883

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FAMILY EUTHYPLOCIIDAE • Genus Campylocia • Campylocia anceps (EATON, 1883) • Campylocia bocainensis PEREIRA & DA-SILVA, 1990 • Campylocia dochmia BERNER & THEW, 1961 • Genus Euthyplocia • Euthyplocia haenschi ULMER, 1942 • Euthyplocia hecuba (HAGEN, 1861) • Genus Mesoplocia • Mesoplocia intermedia DEMOULIN, 1952 FAMILY LEPTOHYPHIDAE • Genus Allenhyphes • Allenhyphes? asperulus (ALLEN, 1967) • Allenhyphes flinti (ALLEN, 1973) • Allenhyphes? spinosus (ALLEN & ROBACK, 1969) • Genus Haplohyphes • Haplohyphes aquilonius LUGO-ORTIZ & MCCAFFERTY, 1995 • Haplohyphes baritu DOMÍNGUEZ, 1984 • Haplohyphes dominguezi MOLINERI, 1999 • Haplohyphes huallaga ALLEN, 1966 • Haplohyphes mithras (TRAVER, 1958) • Haplohyphes yanahuicsa MOLINERI, 2003 • Genus Leptohyphes • Leptohyphes carinus ALLEN, 1973 • Leptohyphes cornutus ALLEN, 1967 • Leptohyphes ecuador MAYO, 1968 • Leptohyphes eximius EATON, 1882 • Leptohyphes illiesi ALLEN, 1967 • Leptohyphes invictus ALLEN, 1973 • Leptohyphes jodiannae ALLEN, 1967 • Leptohyphes liniti WANG, SITES & MCCAFFERTTY, 1998 • Leptohyphes maculatus ALLEN, 1967 • Leptohyphes mollipes NEEDHAM & MURPHY, 1924 • Leptohyphes nigripunctum TRAVER, 1943 • Leptohyphes peterseni ULMER, 1920 • Leptohyphes petersi ALLEN, 1967 • Leptohyphes plaumanni ALLEN, 1967 • Leptohyphes populus ALLEN, 1973 • Leptohyphes setosus ALLEN, 1967 • Leptohyphes tacajalo MAYO, 1968 • Leptohyphes tuberculatus ALLEN, 1967 • Genus Leptohyphodes • Leptohyphodes inanis (PICTET, 1843)

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• Genus Lumahyphes • Lumahyphes guacra MOLINERI, 2004 • Lumahyphes pijcha MOLINERI, 2004 • Lumahyphes yagua MOLINERI & ZÚÑIGA, 2004 • Genus Traverhyphes • Traverhyphes (Byrsahyphes) nanus (ALLEN, 1967) • Traverhyphes (Byrsahyphes) yuqui MOLINERI, 2004 • Traverhyphes (Mocoihyphes) edmundsi (ALLEN, 1973) • Traverhyphes (Mocoihyphes) yuati MOLINERI, 2004 • Traverhyphes (Traverhyphes) chiquitano MOLINERI, 2004 • Traverhyphes (Traverhyphes) indicator (NEEDHAM & MURPHY, 1924) • Traverhyphes (Traverhyphes) pirai MOLINERI, 2001 • Genus Tricorythodes • Tricorythodes arequita TRAVER, 1959 • Tricorythodes australis (BANKS, 1913) • Tricorythodes barbus ALLEN, 1967 • Tricorythodes bullus ALLEN, 1967 • Tricorythodes cristatus ALLEN, 1967 • Tricorythodes hiemalis MOLINERI, 2001 • Tricorythodes lichyi TRAVER, 1943 • Tricorythodes mirca MOLINERI, 2002 • Tricorythodes nicholsae (WANG, SITES & MCCAFFERTTY, 1998) • Tricorythodes ocellus ALLEN & ROBACK, 1969 • Tricorythodes popayanicus DOMÍNGUEZ, 1982 • Tricorythodes quizeri MOLINERI, 2002 • Tricorythodes santarita TRAVER, 1959 • Tricorythodes yura MOLINERI, 2002 • Tricorythodes zunigae MOLINERI, 2002 • Genus Tricorythopsis • Tricorythopsis artigas TRAVER, 1958 • Tricorythopsis chiriguano MOLINERI, 2001 • Tricorythopsis gibbus (ALLEN, 1967) • Tricorythopsis minimus (ALLEN, 1973) • Tricorythopsis sigillatus MOLINERI, 1999 • Tricorythopsis undulatus (ALLEN, 1967) • Tricorythopsis volsellus MOLINERI, 1999 • Tricorythopsis yacutinga MOLINERI, 2001 • Genus Vacupernius • Vacupernius sp. EMMERICH, 2004 • Genus Yaurina • Yaurina mota MOLINERI, 2001 • Yaurina ralla (ALLEN, 1967) • Yaurina yapa MOLINERI, 2001 • Yaurina yuta MOLINERI, 2001

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FAMILY LEPTOPHLEBIIDAE • Genus Archethraulodes • Archethraulodes spatulus PESCADOR & PETERS, 1982 • Genus Askola • Askola froehlichi PETERS, 1969 • Genus Atopophlebia • Atopophlebia flowersi DOMÍNGUEZ & MOLINERI, 1996 • Atopophlebia fortunensis FLOWERS, 1980 • Atopophlebia obrienorum FLOWERS, 1987 • Atopophlebia yarinacocha FLOWERS, 1987 • Genus Bessierus • Bessierus doloris THOMAS & ORTH, 2000 • Genus Choroterpes • Choroterpes sp. • Genus Dactylophlebia • Dactylophlebia carnulenta PESCADOR & PETERS, 1980 • Genus Demoulinellus • Demoulinellus coloratus PESCADOR & PETERS, 1982 • Genus Ecuaphlebia • Ecuaphlebia rumignaui DOMÍNGUEZ, 1988 • Ecuaphlebia sp. A DOMÍNGUEZ, 1988 • Genus Farrodes • Farrodes caribbianus (TRAVER, 1943) • Farrodes carioca DOMÍNGUEZ, MOLINERI & PETERS, 1996 • Farrodes iguazuanus DOMÍNGUEZ & SAVAGE, 1987 • Farrodes longispinus DOMÍNGUEZ, MOLINERI & PETERS, 1996 • Farrodes maculatus (NEEDHAM & MURHPY, 1924) • Farrodes ochraceous DOMÍNGUEZ, MOLINERI & PETERS, 1996 • Farrodes pakitza DOMÍNGUEZ, MOLINERI & PETERS, 1996 • Farrodes roundsi (TRAVER, 1947) • Farrodes savagei DOMÍNGUEZ, 1999 • Farrodes tepui DOMÍNGUEZ, MOLINERI & PETERS, 1996 • Farrodes xingu DOMÍNGUEZ, MOLINERI & PETERS, 1996 • Farrodes yungaensis DOMÍNGUEZ & SAVAGE, 1987 • Genus Fittkaulus • Fittkaulus cuiabae SAVAGE, 1986 • Fittkaulus cururuensis SAVAGE, 1986 • Fittkaulus maculatus SAVAGE & PETERS, 1978 • Genus Gonserellus • Gonserellus atopus PESCADOR, 1997 • Genus Hagenulopsis • Hagenulopsis diptera ULMER, 1920 • Hagenulopsis minuta SPIETH, 1943

ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America

• Genus Hagenulus • Hagenulus marshalli PETERS, FLOWERS, HUBBARD, DOMÍNGUEZ & SAVAGE, 2005 • Genus Hapsiphlebia • Hapsiphlebia anastomosis (DEMOULIN, 1955) • Genus Hermanella • Hermanella costalis (Navás, 1934) • Hermanella froehlichi FERREIRA & DOMÍNGUEZ, 1992 • Hermanella grandis DOMÍNGUEZ & FLOWERS, 1989 • Hermanella maculipennis (ULMER, 1920) • Hermanella guttata DOMÍNGUEZ & FLOWERS, 1989 • Hermanella thelma NEEDHAM & MURHPY, 1924 • Genus Hermanellopsis • Hermanellopsis arsia SAVAGE & PETERS, 1983 • Hermanellopsis incertans (SPIETH, 1943) • Genus Homothraulus • Homothraulus larensis (NAVÁS, 1926) • Homothraulus lucretiae TRAVER, 1960 • Homothraulus misionensis (ESBEN-PETERSEN, 1912) • Genus Hydrosmilodon • Hydrosmilodon gilliesae THOMAS & PÉRU, 2004 • Hydrosmilodon mikei THOMAS & BOUTONNET, 2004 • Hydrosmilodon saltensis FLOWERS & DOMÍNGUEZ, 1992 • Genus Hylister • Hylister plaumanni DOMÍNGUEZ & FLOWERS, 1989 • Genus Leentvaaria • Leentvaaria palpalis DEMOULIN, 1966 • Genus Magallanella • Magallanella flinti PESCADOR & PETERS, 1980 • Genus Massartella • Massartella alegrettae ULMER, 1943 • Massartella brieni (LESTAGE, 1924) • Massartella devani DERKA, 2002 • Massartella venezuelensis PESCADOR & PETERS, 1990 • Massartella sp. PESCADOR & PETERS, 1990 • Genus Massartellopsis • Massartellopsis irarrazavali DEMOULIN, 1955 • Genus Meridialaris • Meridialaris biobionica (ULMER, 1938) • Meridialaris chiloeensis (DEMOULIN, 1955) • Meridialaris diguillina (DEMOULIN, 1955) • Meridialaris inflata PESCADOR & PETERS, 1987 • Meridialaris laminata (ULMER, 1920) • Meridialaris lestagei (ULMER, 1938) • Meridialaris patagonica (LESTAGE, 1931)

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• Meridialaris spina PESCADOR & PETERS, 1987 • Meridialaris tintinnabula PESCADOR & PETERS, 1987 • Genus Microphlebia • Microphlebia pallida SAVAGE & PETERS, 1983 • Microphlebia surinamensis SAVAGE & PETERS, 1983 • Genus Miroculis • Miroculis (Atroari) amazonicus SAVAGE & PETERS, 1983 • Miroculis (Atroari) colombiensis SAVAGE & PETERS, 1983 • Miroculis (Atroari) duckensis SAVAGE & PETERS, 1983 • Miroculis (Miroculis) bicoloratus SAVAGE, 1987 • Miroculis (Miroculis) brasiliaensis SAVAGE & PETERS, 1983 • Miroculis (Miroculis) fittkaui SAVAGE & PETERS, 1983 • Miroculis (Miroculis) marauiae SAVAGE & PETERS, 1983 • Miroculis (Miroculis) nebulosus SAVAGE, 1987 • Miroculis (Miroculis) rossi EDMUNDS, 1963 • Miroculis (Ommaethus) froehlichi SAVAGE & PETERS, 1983 • Miroculis (Ommaethus) mourei SAVAGE & PETERS, 1983 • Miroculis (Yaruma) wandae SAVAGE & PETERS, 1983 • Genus Miroculitus • Miroculitus emersoni (NEEDHAM & MURPHY, 1924) • Genus Needhamella • Needhamella ehrhardti (ULMER, 1920) • Genus Nousia • Nousia bella PESCADOR & PETERS, 1985 • Nousia crena PESCADOR & PETERS, 1985 • Nousia delicata NAVÁS, 1918 • Nousia grandis (DEMOULIN, 1955) • Nousia maculata (DEMOULIN, 1955) • Nousia minor (DEMOULIN, 1955) • Genus Paramaka • Paramaka convexa (SPIETH, 1943) • Genus Penaphlebia • Penaphlebia (Megalophlebia) vinosa (DEMOULIN, 1955) • Penaphlebia (Penaphlebia) barriai PESCADOR & PETERS, 1991 • Penaphlebia (Penaphlebia) chilensis (EATON, 1883) • Penaphlebia (Penaphlebia) exigua DOMÍNGUEZ & PESCADOR, 1983 • Penaphlebia (Penaphlebia) flavidula PESCADOR & PETERS, 1991 • Penaphlebia (Penaphlebia) fulvipes (NEEDHAM & MURPHY, 1924) • Genus Perissophlebiodes • Perissophlebiodes flinti (SAVAGE, 1982) • Genus Rhigotopus • Rhigotopus andinensis PESCADOR & PETERS, 1982 • Genus Secochela • Secochela illiesi PESCADOR & PETERS, 1982

ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America

• Genus Simothraulopsis • Simothraulopsis demerara (TRAVER, 1947) • Genus Terpides • Terpides guyanensis DEMOULIN, 1966 • Genus Thraulodes • Thraulodes bolivianus DOMÍNGUEZ, 1986 • Thraulodes bomplandi (ESBEN-PETERSEN, 1912) • Thraulodes cochunaensis DOMÍNGUEZ, 1987 • Thraulodes colombiae (WALKER, 1853) • Thraulodes consortis DOMÍNGUEZ, 1987 • Thraulodes daidaleus THEW, 1960 • Thraulodes flinti DOMÍNGUEZ, 1987 • Thraulodes furficulus TRAVER, 1946 • Thraulodes guanare CHACÓN, SEGNINI & DOMÍNGUEZ, 1999 • Thraulodes itatiajanus TRAVER & EDMUNDS, 1967 • Thraulodes laetus (EATON, 1883) • Thraulodes limbatus NAVÁS, 1936 • Thraulodes liminaris DOMÍNGUEZ, 1987 • Thraulodes marreroi CHACÓN, SEGNINI & DOMÍNGUEZ, 1999 • Thraulodes mucuy CHACÓN, SEGNINI & DOMÍNGUEZ, 1999 • Thraulodes osiris TRAVER & EDMUNDS, 1967 • Thraulodes papilionis TRAVER & EDMUNDS, 1967 • Thraulodes paysandensis TRAVER, 1964 • Thraulodes regulus TRAVER & EDMUNDS, 1967 • Thraulodes schlingeri TRAVER & EDMUNDS, 1967 • Thraulodes subfasciatus NAVÁS, 1934 • Thraulodes telegraphicus NEEDHAM & MURPHY, 1924 • Thraulodes traverae THEW, 1960 • Thraulodes trijunctus (BANKS, 1918) • Thraulodes ulmeri EDMUNDS, 1950 • Thraulodes venezuelana ULMER, 1943 • Thraulodes vitripennis (BLANCHARD in GAY, 1851) • Genus Tikuna • Tikuna bilineata (NEEDHAM & MURPHY, 1924) • Genus Traverella • Traverella (Traverella) bradleyi (NEEDHAM & MURPHY, 1924) • Traverella (Traverella) montium (ULMER, 1943) • Traverella (Traverella) valdemari (ESBEN-PETERSEN, 1912) • Traverella (Zonda) calingastensis DOMÍNGUEZ, 1995 • Genus Ulmeritoides • Ulmeritoides flavopedes (SPIETH, 1943) • Ulmeritoides haarupi (ESBEN-PETERSEN, 1912) • Ulmeritoides huitoto DOMÍNGUEZ & ZÚÑIGA, 2003 • Ulmeritoides luteotinctus (TRAVER, 1959)

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• Ulmeritoides misionensis DOMÍNGUEZ, 1995 • Ulmeritoides oepa LOPES, DA-SILVA & PY-DANIEL, 2003 • Ulmeritoides patagiatus (THEW, 1960) • Ulmeritoides spinulipenis DOMÍNGUEZ, 1995 • Ulmeritoides uruguayensis (TRAVER, 1959) • Genus Ulmeritus • Ulmeritus balteatus THEW, 1960 • Ulmeritus carbonelli TRAVER, 1956 • Ulmeritus saopaulensis (TRAVER, 1946) FAMILY MELANEMERELLIDAE • Genus Melanemerella • Melanemerella brasiliana ULMER, 1920 FAMILY NESAMELETIDAE • Genus Metamonius • Metamonius anceps (EATON, 1885) FAMILY OLIGONEURIIDAE • Genus Fittkauneuria • Fittkauneuria adusta PESCADOR & PETERS, 1994 • Fittkauneuria carina PESCADOR & PETERS, 1994 • Genus Homoeoneuria • Homoeoneuria (Notachora) fittkaui PESCADOR & PETERS, 1980 • Genus Lachlania • Lachlania boanovae DA-SILVA & PEREIRA, 1993 • Lachlania cacautana (NEEDHAM, 1932) • Lachlania dominguezi PEREIRA, 1989 • Lachlania garciai (NAVÁS, 1912) • Lachlania pallipes (EATON, 1883) • Lachlania radai (NAVÁS, 1926) • Lachlania santosi PEREIRA, 1987 • Genus Oligoneuria • Oligoneuria anomala PICTET, 1843 • Genus Oligoneurioides • Oligoneurioides amazonicus DEMOULIN, 1955 • Genus Spaniophlebia • Spaniophlebia assimilis BANKS, 1913 • Spaniophlebia escomeli COCKERELL, 1926 • Spaniophlebia trailiae EATON, 1881 FAMILY ONISCIGASTRIDAE • Genus Siphlonella • Siphlonella ventilans NEEDHAM & MURPHY, 1924 FAMILY POLYMITARCYIDAE SUBFAMILY ASTHENOPODINAE • Genus Asthenopus • Asthenopus curtus (HAGEN, 1861)

ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America

• Asthenopus gilliesi DOMÍNGUEZ, 1988 • Asthenopus picteti (HUBBARD, 1975) SUBFAMILY CAMPSURINAE • Genus Campsurus • Campsurus albicans (PERCHERON in GUERIN & PERCHERON, 1838) • Campsurus albifilum (WALKER, 1853) • Campsurus argentinus ESBEN-PETERSEN, 1912 • Campsurus assimilis TRAVER, 1944 • Campsurus brasiliensis TRAVER, 1944 • Campsurus burmeisteri ULMER, 1942 • Campsurus claudus NEEDHAM & MURPHY, 1924 • Campsurus corumbanus NEEDHAM & MURPHY, 1924 • Campsurus cuyuniensis TRAVER, 1947 • Campsurus dallasi NAVÁS, 1927 • Campsurus dorsalis (BURMEISTER, 1839) • Campsurus duplicatus SPIETH, 1943 • Campsurus essequibo TRAVER, 1947 • Campsurus evanidus NEEDHAM & MURPHY, 1924 • Campsurus indivisus ULMER, 1942 • Campsurus jorgenseni ESBEN-PETERSEN, 1912 • Campsurus juradinus NAVÁS, 1930 • Campsurus latipennis (WALKER, 1853) • Campsurus litaninensis SPIETH, 1943 • Campsurus longicauda NAVÁS, 1931 • Campsurus lucidus NEEDHAM & MURPHY, 1924 • Campsurus mahunkai PUTHZ, 1973 • Campsurus major NEEDHAM & MURPHY, 1924 • Campsurus melanocephalus PEREIRA & DA-SILVA, 1991 • Campsurus meyeri NAVÁS, 1934 • Campsurus mutilus NEEDHAM & MURPHY, 1924 • Campsurus notatus NEEDHAM & MURPHY, 1924 • Campsurus pallidus NEEDHAM & MURPHY, 1924 • Campsurus paranensis NAVÁS, 1932 • Campsurus paraquarius NAVÁS, 1920 • Campsurus pedicellarius SPIETH, 1943 • Campsurus pfeifferi NAVÁS, 1931 • Campsurus quadridentatus EATON, 1871 • Campsurus scutellaris NEEDHAM & MURPHY, 1924 • Campsurus segnis NEEDHAM & MURPHY, 1924 • Campsurus striatus NEEDHAM & MURPHY, 1924 • Campsurus truncatus ULMER, 1920 • Campsurus ulmeri TRAVER, 1950 • Campsurus violaceus NEEDHAM & MURPHY, 1924 • Campsurus zikani NAVÁS, 1934

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• Genus Tortopus • Tortopus bruchianus (NAVÁS, 1926) • Tortopus harrisi TRAVER, 1950 • Tortopus igaranus NEEDHAM & MURPHY, 1924 • Tortopus obscuripennis DOMÍNGUEZ, 1985 • Tortopus parishi (BANKS, 1918) • Tortopus sarae DOMÍNGUEZ, 1985 • Tortopus unguiculatus (ULMER, 1920) • Tortopus zottai (NAVÁS, 1920)

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8 SYSTEMATICS KEYS TO SOUTH AMERICAN FAMILIES Adults 1 – 2 (1)

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3 (2) – 4(3) – 5(2)

–

Wing venation greatly reduced, three to four longitudinal veins present behind R1 (Figs. 204A, 205A) ................................................................................... Oligoneuriidae Wing venation complete or only moderately reduced, numerous longitudinal veins present behind R1 (Figs. 8A, 72A, 80A, 113A, 123D, 212A) .................................... 2 Base of veins MP2 and CuA strongly divergent from base of MP1 (Figs. 82A, 84A, 212A, 217A); hind wings with numerous longitudinal and cross-veins (Figs. 82B, 84B, 212F); vein MA of hind wings unforked (Figs. 82B, 84B, 212F) ..................................................................................................................................... 3 Base of veins MP2 and CuA slightly divergent from vein MP1 (vein MP2 only may diverge from MP1) (Figs. 8A, 72A, 80A, 113A, 123D); hind wings variable, may be reduced or absent; vein MA of hind wings forked or unforked (Figs. 11B, 78B, 88C, 175D, 202B) ............................................................................................................... 5 Middle and hind legs of male and all legs of female atrophied, nonfunctional .................................................................................................................. Polymitarcyidae All legs of both sexes well-developed, functional ...................................................... 4 Fork of vein MA of fore wings located in basal third of wing (Fig. 84A); male forceps with one long basal segment, terminal segment short (Fig. 85H) or absent (Fig. 84C) ................................................................................................. Euthyplociidae Fork of vein MA of fore wings located at middle of wing (Fig. 82A); male forceps with two long basal segments and a small apical segment (Fig. 82C) ....................... ........................................................................ Ephemeridae.....Hexagenia (Pseudeatonica) Veins MA2 and MP2 basally detached from their respective stems (Figs. 8A, 11A); one (Fig. 27A) or two (Fig. 8A) detached marginal intercalaries present between longitudinal veins; penes hardly visible or not visible externally (Figs. 11C, 27C) ................................................................................................................................. Baetidae Veins MA2 and often MP2 attached basally to their respective stems (Figs. 72A, 88A, 123D, 202A); if present, marginal intercalaries generally attached to longitudinal veins (Figs. 123D, 175C); penes usually prominent and visible externally (Figs. 88D, 123H) ............................................................................................ 6

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Hind wings usually absent (Figs. 72A, 80A, 113A-C); if present (Figs. 90A-F, 93AF), hind wings with long, recurved costal projection located basally (Figs. 90N-P, 93E-F); male eyes generally undivided (Figs. 80C, 102O), rarely divided (Fig. 97D), dorsal portion with hexagonal facets ................................................................. 7 – Hind wings usually present (Figs. 5A-B, 78A-B, 123A-F, 141A-C, 202A-C), rarely absent (Figs. 121A, 140A); costal projection, if present, relatively small and located distal to base (Figs. 4B, 78B, 134B-C, 137G, 141B-C, 142B, 164B-C, 202BC); male eyes distinctly divided into lower and upper portions (Figs. 2A, 218A-C, 225A-B), dorsal portion with square facets (Fig. 140C) ............................................ 9 7 (6) Medial area of mesonotum with a conspicuous circular or oval membranous structure, generally much paler than remaining area (“ommation” Fig. 72B); hind wings absent; fore wings with few cross veins (Fig. 72A); male forceps onesegmented (Fig. 72D) ........................................................................................ Caenidae – Mesonotum not as above; hind wings present or absent; fore wings usually with numerous cross veins; male forceps two- or three- segmented ............................... 8 8 (7) Cubital intercalaries of fore wings absent (Figs. 80A-B); tarsal claws of fore legs of male imago dissimilar, one blunt, other acute (Fig. 80D); male genitalia with short, unsclerotized forceps, segments partially fused (Fig. 80E) ............................................ ................................................................................................ Coryphoridae.....Coryphorus – Cubital intercalaries of fore wings present (Figs. 88A, 113A); tarsal claws of fore leg of male imago similar, both blunt; forceps with at least two distinct segments (Figs. 90M, 97F, 103E) .......................................................................... Leptohyphidae 9(6) Cubital intercalaries of fore wings consisting of series of veinlets, often forked or sinuous, joining vein CuA to the posterior margin (Figs. 4A, 78A,) ..................... 11 – Cubital intercalaries of fore wings variable, but not as above, sometimes absent .............................................................................................................................................. 10 10(9) Two or three basally free short marginal intercalaries present between veins along entire posterior margin of fore wings (Fig. 202A) ....................................................... ................................................................................... Melanemerellidae.....Melanemerella – Marginal intercalaries basally attached or absent (Figs. 123D, 134A, 175C) ........... .................................................................................................................. Leptophlebiidae 11(9) Vein CuP of fore wings strongly recurved (Fig. 78A) ................................................. ............................................................................................. Coloburiscidae.....Murphyella – Vein CuP of fore wings straight or slightly recurved (Fig. 4A) ............................. 12 12(11) Basitarsus (segment I of tarsus partially fused to tibia) of hind legs subequal or longer than tibia; male forceps four-segmented (Fig. 211C); fore wings with dark spots on costal membrane (Fig. 211A) ............................................................... .............................................................................................Oniscigastridae.....Siphlonella – Basitarsus of hind legs shorter than tibia; male forceps three-segmented (Fig. 4C), or if four-segmented (Figs. 5C, 203C); fore wings without dark spots as above (Figs. 5A, 203A) ................................................................................................................ 13

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13(12) Tarsal claws of a pair similar, both apically acute; wings as in Figs. 203A-B; male genitalia as in Fig. 203C ................................................ Nesameletidae.....Metamonius – Tarsal claws of a pair dissimilar, one apically hooked, the other apically blunt; wings as in Figs. 4A-B and 5A-B; male genitalia as in Figs. 4C and 5C .................. ..................................................................................................................... Ameletopsidae Nymphs 1 – 2(1) – 3(2) – 4(3) – 5(2) – 6(5) – 7(6)

Abdominal gills absent (Fig. 79A), thoracic sterna with respiratory evaginations (accessory gills); dense row of setae on anterior margin of femora and tibiae of fore legs (Figs. 3A, 79A) ................................................ Coloburiscidae.....Murphyella Abdominal gills present; setation of anterior margin of femora and tibiae of fore legs not as above ................................................................................................................ 2 Abdominal gills on segments II-VII deeply forked, with margins heavily fringed (Fig. 3L), gills on segment I variable or absent; mandibular tusks projected forward, visible on dorsal view (Figs. 3E-G, T-U) ...................................................... 3 Gills on abdominal segments variable, with margins fringed or not; mandibular tusks absent ......................................................................................................................... 5 Abdominal gills lateral (Fig. 86A); fore legs not modified for burrowing, tibiae cylindrical; mandibular tusks with numerous long setae (Fig. 3E) ......................... .................................................................................................................... Euthyplociidae Abdominal gills dorsal (Fig. 83A); fore legs modified for burrowing, tibiae more or less flattened (Figs. 3B-C); mandibular tusks variable (Figs. 3F-G) ................... 4 Ventral apex of hind tibiae pointed (see arrow in Fig. 3B); mandibular tusks curved upward as viewed laterally (Fig. 3F) ................................................................. ........................................................................ Ephemeridae.....Hexagenia (Pseudeatonica) Ventral apex of hind tibiae rounded (Fig. 3C); mandibular tusks almost straight or with apices curved downward as viewed laterally (Fig. 3G) ............. .................................................................................................................. Polymitarcyidae Abdominal gills on segment I or II operculate, covering succeeding pairs (Figs. 3H-K) .................................................................................................................................... 6 Abdominal gills variable, those on segments I or II never operculate ................... 8 Abdominal gills on segment I operculate, covering succeeding pairs (Fig. 3H); gills present on segments I-IV; abdomen with a median row of projections on anterior terga ...................................................................Oniscigastridae.....Siphlonella Abdominal gills on segment II operculate (Figs. 3I-K); gills present on segments II-V or II-VI; abdomen with median projections on anterior terga variable ................................................................................................................................. 7 Abdominal gills on segments I-VI; gills on segment I reduced, filiform (almost above hind coxae) (Fig. 75A); operculate gills on segment II large, subquadrangular, with two ridges forming an Y-shaped figure (Fig. 3I); gills III-VI with fringed margins and single lamella (Fig. 3M) ...................................... Caenidae

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T

U

Plate 3. Figs. 3A-U. Nymphs: 3A, Murphyella (Coloburiscidae), fore leg; 3B, Hexagenia (Ephemeridae), hind leg; 3C, Tortopus (Polymitarcyidae: Campsurinae), hind leg; 3D, Lachlania (Oligoneuriidae), fore leg; 3E, Euthyplocia (Euthyplociidae), head d.v.; 3F, Hexagenia (Ephemeridae), head l.v.; 3G, Ephoron (Polymitarcyidae : Polymitarcyinae), head l.v.; 3H, Siphlonella (Oniscigastridae), abdominal segments I-V, d.v.; 3I, Caenis (Caenidae), abdomen d.v.; 3J, Coryphorus (Coryphoridae), abdomen d.v.; 3K, Tricorythodes (Leptohyphidae), abdomen, d.v.; 3L, Euthyplocia (Euthyplociidae), gill II; 3M, Caenis (Caenidae), gill III; 3N, Chaquihua (Ameletopsidae), mandible; 3O, Chaquihua (Ameletopsidae), maxillae; 3P, Metamonius (Nesameletidae), mandible; 3Q, Metamonius (Nesameletidae), maxillae; 3R, Baetidae, head; 3S, Nesameletidae, head; 3T, Tortopus (Polymitarcyidae : Campsurinae), head; 3U, Asthenopus (Polymitarcyidae: Asthenopodinae), head. (C = canines; ES = epicranial suture; OG = operculate gill; P = maxillary palpi; T = mandibular tusk).

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Abdominal gills present only on segments II-V or II-VI, operculate gill on segment II variable, but not as above (Figs. 3J-K); gills margins entire or with imbricate lobes and with two or more lamellae ........................................................ 12 8(5) Maxillary and labial palpi multisegmented and filiform; mandibles and maxillae strongly modified for predation (Figs. 3N-O) .................................. Ameletopsidae – Maxillary and labial palpi at most three-segmented; mandibles and maxillae not strongly modified for predation (Figs. 3P-Q) .............................................................. 9 9(8) Internal edge of fore femora with double row of long setae (Fig. 3D); maxillary and labial palpi two-segmented; gill tufts at base of maxillae ...... Oligoneuriidae – Fore femora not as above; maxillary and labial palpi three-segmented; without gills at base of maxillae ................................................................................................... 10 10 (9) Clypeus fused with frons; head usually prognathous (Fig. 124A), rarely hypognathous (Fig. 231A); abdominal gills various but generally composed of narrow, leaf-like, or fringed ventral and dorsal lamellae .............. Leptophlebiidae – Clypeus not fused with frons; head hypognathous (Figs. 25B, 203D); abdominal gills unilamellate, usually oval, sometimes folded at base ....................................... 11 11(10) Lateral branches of epicranial suture ending anterior to lateral ocelli (Fig. 3R); antennae long, at least twice as long as width of head; gills on abdominal segments I-V, I-VII, or II-VII; posterolateral corners of abdominal segments not expanded in form of flat lateral projections, or, if expanded, projections little developed .................................................................................................... Baetidae – Lateral branches of epicranial suture ending at lateral ocelli (Fig. 3S); antennae relatively short (Fig. 3S); gills present on abdominal segments I-VII (Fig. 203D); posterolateral corners of abdominal segments expanded in form of flat lateral projections ....................................................................... Nesameletidae.....Metamonius 12(7) Abdominal terga I-IX with paired submedian tubercles (Fig. 201A) ....................... ................................................................................... Melanemerellidae.....Melanemerella – Abdominal terga generally lacking tubercles, or if present with single median tubercle ............................................................................................................................... 13 13 (12) Opercular gills quadrangular (Fig. 3J); tubercles present on head, pronotum and abdominal terga I-III and VII-IX (Figs. 80F, 81A); gills enclosed in a gill chamber ................................................................................................ Coryphoridae.....Coryphorus – Opercular gills subquadrangular, triangular, subtriangular or oval; tubercles generally absent, if present never on all three tagmata simultaneously; without gill chamber .................................................................................................... Leptohyphidae

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CLAVES

PARA LAS

FAMILIAS

SUDAMERICANAS

Adultos 1 – 2 (1)

–

3 (2) – 4 (3) – 5 (2)

–

6 (5)

–

Venación alar grandemente reducida, sólo tres o cuatro venas longitudinales presentes detrás de R1 (Figs. 204A, 205A) ........................................ Oligoneuriidae Venación alar completa o sólo moderadamente reducida, numerosas venas longitudinales presentes detrás de R1 (Figs. 8A, 72A, 80A, 113A, 123D, 212A) ........................ 2 Base de las venas MP2 y CuA fuertemente divergentes de la base de MP1 (Figs. 82A, 84A, 212A, 217A); alas posteriores con numerosas venas longitudinales y transversales (Figs. 82B, 84B, 212F); vena MA del ala posterior no bifurcada (Figs. 82B, 84B, 212F) .................................................................................................................. 3 Base de las venas MP2 y CuA levemente divergentes de la vena MP1 (puede divergir solamente la vena MP2 de MP1) (Figs. 8A, 72A, 80A, 113A, 123D); alas posteriores variables, pudiendo estar reducidas o ausentes; vena MA del ala posterior bifurcada o no bifurcada (Figs. 11B, 78B, 88C, 175D, 202B) ................. 5 Patas medias y posteriores del macho y todas las patas de la hembra débiles, no funcionales ............................................................................................. Polymitarcyidae Todas las patas en ambos sexos bien desarrolladas, funcionales ............................. 4 Horquilla de la vena MA de las alas anteriores ubicada en el tercio basal del ala (Fig. 84A); fórceps genitales del macho con un largo segmento basal y un corto segmento terminal (Fig. 85H) o ninguno (Fig. 84C) ...................... Euthyplociidae Horquilla de la vena MA del ala anterior ubicada en la mitad del ala (Fig. 82A); fórceps genitales del macho con dos largos segmentos basales y un corto segmento apical (Fig. 82C) ....................... Ephemeridae.....Hexagenia (Pseudeatonica) Venas MA2 y MP2 separadas basalmente de sus troncos respectivos (Figs. 8A, 11A); una (Fig. 27A) o dos (Fig. 8A) intercalares marginales sueltas entre las venas longitudinales; penes escasamente visibles o no visibles externamente (Figs. 11C, 27C) ........................................................................................................................ Baetidae Venas MA2 y a menudo MP2 unidas basalmente a sus troncos respectivos (Figs. 72A, 88A, 123D, 202A); intercalares marginales si presentes, generalmente unidas a venas longitudinales (Figs. 123D, 175C); penes usualmente bien desarrollados y visibles externamente (Figs. 88D, 123H) ...................................................................... 6 Alas posteriores generalmente ausentes (Figs. 72A, 80A, 113A-C); si están presentes (Figs. 90A-F, 93A-F) proyección costal larga y recurvada, ubicada basalmente (Figs. 90N-P, 93E-F); ojos del macho generalmente no divididos (Figs. 80C, 102O), raramente divididos (Fig. 97D), porción dorsal con facetas hexagonales ......................................................................................................................... 7 Alas posteriores generalmente presentes (Figs. 5A-B, 78A-B, 123A-F, 141A-C, 202A-C), raramente ausentes (Figs. 121A, 140A); proyección costal, si presente, relativamente pequeña y ubicada lejos de la base (Figs. 4B, 78B, 134B-C, 137G, 141B-C, 142B, 164B-C, 202B-C); ojos del macho marcadamente divididos en porciones dorsal y ventral (Figs. 2A, 218A-C, 225A-B), porción dorsal con facetas cuadrangulares (Fig. 140C) ............................................................................................... 9

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– 8 (7)

– 9 (6) – 10 (9) – 11 (9) – 12 (11)

–

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Zona media del mesonoto con una estructura circular u ovalada notoria, generalmente mucho más clara que el resto (“ommation” Fig. 72B); alas posteriores ausentes; alas anteriores con pocas venas transversas (Fig. 72A); fórceps del macho unisegmentados (Fig. 72D) ......................................... Caenidae Mesonoto no como en el caso anterior; ala posterior ausente o presente; ala anterior usualmente con numerosas venas transversas; fórceps del macho bi- o trisegmentados .................................................................................................................... 8 Intercalares cubitales del ala anterior ausentes (Figs. 80A-B); uñas tarsales de la pata anterior del imago macho desiguales, una roma y la otra aguda (Fig. 80D); genitalia masculina con fórceps cortos no esclerosados, segmentos parcialmente fusionados (Fig. 80E) ........................................................ Coryphoridae.....Coryphorus Intercalares cubitales del ala anterior presentes (Figs. 88A, 113A); uñas tarsales de la pata anterior del imago macho iguales, ambas romas; fórceps al menos con dos segmentos evidentes (Figs. 90M, 97F, 103E) ................................... Leptohyphidae Las intercalares cubitales del ala anterior consisten en una serie de venillas, a menudo bifurcadas o sinuosas, que unen la vena CuA al margen posterior (Figs. 4A, 78A) ............................................................................................................................. 11 Intercalares cubitales del ala anterior variables pero no como en el dilema anterior, algunas veces ausentes .................................................................................................... 10 Dos o tres cortas intercalares marginales sueltas basalmente presentes entre las venas a lo largo de todo el margen externo del ala (Fig. 202A) ................................ ................................................................................... Melanemerellidae.....Melanemerella Marginales intercalares generalmente unidas basalmente o ausentes (Figs. 123D, 134A, 175C) .......................................................................................... Leptophlebiidae Vena CuP del ala anterior fuertemente recurvada (Fig. 78A) .................................... ............................................................................................. Coloburiscidae.....Murphyella Vena CuP del ala anterior recta o levemente recurvada (Fig. 4A) ........................ 12 Basitarso (segmento I del tarso parcialmente fusionado a la tibia) de las patas posteriores subigual o más largo que la tibia; fórceps de la genitalia masculina tetra segmentados (Fig. 211C); alas anteriores con manchas oscuras en la membrana costal (Fig. 211A) ............................................................Oniscigastridae.....Siphlonella Basitarso de las patas posteriores más corto que la tibia; fórceps de la genitalia masculina trisegmentados (Fig.4C), o si tetrasegmentados (Figs. 5C, 203C), las alas anteriores no poseen marcas oscuras tan notorias como en el dilema anterior (Figs. 5A, 203A) ................................................................................................................ 13 Uñas tarsales del par similares, ambas apicalmente aguzadas; alas como en las Figs. 203A-B; genitalia masculina como en la Fig. 203C ...................................................... ............................................................................................ Nesameletidae.....Metamonius Uñas tarsales del par diferentes entre si, una aguzada apicalmente, la otra roma; alas como en las Figs.4A-B, 5A-B; genitalia masculina como en las Figs. 4C, 5C ..................................................................................................................... Ameletopsidae

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Ninfas 1

– 2(1)

– 3(2) – 4(3) – 5(2) – 6(5)

– 7(6)

–

Branquias abdominales ausentes (Fig. 79A), con evaginaciones respiratorias (branquias accesorias) en los esternos torácicos; fémures y tibias de las patas anteriores con una densa hilera de setas sobre el borde anterior (Figs. 3A, 79A) ............................................................................................. Coloburiscidae.....Murphyella Branquias abdominales presentes, borde anterior de las patas anteriores con setas variables ................................................................................................................................ 2 Branquias abdominales en los segmentos II-VII profundamente hendidas y con los márgenes con muchos flecos (Fig. 3L), branquias en el segmento I variables o ausentes; colmillo mandibular proyectándose por el frente de la cabeza (Figs. 3EG, T-U) ................................................................................................................................. 3 Branquias abdominales variables, márgenes de las branquias con o sin flecos; colmillos mandibulares ausentes ..................................................................................... 5 Branquias abdominales laterales (Fig. 86A); las patas anteriores no modificadas para cavar, tibias cilíndricas; colmillos mandibulares con numerosas setas largas (Fig. 3E) .................................................................................................... Euthyplociidae Branquias abdominales dorsales (Fig. 83A); las patas anteriores modificadas para cavar, tibias aplanadas (Figs. 3B-C); colmillos mandibulares variables ................... 4 Ápice ventral de las tibias posteriores terminado en punta (flecha en Fig. 3B); en vista lateral los colmillos mandibulares curvados hacia arriba (Fig. 3F) .................. ........................................................................ Ephemeridae.....Hexagenia (Pseudeatonica) Apice ventral de las tibias posteriores redondeado (Fig. 3C); en vista lateral los colmillos mandibulares casi rectos o curvados hacia abajo apicalmente (Fig. 3G) .................................................................................................................. Polymitarcyidae Branquias abdominales en el segmento I ó II operculares, cubriendo las restantes (Figs. 3H-K) ....................................................................................................................... 6 Branquias abdominales variables, pero las del segmento I ó II nunca operculares ................................................................................................................................................ 8 Branquias abdominales del segmento I operculares, cubriendo las restantes (Fig. 3H); branquias en los segmentos abdominales II-V; abdomen con una hilera mediana de proyecciones sobre los tergos anteriores ................................................ .............................................................................................Oniscigastridae.....Siphlonella Branquias abdominales del segmento II operculares (Figs. 3I-K); branquias presentes en los segmentos II-V ó II-VI; proyecciones medianas en los tergos anteriores variadas .............................................................................................................. 7 Branquias abdominales en los segmentos I-VI, las branquias del segmento I reducidas, filiformes (casi sobre coxas posteriores) (Fig. 75A); branquias operculares del segmento II grandes subcuadrangulares, con dos costillas unidas formando una figura en forma de Y (Fig. 3I); branquias de los segmentos III-VI con los márgenes con flecos y lamelas únicas (Fig. 3M) ......................................... Caenidae Branquias abdominales en los segmentos II-V o II-VI; branquias operculares del segmento II variadas pero nunca como arriba (Figs. 3J-K); branquias enteras o con pequeños lóbulos imbricados y con dos o más lamelas .................................. 12

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Palpos maxilares y labiales multisegmentados y filiformes; mandíbulas y maxilas marcadamente modificadas para depredar (Figs. 3N-O) ............... Ameletopsidae – Palpos maxilares y labiales como máximo trisegmentados, mandíbulas y maxilas no fuertemente modificadas para depredar (Figs. 3P-Q) .......................................... 9 9(8) Fémures anteriores con una doble hilera de largas setas sobre el lado interno (Fig. 3D); palpos maxilares y labiales bi-segmentados; un penacho de branquias unido a la base de las maxilas ............................................................................. Oligoneuriidae – Fémures anteriores sin tales setas; palpos maxilares y labiales trisegmentados; sin branquias en la base de las maxilas .............................................................................. 10 10(9) Clipeo fusionado a la frente; cabeza usualmente prognata (Fig. 124A), raramente hipognata (Fig. 231A); branquias abdominales variables pero generalmente compuestas de una lámina ventral y una dorsal, delgadas, en forma de hoja o festoneadas ............................................................................................ Leptophlebiidae – Clipeo no fusionado a la frente; cabeza hipognata (Figs. 25B, 203D); branquias abdominales usualmente ovales, compuestas de una única lámina, a veces plegada sobre sí misma en la base ............................................................................................... 11 11(10) Ramas laterales de la sutura epicranial terminando anteriormente a los ocelos laterales (Fig. 3R); antenas largas, dos o más veces el ancho de la cabeza; branquias en los segmentos abdominales I-V, I-VII, o II-VII; ángulos posterolaterales de los segmentos abdominales no expandidos en proyecciones laterales planas, o si están presentes, poco desarrolladas ........................................................................... Baetidae – Ramas laterales de la sutura epicranial terminando en los ocelos laterales (Fig. 3S); antenas relativamente cortas (Fig. 3S); branquias en los segmentos abdominales IVII (Fig. 203D); ángulos posterolaterales de los segmentos abdominales expandidos en proyecciones laterales planas .......................... Nesameletidae.....Metamonius 12(7) Tubérculos submedianos pares presentes en los tergos abdominales I-IX (Fig. 201A) ....................................................................... Melanemerellidae.....Melanemerella – Generalmente sin tubérculos abdominales, si están presentes son únicos y de posición media .................................................................................................................. 13 13(12) Branquias operculares cuadrangulares (Fig. 3J); con tubérculos en cabeza, pronoto y tergos abdominales I-III y VII-IX (Figs. 80F, 81A); branquias encerradas en una cámara branquial ................................................................ Coryphoridae.....Coryphorus – Branquias operculares subcuadrangulares, triangulares, subtriangulares u ovales; tubérculos generalmente ausentes, si presentes nunca en los tres tagmas simultáneamente; sin cámara branquial .......................................................... Leptohyphidae

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FAMILY AMELETOPSIDAE Taxonomy: The family Ameletopsidae contains two subfamilies, Ameletopsinae and Chiloporterinae, both known from South America. This family comprises five genera, two from Australia (the fossil Promirara, and recent Mirawara), one from New Zealand (Ameletopsis), and two from South America (Chaquihua and Chiloporter). Adult Characteristics: Fore wings with numerous veins (Figs. 4A, 5A); hind wings large (Figs. 4B, 5B); vein CuA of fore wings connected to hind margin of wing by several veinlets (Figs. 4A, 5A); tarsal claws of each pair dissimilar, one blunt the other acute. Nymphal Characteristics: Head enlarged, wide, with predatory mouthparts; labial and maxillary palpi are multisegmented and filiform; tarsus 2 or 3 segmented (Fig. 5D); caudal filaments relatively short. Distribution: Australia, New Zealand, Southern Argentina and Chile. Biology: The nymphs of this family are predatory and inhabit cold rivers and streams at high latitudes in the South. KEYS TO SOUTH AMERICAN GENERA Adults 1 –

Terminal filament much shorter than cerci; genital forceps three-segmented (Fig. 4C) ............................................................................... Ameletopsinae.....Chaquihua Terminal filament subequal in length to cerci; genital forceps four-segmented (Fig. 5C) ............................................................................. Chiloporterinae.....Chiloporter

Nymphs 1 –

CLAVES

Abdominal gills consisting of dorsal and ventral portions, ventral lamella with fringed margins, dorsal lamella circular, pigmented, covering the tergum (Figs. 6A, 227A) .......................................................................... Chiloporterinae.....Chiloporter Abdominal gills consisting of single entire lamella, oval, non-pigmented, laterally positioned, not covering the abdomen (Fig. 227B) .... Ameletopsinae.....Chaquihua PARA LOS

GÉNEROS

SUDAMERICANOS

Adultos 1 –

Filamento terminal mucho más corto que los cercos; fórceps genitales trisegmentados (Fig. 4C) ....................................................... Ameletopsinae.....Chaquihua Filamento terminal subigual en longitud que los cercos; fórceps genitales tetrasegmentados (Fig. 5C) ...................................................... Chiloporterinae.....Chiloporter

Ninfas 1

Branquias abdominales consistiendo de láminas dorsal y ventral, lámina ventral con los márgenes con flecos, lámina dorsal circular, pigmentada, cubriendo el tergo (Figs. 6A, 227A) ...................................................... Chiloporterinae.....Chiloporter

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Branquias abdominales consistiendo de una única lámina, ovaladas, no pigmentadas, de posición lateral, no cubriendo el abdomen (Fig. 227B) ................ ................................................................................................ Ameletopsinae.....Chaquihua SUBFAMILY AMELETOPSINAE

The subfamily Ameletopsinae contains the genus Chaquihua DEMOULIN, known only from Chile, and three other genera known from Australia, New Zealand, and the Baltic Eocene. Genus Chaquihua DEMOULIN Chaquihua DEMOULIN, 1955d: 11. (Type-species: Chaquihua penai DEMOULIN, original designation).

Taxonomy: Chaquihua has been described from the adult and egg stages. Adult Characteristics: 1) Genital forceps three-segmented (Fig. 4C); 2) sternum IX of female rounded with a short median incision; and 3) costal projection of hind wing acute (Fig. 4B). Nymphal Characteristics (Fig. 227B): Although the nymphs remain undescribed in the literature, we have seen nymphal specimens of this genus. The nymphs can be characterized by the following combination of characters: 1) Head wide, almost as broad as thorax; and 2) abdominal gills with unpigmented lamella and pigmented tracheoles, and non-fringed or smooth along lamellar margins. Distribution: Southern Chile. Biology: Nymphs are carnivorous and inhabit clean and cold streams.

A

B

C

Plate 4. Chaquihua penai. Figs. 4A-C, male imago: 4A, fore wing; 4B, hind wing; 4C, genitalia v.v. Figs. 4A-C modified from DEMOULIN, 1955d.

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KEY

TO

CHAQUIHUA

Adults 1 – CLAVE

Wing membranes and veins orangish ............................................... Chaquihua bullocki Wing membranes hyaline, veins brownish ........................................... Chaquihua penai PARA

CHAQUIHUA

Adultos 1 –

Membranas y venas de las alas anaranjadas .................................... Chaquihua bullocki Membranas de las alas claras, venas castañas ...................................... Chaquihua penai

Chaquihua bullocki (NAVÁS) Euthyplocia bullocki NAVÁS, 1930a: 328 (female); LESTAGE, 1931b: 59; ULMER, 1938: 102. Chaquihua bullocki; EDMUNDS et al., 1976: 281; HUBBARD, 1982a: 272.

Taxonomy: The adults of Chaquihua bullocki can be distinguished from Chaquihua penai by the generally pale orange color of the wing membrane and veins, body, and legs. Distribution and Biology: Chile. Adults were collected in February. Chaquihua penai DEMOULIN Chaquihua peñai DEMOULIN, 1955d: 11 (male, female). Chaquihua penai; KOSS & EDMUNDS, 1974: 299 (egg); HUBBARD, 1982a: 272.

Taxonomy: The adults of Chaquihua penai can be distinguished from Chaquihua bullocki by the following combination of characters: 1) Wing membrane hyaline, with veins brown; 2) thorax brownish; and 3) abdomen purplish with a wide median longitudinal band, which attenuates in last three segments. Distribution and Biology: Chile (Río Chaquihua, Enco, X Region). Adults were collected in February and March. SUBFAMILY CHILOPORTERINAE The subfamily Chiloporterinae is monogeneric, and the genus Chiloporter LESTAGE, is known only from Chile and Argentina. Genus Chiloporter LESTAGE Chiloporter LESTAGE, 1931b: 50; DEMOULIN, 1955d: 5; DOMÍNGUEZ et al., 1994: 32; MERCADO & ELLIOTT, 2005: 230. (Type-species: Chiloporter eatoni LESTAGE, original designation). Ameletopsis [partim]; DEMOULIN, 1952c: 171.

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Taxonomy: Chiloporter is known from nymphs and adults. The two species included in the genus, C. eatoni and C. penai were recently synonymized by MERCADO & ELLIOTT (2005), with C. eatoni as the senior synonym. Adult Characteristics: 1) Costal projection of hind wing rounded (Fig. 5B); 2) forceps of genitalia four-segmented (Fig. 5C); and 3) tarsal claws dissimilar. Nymphal Characteristics (Fig. 227A): 1) Head wider than thorax (Fig. 6A); 2) ventral lamellae of abdominal gills fringed, dorsal lamellae pigmented, subcircular flat and covering abdominal terga; 3) caudal filaments fringed (Fig. 6A); and 4) large eyes covering approximately 2/3 of head. Distribution: Argentina (Neuquén Prov., 40ºS to Chubut Prov., approximately 42ºS) and Chile (VII Region, 35º24´S to XI Región, 45º34´S). Biology: The carnivorous nymphs are found in well oxygenated rivers. The nymphs are mainly crawlers, but also are capable of short bursts of fast swimming, using the caudal filaments as a fin. Based on gut content analysis, the nymphs appear to feed mainly on other mayfly nymphs, caddisfly and chironomid (Diptera) larvae. Adults emerged from December to mid-January. Emergence and mating flight occur at dusk, mating adults were observed flying in an up and down fashion. Chiloporter eatoni LESTAGE “Genus and species undetermined”; EATON, 1885: 229 (nymphal exuviae). Chiloporter eatoni LESTAGE, 1931b: 50; LESTAGE, 1935a: 97; LESTAGE, 1938: 176; DEMOULIN, 1955c: 3; DEMOULIN, 1955d: 6; HUBBARD, 1982a: 272; WAIS, 1985: 2170 (nymph); WAIS & BONETTO, 1988: 1457; HUBBARD et al., 1992: 209; DOMÍNGUEZ et al., 1994: 32 (nymph). Ameletopsis eatoni; DEMOULIN, 1952c: 171.

A

B

C

D

Plate 5. Chiloporter eatoni. Figs. 5A-D, male imago: 5A, fore wing and detail; 5B, hind wing; 5C, genitalia v.v. Fig. 5D, nymph: fore tarsus. Figs. 5A-D modified from DEMOULIN, 1955d.

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Chiloporter penai DEMOULIN, 1955d: 6 (male, nymph); KOSS & EDMUNDS, 1974: 299 (egg); HUBBARD, 1982a: 272; DOMÍNGUEZ et al., 1994: 32 (male); MERCADO & ELLIOTT, 2005: 231.

Taxonomy: Chiloporter eatoni is the only known species in the genus, and can be characterized by the features mentioned in the generic discussion. Distribution and Biology: As in generic discussion.

A

Plate 6. Chiloporter eatoni, nymph. Fig. 6A: habitus. Drawing by A. Dupuy.

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FAMILY BAETIDAE Taxonomy: The family Baetidae, established by LEACH in 1815, has a nearly cosmopolitan dstribution and presently includes almost 100 genera worldwide. South America has nearly one third of all currently described genera in the family, and the Baetidae are second only to the family Leptophlebiidae in taxa diversity in the region. Adult Characteristics: 1) Males generally with compound eyes turbinate, divided into a dorsal and a ventral region (Figs. 20A, D, 40A, 64A); 2) fore wings with veins IMA, MA2, IMP and MP2 detached basally (Fig. 8A); 3) hind wings small or absent; 4) legs with two tarsal claws: one blunt, one acute; 5) genitalia: forceps three or four segmented and sclerotized, penes membranous; 6) terminal filament absent. Nymphal Characteristics: 1) Epicranial suture extended beyond lateral ocelli; 2) antennae longer than head capsule; 3) abdominal gills not operculate, with only one lamella on abdominal segments I-VII, II-VII, I-V or I-VI; 4) terminal filament of variable length. Distribution: Cosmopolitan, except some oceanic islands, New Zealand and Antarctica. Biology: The nymphs inhabit a variety of lotic and lentic habitats with sandy or rocky substrates. Baetid nymphs are found in the full range of lotic habitats from riffles to pools and nearly all possible microhabitats including rocks, leaf packs, and woody debris. Many nymphs live among emergent vegetation and others can be found in the splash zone of waterfalls. Moreover, some nymphs live associated with bivalves, in mutualism. Some nymphs are not tolerant of poorly oxygenated water while others can live in waters where oxygen levels are low. Most nymphs feed on particulate organic matter suspended in the water column or deposited on the substrate while others are grazers or scrapers of algae and bacteria. A few nymphs are predators, feeding on larvae of Simuliidae or even small nymphs of Odonata. The subimagos generally emerge in the afternoon and evening. The nuptial flight can be observed at noon or especially in the afternoon. KEYS TO SOUTH AMERICAN GENERA Male Imago 1 – 2(1) – 3(2) – 4(3) – 5(4) –

Males with compound eyes not turbinate; basal segment of forceps longer than segment II (Figs. 18D, H) .................................................................................... Aturbina Males with compound eyes turbinate (Figs. 20A, D, 40A, 64A); basal segment of forceps shorter than segment II (Fig. 62D) ................................................................. 2 Body with small reddish brown spots ............................................................ Callibaetis Body without small spots ................................................................................................. 3 Fore wings with spots ........................................................................................ Moribaetis Fore wings without spots (Fig. 62B) .............................................................................. 4 Hind wings absent ............................................................................................................. 5 Hind wings present .......................................................................................................... 11 Basal segment of forceps with distomedial projection (Fig. 20C) .......................... 6 Basal segment of forceps without projection (Fig. 47B) ........................................... 7

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6(5) – 7(5) – 8(7) – 9(7) – 10(9) – 11(4) – 12(11) – 13(12) – 14(13) – 15(14) – 16(14) –

Postero lateral margin of styliger plate with small projection (Fig. 64C) ................ ...................................................................................................................................... Varipes Posterior margin of styliger plate without projection (Fig. 20C) ................. Baetodes Segment II of forceps with distomedial projection (Fig. 47B) ................................ 8 Segment II of forceps without projection ................................................................... 9 Apical margin of styliger plate with small distomedial projection (Fig. 47B); specimens larger (5.0 to 5.5 mm) .................................................................... Guajirolus Apical margin of styliger plate without projection (Fig. 40C); specimens smaller (3.5 to 4.0 mm) ........................................................................................................... Chane Apical margin of styliger plate with a subtriangular projection (Fig. 16A) ............. .................................................................................................................................. Apobaetis Styliger plate without projection (Fig. 42D) ............................................................... 10 Segment III of forceps short and rounded (Fig. 42D) ................... Cloeodes (in part) Segment III of forceps elongated (Fig. 8B) ............................................. Americabaetis Hind wings with 3 longitudinal veins, the second vein bifurcate (Fig. 11B) ........... .................................................................................................................................. Andesiops Hind wings with 2 or 3 longitudinal veins, the second vein never bifurcate (Fig. 30C) .............................................................................................................................................. 12 Forceps apparently four-segmented ..................................................................... Fallceon Forceps three-segmented (Fig. 54C) ............................................................................ 13 Hind wings with subquadrangular costal projection (Fig. 54F) .............. Paracloeodes Hind wings with pointed costal projection (Fig. 30C) ............................................. 14 Segment III of forceps short and rounded (Fig. 42D) ............................................ 15 Segment III of forceps elongate (Fig. 30E) ............................................................... 16 Forceps with bases narrowly separated, basal segment without projection (Fig. 42D); fore wings with vein A1 not strongly curved (Fig. 42A) ..... Cloeodes (in part) Forceps with bases widely separated, basal segment with small distomedial projection (Fig. 51D); fore wings with vein A1 strongly curved (Fig. 51A) ............ ................................................................................................................................ Mayobaetis Fore wings with anal margin rounded (Fig. 30A) ................................Camelobaetidius Fore wings with anal margin relatively straight (Fig. 62B) ............................. Tupiara

Nymphs 1 – 2(1) – 3(2) – 4(3) – 5(4)

Tarsal claws spatulate (Fig. 32I), with 5 to 40 denticles .....................Camelobaetidius Tarsal claws apically pointed (Fig. 19H) ........................................................................ 2 Gills on abdominal segments I-V ........................................................................ Baetodes Gills on abdominal segments I-VI, I-VII or II-VII .................................................... 3 Abdominal gills with two lamellae (Figs. 29M-O) ....................................... Callibaetis Abdominal gills with a single lamella (Fig. 9J) ............................................................. 4 Tarsal claws without denticles (Fig. 44J) ....................................................................... 5 Tarsal claws with denticles (Figs. 9D, 14D) although these can be very small (Fig. 19H) ...................................................................................................................................... 6 Tarsal claws long (0.75 to 0.8 times length of tarsi) (Fig. 16I); tibiae without an arc of long setae (Fig. 16I) ....................................................................................... Apobaetis

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Tarsal claws short (less than 0.6 times length of tarsi) (Fig. 44J); tibiae with an arc of long setae basally (Fig. 44H) ...................................................... Cloeodes/ Bernerius 1 Tarsal claws longer than 1/2 the length of tarsi (Fig. 55H) .................................... ............................................................................................................. Iguaira/ Paracloeodes 2 Tarsal claws shorter than 1/2 the length of tarsi (Fig. 49N) ................................... 7 Mandibles long and broad with very broad and protruding molar region (Figs. 49D-G); maxillary canines strongly protruding, the second one half the length of others (carnivorous kind) (Fig. 49K) ......................................................... Harpagobaetis Mandibles more or less triangular, molar region wide and not protruding (Figs. 63C-D); maxillary canines normal and subequal in length (herbivorous kind) (Fig. 63F) ....................................................................................................................................... 8 Tarsal claws with subapical denticle longer than others (Fig. 48O); prosthecae of mandibles inserted in concavity (Figs. 48B-C) ............................................................ 9 Tarsal claws with subapical denticle subequal in length to others (Fig. 13L); prosthecae of mandibles not inserted in concavity (Figs. 13C, E) ....................... 10 Segment I of labial palpi never transverse to glossae and paraglossae, segment II and III without rows of long setae (Fig. 48F); glossae dorsally with transverse row of strong setae (Fig. 48F) .................................................................................. Guajirolus Segment I of labial palpi transverse to glossae and paraglossae, segment II and III with two rows of long setae (Fig. 41H); glossae without transverse row of strong setae (Fig. 41H) ........................................................................................................... Chane Tarsal claws with 1 or 2 strong setae near base of subapical denticle on external margin (Figs. 21K, 52I, 57J) .......................................................................................... 11 Tarsal claws without strong setae (Fig. 9H) or with pair of subapical fine setae (Fig. 13L) ............................................................................................................................ 14 Tarsal claws with 2 rows of denticles (Fig. 60I) ............................................. Spiritiops Tarsal claws with 1 row of denticles (Fig. 21K) ........................................................ 12 Scape and pedicel dorsoventrally flattened; scape 3 times the length of the pedicel (Fig. 52A); cerci with row of spines on external margin (Fig. 52J) ......... Mayobaetis Scape and pedicel cylindrical or subcylindrical; scape 1.0 to 1.5 times the length of the pedicel; cerci without spines on external margin ............................................... 13 Hind wing pads rudimentary if present; terminal filament < 0.25 the length of cerci; paraprocts with posterior margins smooth ........................................ Prebaetodes Hind wing pads well developed; terminal filament 0.5 the length of cerci; paraprocts wih spines on posterior margins ................................................. Moribaetis Gills present on abdominal segments I-VI; labrum trapezoidal (Fig. 61A), with lateral margins divergent from the base ........................................................ Tomedontus Gills present on abdominal segments I or II-VII; labrum variable but not as above .............................................................................................................................................. 15

Cloeodes and Bernerius are in revision at this moment, and there are some doubts about the validity of the last one. 2 Paracloeodes and Iguaira are in revision at this moment, and there are some doubts about the validity of the last one, as both genera appear indistinguishable. 1

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15(14) Tarsal claws apically with paired bristle setae (Fig. 13L) ......................................... 16 – Tarsal claws without setae (Fig. 9H) ............................................................................ 17 16(15) Mandibles with setae between mola and prostheca; tarsal claws with 1 row of denticles ......................................................................................................Fallceon (in part) – Mandibles without setae between mola and prostheca (Figs.13C, E); tarsal claws with 2 rows of denticles, although the second row can be reduced (Figs. 13L, 14H) .................................................................................................................................. Andesiops 17(15) Femora with abundant strong and long spine-like setae on dorsal and ventral margins (Figs. 65I, K) ...................................................................................................... 18 – Femora with or without long spine-like setae, if such setae present they are not abundant on either dorsal or ventral margins (Fig. 9G) .......................................... 19 18(17) Femora II and III with a transverse row of long spine-like setae on proximal third (Figs. 65K; 66M; 67H); left mandible with incisors completely fused (Fig. 65C); segment III of labial palpi rounded (Fig. 65H) ................................................. Varipes – Femora II and III without a transverse row of long spine-like setae on proximal third; left mandible with incisors cleft (Fig. 59C); segment III of labial palpi pointed (Fig. 59G) .................................................................................................. Rivudiva 19(17) Gills present on abdominal segments II-VII ............................................................. 20 – Gills present on abdominal segments I-VII ............................................................... 21 20(19) Segment II of labial palpi with distomedial projection (Fig. 9F); mandibles with setae between prostheca and mola (Figs. 9B, C) ..................................... Americabaetis – Segment II of labial palpi not as above (Fig. 69F); mandibles without setae between prostheca and mola (Figs. 69B-C) ......................................................... Zelusia 21(19) Hind wing pads present .................................................................................................. 22 – Hind wing pads absent .................................................................................................... 25 22(21) Labrum with dorsal subapical row of flattened setae that are basally broad and apically pointed (Figs. 19A-B); maxillary palpi three-segmented (Fig. 19F) ............ ................................................................................................................................... Aturbina – Labrum without dorsal subapical row of flattened setae (Figs. 53A-B); maxillary palpi two-segmented (Fig. 53E) .................................................................................... 23 23(22) Tarsal claws with 2 rows of denticles (Fig. 63I) ............................................... Tupiara – Tarsal claws with 1 row of denticles (Fig. 53J) .......................................................... 24 24(23) Mandibles with setae between prostheca and mola ..........................Fallceon (in part) – Mandibles without setae between prostheca and mola (Figs. 53C, D) ....... Nanomis 25(21) Thoracic mesosternum with a projection (Fig. 68Q); prostheca of right mandible bifid (Fig. 68F) ................................................................................................. Waltzoyphius – Thoracic mesosternum without projection; prostheca of right mandible robust with denticles apically and with a transverse seta (Fig. 46E) .................................. 26 26(25) Mandibles with incisors fused basally (Figs. 46C-D); labrum with a small central lobe on anterior margin (Fig. 46A) ............................................................ Cryptonympha – Mandible with incisors cleft in two sets, inner set of incisors positioned at right angle to outer set (Figs. 7B-C); labrum with a small central lobe on anterior margin and one large and one small lobes on both sides (Fig. 7A) ........................ Adebrotus

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SUDAMERICANOS

Adultos 1 – 2(1) – 3(2) – 4(3) – 5(4) – 6(5) – 7(5) – 8(7) – 9(7) – 10(9) – 11(4) – 12(11) – 13(12) – 14(13) – 15(14) –

Ojos compuestos del macho no turbinados; fórceps con el segmento I más grande que el segmento II (Figs. 18D, H) ..................................................................... Aturbina Ojos del macho turbinados (Figs. 20A, D, 40A, 64A); fórceps con el segmento I más corto que el segmento II (Fig. 62D) ...................................................................... 2 Cuerpo con pequeñas manchas castaño-rojizas ........................................... Callibaetis Cuerpo sin tales manchas ................................................................................................. 3 Alas anteriores con manchas ............................................................................ Moribaetis Alas anteriores sin manchas (Fig. 62B) .......................................................................... 4 Alas posteriores ausentes .................................................................................................. 5 Alas posteriores presentes .............................................................................................. 11 Segmento I de los fórceps con una proyección lateral (Fig. 20C) ........................... 6 Segmento I de los fórceps sin proyección (Fig. 47B) ................................................. 7 Margen postero-lateral de la placa estilígera con una pequeña proyección (Fig. 64C) ............................................................................................................................. Varipes Margen postero-lateral de la placa estilígera sin proyección (Fig. 20C) ...... Baetodes Segmento II de los fórceps con una proyección distomedial (Fig. 47B) ............... 8 Segmento II de los fórceps sin proyección .................................................................. 9 Margen apical de la placa estilígera con una pequeña proyección distomedial(Fig. 47B); individuos grandes (5,0 a 5,5 mm) ....................................................... Guajirolus Margen apical de la placa estilígera sin proyección (Fig. 40C); individuos pequeños (3,5 a 4,0 mm) ............................................................................................................. Chane Margen apical de la placa estilígera con una fuerte proyección subcuadrangular (Fig. 16A) ............................................................................................................... Apobaetis Margen apical de la placa estilígera sin proyección (Fig. 42D) .............................. 10 Fórceps con segmento III corto y redondeado (Fig. 42D) ........ Cloeodes (en parte) Fórceps con segmento III alargado (Fig. 8B) .......................................... Americabaetis Alas posteriores con 3 venas longitudinales, la segunda bifurcada (Fig. 11B) ........ .................................................................................................................................. Andesiops Alas posteriores con 2 o 3 venas longitudinales, la segunda nunca bifurcada (Fig. 30C) ..................................................................................................................................... 12 Fórceps aparentemente tetra-segmentados ........................................................ Fallceon Fórceps tri-segmentados (Fig. 54C) ............................................................................. 13 Proyección costal del ala posterior subcuadrangular (Fig. 54F) .............. Paracloeodes Proyección costal del ala posterior aguda (Fig. 30C) ............................................... 14 Fórceps con el segmento III corto y redondeado (Fig. 42D) ................................ 15 Fórceps con el segmento III alargado (Fig. 30E) ..................................................... 16 Base de los fórceps muy próximas entre sí, segmento I sin proyección (Fig. 42D); ala anterior con vena A1 no fuertemente curvada (Fig. 42A) ..... Cloeodes (en parte) Base de los fórceps separadas entre sí, segmento I con una pequeña proyección distomedial (Fig. 51D); ala anterior con vena A1 fuertemente curvada (Fig. 51A) ................................................................................................................................ Mayobaetis

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16(14) Ala anterior con la región anal redondeada (Fig. 30A) .......................Camelobaetidius – Ala anterior con la región anal relativamente recta (Fig. 62B) ...................... Tupiara Ninfas 1 – 2(1) – 3(2) – 4(3) – 5(4)

Uñas tarsales espatuladas, con 5 a 40 dentículos (Fig. 32I) ...............Camelobaetidius Uñas tarsales aguzadas hacia el ápice (Fig. 19H) ......................................................... 2 Branquias en los segmentos I-V solamente ....................................................... Baetodes Branquias en los segmentos abdominales I-VI, I-VII ó II-VII ............................... 3 Branquias abdominales con dos láminas (Figs. 29M-O) ............................ Callibaetis Branquias abdominales con una lámina (Fig. 9J) ........................................................ 4 Uñas tarsales sin dentículos (Fig. 44J) ........................................................................... 5 Uñas tarsales con dentículos, aunque pueden ser pequeños (Fig. 19H) ................ 6 Uñas tarsales largas (0,75 a 0,80 veces la longitud del tarso) (Fig. 16I); tibia sin un arco de setas largas (Fig. 16I) ........................................................................... Apobaetis – Uñas tarsales cortas (menos de 0,6 veces la longitud del tarso) (Fig. 44J) tibia con un arco de setas largas (Fig. 44H) .................................................. Cloeodes/ Bernerius 3 6(4) Uñas tarsales más de la mitad de la longitud del tarso (Fig. 55H) ............................ ............................................................................................................. Iguaira/ Paracloeodes 4 – Uñas tarsales menos de la mitad de la longitud del tarso (Fig. 49N) ..................... 7 7(6) Mandíbulas largas y angostas con región molar muy angosta y protuyente (Figs. 49D-G); caninos maxilares fuertemente desarrollados, el segundo la mitad de tamaño que los restantes (tipo carnívoro) (Fig. 49K) ........................... Harpagobaetis – Mandíbulas más bien triangulares, con región molar ancha y no protuyente (Figs. 63CD); caninos maxilares normales y de igual longitud (tipo herbívoro) (Fig. 63F) ............ 8 8(7) Dentículo subapical de las uñas tarsales más grande que los restantes (Fig. 48O); prosteca mandibular insertada en una concavidad (Figs. 48B-C) ............................ 9 – Dentículo subapical de las uñas tarsales subigual a los otros (Fig.13L); prosteca mandibular no insertada en una concavidad (Figs. 13C, E) ................................... 10 9(8) Segmento I del palpo labial nunca transversal a las glosas y paraglosas, segmento II y III sin hilera de setas largas (Fig. 48F); glosas dorsalmente con una hilera transversal de setas fuertes (Fig. 48F) ............................................................. Guajirolus – Segmento I del palpo labial transversal a las glosas y paraglosas, segmento II y III con dos hileras de setas largas (Fig. 41H); glosas sin hilera transversal de setas (Fig. 41H) .............................................................................................................................. Chane 10(8) Uñas tarsales con 1 o 2 setas fuertes en la base del dentículo subapical del margen externo (Figs. 21K, 52I, 57J) ......................................................................................... 11 – Uñas tarsales sin tales setas (Fig. 9H) o bien con un par de setas débiles apicales (Fig. 13L) ............................................................................................................................ 14 11(10) Uñas tarsales con 2 hileras de dentículos (Fig. 60I) ...................................... Spiritiops – Uñas tarsales con 1 hilera de dentículos (Fig. 21K) ................................................. 12 3 4

Cloeodes y Bernerius están en revisión y existen dudas respecto a la validez de éste último. Paracloeodes e Iguaira están en revisión y existen dudas respecto a la validez de éste último, ambos géneros parecen indistinguibles.

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12(11) Escapo y pedicelo aplanados dorsoventralmente; escapo 3 veces la longitud del pedicelo (Fig. 52A); cercos con una hilera de espinas en el margen externo (Fig. 52J) ................................................................................................................................ Mayobaetis – Escapo y pedicelo cilíndricos o subcilíndricos; escapo 1,0 a 1,5 veces la longitud del pedicelo; cercos sin espinas en el margen externo ............................................. 13 13(12) Pterotecas posteriores si presentes rudimentarias; filamento terminal < 0,25 de la longitud de los cercos; paraproctos con sus márgenes posteriores lisos Prebaetodes – Pterotecas posteriores bien desarrolladas; filamento terminal 0,5 de la longitud de los cercos; paraproctos con sus márgenes posteriores terminados en puntas .... ................................................................................................................................. Moribaetis 14(10) Branquias presentes en los segmentos abdominales I-VI; labro trapezoidal (Fig. 61A) con márgenes laterales divergentes desde la base ............................. Tomedontus – Branquias presentes en los segmentos abdominales I ó II-VII; labro variado, pero no como arriba ................................................................................................................. 15 15(14) Uñas tarsales con un par de setas débiles (Fig. 13L) ................................................ 16 – Uñas tarsales sin tales setas (Fig. 9H) .......................................................................... 17 16(15) Setas simples entre la prosteca y la mola mandibular; uñas tarsales con 1 sola hilera de dentículos .......................................................................................... Fallceon (en parte) – Sin setas entre la prosteca y la mola mandibular (Figs. 13C, E); uñas tarsales con 2 hileras de dentículos aunque la segunda hilera puede estar reducida (Figs. 13L, 14H) .................................................................................................................................. Andesiops 17(15) Fémures con abundantes setas largas y robustas en los márgenes dorsal y ventral (Figs. 65I, K) ..................................................................................................................... 18 – Fémures con o sin setas largas, pero si están presentes no abundantes ni situadas en los márgenes dorsal y ventral (Fig. 9G) ................................................................. 19 18(17) Fémures II y III con una hilera transversa de setas como espinas largas en el tercio proximal (Figs. 65K; 66M; 67H); incisivos de la mandíbula izquierda completamente fusionados (Fig. 65C); segmento III del palpo labial redondeado (Fig. 65H) ................................................................................................................... Varipes – Fémures II y III sin hilera transversal de setas como espinas; incisivos de la mandíbula izquierda completamente divididos (Fig. 59C); segmento III del palpo labial agudo (Fig. 59G) .......................................................................................... Rivudiva 19(17) Branquias presentes en los segmentos abdominales II-VII .................................... 20 – Branquias presentes en los segmentos abdominales I-VII ...................................... 21 20(19) Segmento II del palpo labial con una proyección digitiforme distomedial (Fig. 9F); mandíbulas con setas simples entre la prosteca y la mola (Figs. 9B-C) ................... ............................................................................................................................ Americabaetis – Segmento II del palpo labial no como arriba (Fig. 69F); mandíbulas sin setas entre la prosteca y la mola (Figs. 69B-C) ........................................................................ Zelusia 21(19) Pterotecas posteriores presentes ................................................................................... 22 – Pterotecas posteriores ausentes ..................................................................................... 25 22(21) Labro dorsalmente con una hilera de setas subapicales aplanadas, anchas basalmente y agudas apicalmente (Figs. 19A-B); palpos maxilares tri-segmentado (Fig. 19F) ................................................................................................................. Aturbina

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– 23(22) – 24(23) – 25(21) – 26(25) –

Labro dorsalmente sin setas aplanadas (Figs. 53A-B); palpos maxilares bisegmentados (Fig. 53E) ................................................................................................... 23 Uñas tarsales con 2 hileras de dentículos (Fig. 63I) ........................................ Tupiara Uñas tarsales con 1 hilera de dentículos (Fig. 53J) ................................................... 24 Mandíbulas con setas entre la prosteca y la mola .......................... Fallceon (en parte) Mandíbulas sin setas entre la prosteca y la mola (Figs. 53C-D) .................. Nanomis Mesoesterno torácico con una proyección (Fig. 68Q); prosteca de la mandíbula derecha bífida (Fig. 68F) ................................................................................ Waltzoyphius Mesoesterno torácico sin proyección; prosteca de la mandíbula derecha robusta, con dentículos apicalmente y con una seta transversa (Fig. 46E) ......................... 26 Mandíbulas con incisivos fusionados basalmente (Figs. 46C-D); labro con un pequeño lóbulo central en el margen anterior (Fig. 46A) .................... Cryptonympha Mandíbula con incisivos divididos en dos grupos, incisivos internos ubicados en ángulo recto con respecto a los externos (Figs. 7B-C); labro con un pequeño lóbulo central sobre el margen anterior y un lóbulo grande y uno pequeño a ambos lados (Fig. 7A) ......................................................................................... Adebrotus

Genus Adebrotus LUGO-ORTIZ & MCCAFFERTY Adebrotus LUGO-ORTIZ & MCCAFFERTY, 1995a: 234; ORTH et al., 2000: 34. (Type-species: Adebrotus amazonicus LUGO-ORTIZ & MCCAFFERTY, original designation).

Taxonomy: Adebrotus was established by LUGO-ORTIZ & MCCAFFERTY (1995a). Only one species was included in the genus: A. amazonicus, known from nymphs. Subsequently, ORTH et al. (2000), reported this genus from French Guiana, although they did not specify the species. Adult Characteristics: Unknown. Nymphal Characteristics: 1) Labrum apically with a small central lobe and a large and a small lobes on either side, dorsal surface with 9-10 robust setae at margin on either side of midline (Fig. 7A); 2) left mandible with inner incisors rotated at right angle to outer incisors (Fig. 7B); 3) right mandible with inner incisors rotated to a right angle, prostheca with a transverse branched seta (Fig. 7C); 4) hypopharynx with lingua subequal in length to superlinguae (Fig. 7D); 5) maxillae with palpi two-segmented, 1.5 times length of galea-lacinia (Fig. 7E); 6) labium (Fig. 7F) glossae subequal in length and broader than paraglossae, segment II of palpi elongate without projection, segment III conical; 7) tarsal claws with one row of denticles (Fig. 7H). Distribution: Brazil, Colombia and French Guiana. Adebrotus amazonicus LUGO-ORTIZ & MCCAFFERTY A. amazonicus LUGO-ORTIZ & MCCAFFERTY, 1995a: 235; DOMÍNGUEZ et al., 2002: 462; SALLES et al., 2004: 2.

Taxonomy: Adebrotus amazonicus, known only from nymphs, can be distinguished by the following combination of characters: 1) Mandibles with incisors cleft in two sets and with setae between prostheca and mola (Figs. 7B-C); 2) mola of right mandible with

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SL

TS

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P

L

B

C

III

D II

I

F E

H D

I

G

Plate 7. Adebrotus amazonicus, nymph. Figs. 7A-I. Figs. 7A-F, mouthparts: 7A, labrum d.v.; 7B, left mandible v.v.; 7C, right mandible v.v.; 7D, hypopharynx v.v.; 7E, maxilla v.v.; 7F, labium, left d.v., right v.v. 7G, leg I v.v.; 7H, tarsal claw I; 7I, tergum IV v.d. (D = denticles; I = incisors; L = lingua; P = prostheca; SL = superlingua; TS = transverse seta; Roman numbers = segment numbers). Figs. 7AI modified from LUGO-ORTIZ & MCCAFFERTY, 1995a.

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bipectinate setae (Fig. 7C); 4) dorsal edge of femora with a row of short spines (Fig. 7G); 5) posterior margin of terga with irregular spines as in Fig. 7I. Distribution: Brazil. Genus Americabaetis KLUGE Baetis (Americabaetis) KLUGE, 1992b: 18. Genus 1 nr Pseudocloeon ROBACK, 1966: 134. Acerpenna WALTZ & MCCAFFERTY, in part: LUGO-ORTIZ & MCCAFFERTY, 1994: 66. Americabaetis LUGO-ORTIZ & MCCAFFERTY, 1996a: 157; LUGO-ORTIZ & MCCAFFERTY, 1999a: 260. (Type-species: Baetis naranjoi KLUGE, original designation).

Taxonomy: Americabaetis was originally established as a subgenus of Baetis by KLUGE (1992b), including 2 species from Cuba. LUGO-ORTIZ & MCCAFFERTY (1994) synonymized this subgenus with Acerpenna WALTZ & MCCAFFERTY (1987d). LUGO-ORTIZ & MCCAFFERTY (1996a), after examining material from South America, raised Americabaetis to generic status. All species distributed in Central and South America were transferred to this genus while the species distributed in North America remained in Acerpenna. In South America are 11 species described, five known from adults and four from nymphs alone, one from nymph and female imago and one from nymph and adults. However, as the species described from adults cannot be reliably distinguished based on the known characters, we preferred not to include a key. Adult Characteristics: 1) Eyes of male conical; 2) fore wings with paired marginal intercalaries (Fig. 8A); 3) hind wings absent; 4) genitalia: forceps three-segmented, segment III elongate (Fig. 8B). Nymphal Characteristics: 1) Head longer than wide; 2) frontal keel absent or present; 3) labrum (Fig. 9A), dorsally with 1 pair of subapical setae near midline, apical margin with two kinds of setae: bifid near midline and branched near lateral margin; 4) mandibles (Figs. 9B-C)

A

IV IV

B Plate 8. Americabaetis alphus, male imago. Figs. 8A-B. 8A, fore wing; 8B, genitalia. (IV = intercalary veins).

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with setae between prostheca and mola, incisors fused apically, prosthecae robust with denticles apically; 5) hypopharynx (Fig. 9D) with lingua subequal to superlinguae; 6) maxillae (Fig. 9E) with palpi two-segmented; 7) labium (Fig. 9F): glossae 0.6-0.8 times length and half the width of paraglossae, glossae with a row of long spine-like setae, paraglossae rectangular with 3 rows of long spine-like setae, segment II of palpi with a distomedial projection; 8) legs robust (Fig. 9G), dorsal edge of femora with a row of spines, tibiae 1.4-1.5 times length of tarsi; 9) tarsal claws with a single row of denticles (Fig. 9H); 10) hind wing pads absent; 11) abdominal gills (Fig. 9J) presents on segment II-VII and elongate; 12) posterior margin of paraprocts with spines (Fig. 9K); 13) terminal filament 0.8-1.0 times length of cerci. Distribution: Argentina, Bolivia, Brazil, Chile, Colombia, Ecuador, French Guiana, Paraguay, Peru and Uruguay. KEY

TO

SOUTH AMERICAN AMERICABAETIS

Nymphs 1 – 2(1)

Frontal keel absent ............................................................................................................ 2 Frontal keel present ........................................................................................................... 4 Thorax with paired setose protuberances on meso- and metasterna (Fig. 10K); paraglossae of labium with three rows of simple setae (Fig. 10J) ............................ ............................................................................................................... Americabaetis titthion Thorax without protuberances; paraglossae of labium with three rows of pectinate setae (Fig. 10B) .................................................................................................. 3 Glossae 0.6 times length of paraglossae (Fig. 10B); posterior margin of abdominal terga with rounded spines (Fig. 10D) .......................................... Americabaetis labiosus Glossae 0.8 to 0.9 times length of paraglossae; posterior margin of abdominal terga with pointed spines (similar to Fig. 9I) .............................. Americabaetis robacki Abdominal gills long, 2.5 times length of each segment (Fig. 10H) ......................... ........................................................................................................ Americabaetis maxifolium Abdominal gills short, 1.0 to 1.5 times length of each segment ............................. 5 Sexual dimorphism present in nymphal stage with different color patterns between male and female nymphs; lingua pointed apically (Fig. 9D) ...................... ................................................................................................................ Americabaetis alphus Without sexual dimorphism in nymphal stage; lingua rounded apically (Fig. 10E) .............................................................................................................Americabaetis longetron

– 3(2) – 4(1) – 5(4) –

CLAVE

PARA LOS

AMERICABAETIS

SUDAMERICANOS

Ninfas 1 – 2(1) –

Quilla frontal ausente ........................................................................................................ 2 Quilla frontal presente ...................................................................................................... 4 Tórax con pares de tubérculos setosos en el meso y metaesterno (Fig. 10K); paraglosas del labio con tres hileras de setas simples (Fig. 10J) ...................... Americabaetis titthion Tórax sin tubérculos; paraglosas del labio con tres hileras de setas pectinadas (Fig. 10B) ............................................................................................................................. 3

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P

A

S

L

E

P S

SL

C

B D

III

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F I

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H J D

K

Plate 9. Americabaetis alphus, nymph. Figs. 9A-K. Figs. 9A-F, mouthparts: 9A, labrum, left d.v., right v.v.; 9B, left mandible v.v.; 9C, right mandible v.v.; 9D, hypopharynx v.v.; 9E, maxilla v.v.; 9F, labium, left d.v., right v.v. 9G, leg I; 9H, tarsal claw I; 9I, posterior margin of abdominal tergum IV; 9J, gill IV; 9K, paraproct. (D = denticles; L = lingua; P = prostheca; S = setae; SL = superlingua; Roman numbers = segment numbers).

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Glosas 0,6 la longitud de las paraglosas (Fig. 10B); márgenes posteriores de los tergos abdominales con espinas redondeadas (Fig. 10D) ....... Americabaetis labiosus Glosas 0,8 a 0,9 la longitud de las paraglosas; márgenes posteriores de los tergos abdominales con espinas (como en la Fig. 9I) ............................ Americabaetis robacki Branquias abdominales largas, 2,5 la longitud de cada segmento (Fig. 10H) ......... ........................................................................................................ Americabaetis maxifolium Branquias abdominales cortas, 1,0 a 1,5 la longitud de cada segmento ................. 5 Dimorfismo sexual en las ninfas con diferentes patrones de coloración abdominal entre machos y hembras; lingua (Fig. 9D) aguda apicalmente ...... Americabaetis alphus Sin dimorfismo sexual en las ninfas; lingua (Fig. 10E) redondeada apicalmente .............................................................................................................Americabaetis longetron

Americabaetis alphus LUGO-ORTIZ & MCCAFFERTY Americabaetis alphus LUGO-ORTIZ & MCCAFFERTY, 1996a: 163; SALLES et al., 2004: 2.

Taxonomy: This species, known from nymphs and adults, can be distinguished from the other species of the genus by the following combination of characters. In adults: 1) Compound eyes oval, not contiguous apically. In nymphs: 1) frontal keel present; 2) lingua (Fig. 9D) pointed apically; 3) labium (Fig. 9F) with glossae 0.8 times length of paraglossae, segment II of palpi with a strong distomedial projection; 4) tarsal claw with 14-16 denticles (Fig. 9H); 5) posterior margin of abdominal terga with pointed spines (Fig. 9I). This species is sexually dichromatic in the nymphs, with the abdomen of males having a characteristic color pattern of tergum I white, tergum IV with posterolateral white spots and tergum VII with posterolateral marks, females have white markings laterally on abdominal terga II-VIII. Distribution and Biology: Argentina, Bolivia, Brazil, Chile and Paraguay. This species was collected in many different kinds of rivers, especially in the mountain rain forests from Argentina and Bolivia. Americabaetis bridarolli (NAVÁS) Pseudocloeon bridarolli NAVÁS, 1933d: 113. Americabaetis bridarolli; LUGO-ORTIZ & MCCAFFERTY (1999a): 260.

Taxonomy: This species, known from adults, was originally placed in Pseudocloeon. LUGO-ORTIZ & MCCAFFERTY (1999a) transferred this species to Americabaetis, based on the absence of hind wings, body size and collection data. However, presently known characteristics are not sufficient to distinguish this species from the others. Distribution: Argentina. Americabaetis jorgenseni (ESBEN-PETERSEN) Cloeon jorgenseni ESBEN-PETERSEN, 1909: 551. Pseudocloeon jorgenseni; ULMER, 1920c: 125. Americabaetis jorgenseni; LUGO-ORTIZ & MCCAFFERTY, 1999a: 260.

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Taxonomy: This species was originally described from male and female imagos by ESBENPETERSEN (1909). LUGO-ORTIZ & MCCAFFERTY (1999a) transferred this species to Americabaetis based on the absence of hind wings, body size range and the figure of the male genitalia provided by ESBEN-PETERSEN (1909: Fig. 1). However, although this species could belong to the genus, these characteristics are not enough to distinguish this species from the others. Distribution: Argentina. Americabaetis labiosus LUGO-ORTIZ & MCCAFFERTY Americabaetis labiosus LUGO-ORTIZ & MCCAFFERTY, 1996a: 164. Taxonomy: This species, known from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Frontal keel absent; 2) lingua (Fig. 10A) broadly pointed apically; 3) labium (Fig. 10B) with glossae 0.6 times length of paraglossae, paraglossae with 3 rows of pectinate setae, segment II of palpi with a small distomedial projection; 4) tarsal claw with 10-11 denticles (Fig. 10C); 5) posterior margin of abdominal terga with rounded spines (Fig. 10D). Distribution: Brazil, Paraguay, Uruguay. Americabaetis longetron LUGO-ORTIZ & MCCAFFERTY Americabaetis longetron LUGO-ORTIZ & MCCAFFERTY, 1996a: 165.

Taxonomy: This species, known from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Frontal keel present; 2) lingua (Fig. 10E) rounded apically; 3) labium (Fig. 10F) with glossae 0.8 times length of paraglossae, segment II of palpi with a strong distomedial projection; 4) tarsal claw with 12-14 denticles; 5) posterior margin of abdominal terga with spines; 6) abdomen elongate. Distribution: Brazil, Paraguay and Uruguay. Americabaetis maxifolium LUGO-ORTIZ & MCCAFFERTY Americabaetis maxifolium LUGO-ORTIZ & MCCAFFERTY, 1996a: 166.

Taxonomy: This species, known from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Frontal keel present; 2) lingua rounded apically; 3) labium (Fig. 10G) with glossae 0.8 times length of paraglossae, segment II of palpi with a strong distomedial projection; 4) tarsal claw with 14-16 denticles; 5) posterior margin of abdominal terga with spines; 6) abdominal gills extremely long, 2.5 times length of each segment (Fig. 10H). Distribution: Paraguay. Americabaetis oldendorffi (WEYENBERGH) Oxycypha oldendorffii WEYENBERGH, 1883: 173. Cloeon oldendorffii; EATON, 1885: 191. Baetis oldendorffi; BANKS, 1913: 85.

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B

SL

71

III II

D

A C

I

D L

SL

III II

E III

I II

F

I

G

I

H III II

I II

L SP

I

J

III

K

SP

Plate 10. Americabaetis, nymphs. Figs. 10A-D, A. labiosus: 10A, hypopharynx v.v.; 10B, labium, left d.v., right v.v.; 10C, tarsal claw I; 10D, posterior margin of abdominal tergum IV. Figs. 10E-F, A. longetron: 10E, hypopharynx v.v.; 10F, labium, left d.v., right v.v. Figs. 10G-H, A. maxifolium: 10G, labium, left d.v., right v.v.; 10H, gill IV. Fig. 10I, A. robacki: paraproct. Figs. 10J-L, A. titthion: 10J, labium, left d.v., right v.v.; 10K, thorax: sterna I-III; 10L, posterior margin of abdominal tergum IV. (D = denticles; L = lingua; SL = superlingua; SP = setose protuberance; Roman numbers = segment numbers). Figs. 10E-L modified from LUGO-ORTIZ & MCCAFFERTY, 1996a.

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Pseudocloeon oldendorffii; ULMER, 1920c: 125; NEEDHAM & MURPHY, 1924: 57. Americabaetis oldendorffii; LUGO-ORTIZ & MCCAFFERTY, 1999a: 261; SALLES et al., 2004b: 24.

Taxonomy: This species, known from adults, was originally described by WEYEN(1883) and placed in the genus Oxycypha. This species underwent several shift into different genera until LUGO-ORTIZ & MCCAFFERTY (1999a) transferred it to Americabaetis, based on the absence of hind wings, body size and geographic distribution. Later, SALLES et al. (2004) notice that the specimen identified as Baetis oldendorffi by BANKS (1913) could not belong to this genus because it has hind wing. For this reason this species would not be in Brazil. The original description does not provide specific characteristics to distinguish this species from others in the genus. Distribution: Argentina. BERGH

Americabaetis peterseni (HUBBARD) Cloeon brunneum ESBEN-PETERSEN, 1909: 552, nec C. brunneum RAMBUR, 1842. Pseudocloeon brunneum; ULMER, 1920c: 125. Cloeon peterseni HUBBARD, 1974: 257. Pseudocloeon peterseni; HUBBARD et al., 1992: 204. Americabaetis peterseni; LUGO-ORTIZ & MCCAFFERTY, 1999a: 261.

Taxonomy: This species, known from adults, was originally described by ESBENPETERSEN (1909). LUGO-ORTIZ & MCCAFFERTY (1999a) transferred this species to Americabaetis based on the absence of hind wings, body size, collection data and the generically diagnostic figure of the male genitalia (ESBEN-PETERSEN, 1909: Fig. 3). However, the original description does not provide specific characteristics to distinguish this species from others in the genus. Distribution: Argentina. Americabaetis robacki (LUGO-ORTIZ & MCCAFFERTY) Genus 1 nr Pseudocloeon sp. 5 ROBACK, 1966: 135. Acerpenna robacki LUGO-ORTIZ & MCCAFFERTY, 1994: 73. Americabaetis robacki; LUGO-ORTIZ & MCCAFFERTY, 1996a: 167; DOMÍNGUEZ et al., 2002: 462.

Taxonomy: This species, known from a single nymph, can be distinguished from other species of the genus by the following combination of characters: 1) Frontal keel absent; 2) labium with glossae 0.8-0.9 times length of paraglossae, paraglossae with 3 rows of pectinate setae, segment II of palpi with a thumb-like projection; 4) tarsal claw with 12-13 denticles; 5) posterior margin of abdominal terga with spines; 6) paraprocts with elongate sharp spines (Fig. 10I). Distribution: Colombia and Peru. Americabaetis titthion LUGO-ORTIZ & MCCAFFERTY Americabaetis titthion LUGO-ORTIZ & MCCAFFERTY, 1996a: 167; SALLES et al., 2004b: 23.

Taxonomy: This species, known from nymphs and female imago, can be distinguished from the other species of the genus by the following combination of characters. In female imago, 1) paired setose protuberances on meso- and metasterna; 2) posterior margin of head

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projected. In nymphs, 1) Frontal keel absent; 2) lingua broadly pointed apically (similar to Fig. 10A); 3) labium (Fig. 10J) with glossae 0.8 times length of paraglossae, segment II of palpi with a moderate distomedial projection; 4) tarsal claw with 10-12 denticles; 5) posterior margin of abdominal terga with rounded spines (Fig. 10L); 6) paired setose protuberances on meso- and metasterna (Fig. 10K); 7) abdominal gills with simple marginal setae. Distribution: Brazil. Americabaetis weiseri (NAVÁS) Pseudocloeon weiseri NAVÁS, 1926b: 34. Americabaetis weiseri; LUGO-ORTIZ & MCCAFFERTY, 1999a: 261.

Taxonomy: This species was originally described from a female imago by NAVÁS (1926b). LUGO-ORTIZ & MCCAFFERTY (1999a) transferred this species to Americabaetis based on the absence of hind wings, body size, collection data, and the close similarity to females of other species of Americabaetis. Although this species could belong to the genus, LUGO-ORTIZ & MCCAFFERTY suggested that because the species description was based on females, any future assignments to this species would be difficult. Distribution: Argentina. Americabaetis albinervis (NAVÁS) Nomen Dubium Pseudocloeon albinerve NAVÁS, 1923a: 443. Americabaetis albinervis; LUGO-ORTIZ & MCCAFFERTY, 1999a: 260; MCCAFFERTY, 2000b: 378.

Taxonomy: This species, known from adults, was originally placed in the genus Pseudocloeon (NAVÁS, 1923a). LUGO-ORTIZ & MCCAFFERTY (1999a) transferred this species to Americabaetis. However, these authors suggested that although this species belongs in Americabaetis, the poor condition of the type material, as well as the brief and inconsequential original description, make it impossible to assign any new material to this species in the future. Finally, MCCAFFERTY (2000b) proposed this species as nomen dubium. Distribution: Chile. Americabaetis bruchi (NAVÁS) Nomen Dubium Pseudocloeon bruchi NAVÁS, 1926b: 34. Americabaetis bruchi; LUGO-ORTIZ & MCCAFFERTY, 1999a: 260; MCCAFFERTY, 2000b: 378.

Taxonomy: This species, known from adults, was originally placed in the genus Pseudocloeon (NAVÁS, 1926b). LUGO-ORTIZ & MCCAFFERTY (1999a) transferred this species to Americabaetis based on the absence of hind wings, body size, collection data and the statement by NAVÁS (1926b) that this species was similar to P. jorgenseni, a species clearly assignable to Americabaetis. However the authors suggested that although this species belongs to Americabaetis, the poor condition of the type material, as well as the incomplete original description, make it impossible to assign any new material to this species in the future. Finally, MCCAFFERTY (2000b) proposed this species as nomen dubium. Distribution: Argentina.

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Genus Andesiops LUGO-ORTIZ & MCCAFFERTY Andesiops LUGO-ORTIZ & MCCAFFERTY, 1999c: 89; NIETO, 2004c: 95. (Type-species: Baetis peruvianus ULMER, original designation). Deceptiviosa LUGO-ORTIZ & MCCAFFERTY, 1999c: 93. (Type-species: Deceptiviosa torrens LUGO-ORTIZ & MCCAFFERTY, original designation).

Taxonomy: Andesiops was established by LUGO-ORTIZ & MCCAFFERTY (1999c). A single species, A. peruvianus (ULMER, 1920a) was included in the genus. NIETO (2004c) synonymized Deceptiviosa with this genus, transferring the two species originally placed in Deceptiviosa to Andesiops. Another species, also transferred to Deceptiviosa (MCCAFFERTY, 2000b), was omitted by NIETO (2004c) and is discussed below. Adult Characteristics: 1) Hind wings with a small, sharp-pointed costal process located in basal 1/3 (Figs. 11B, E); 2) hind wings with 3 longitudinal veins, the middle vein forked (Figs. 11B, E); 3) segment III of forceps elongate (Figs. 11C, F). Nymphal Characteristics (Figs. 12A-B): 1) Labrum (Fig. 13A) wider than long, with anteromedial margin cleft with a small central lobe, dorsal surface with two kinds of bipectinate setae on apical margin: basally bifid near midline and apically bifid near lateral

A

IV

IV

B

C

IV

D

IV

E

F

Plate 11. Andesiops, imagos. Figs. 11A-C, A. peruvianus: 11A, fore wing; 11B, hind hing; 11C, male genitalia, v.v. Figs. 11D-F, A. torrens: 11D, fore wing; 11E, hind hing; 11F, male genitalia, v.v. (IV = Intercalary veins). Figs. 11A-F modified from NIETO, 2004c.

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margin (Fig. 13B); with 1 pair of subapical simple setae medially and a row of 2-8 subapical setae laterally; 2) left mandible (Figs. 13C, 14A) with prostheca robust with denticles apically; 3) right mandible with prostheca bifid and with incisors cleft apically (Fig. 13E); 4) maxillae with palpi shorter than (Fig. 13G) or as long as galea-lacinia (Fig. 13J); 5) segment II of labial palpi with a small distomedial projection (Figs. 13H, K); 6) dorsal margin of femora and tibiae with a row of long setae, tibiae twice length of tarsi (Figs. 14C, G); 7) tarsal claws with 2 rows of denticles, first row well developed, second row reduced in size or in number of denticles, with a subapical pair of fine setae (Figs. 13L, 14D, H); 8) gills without tracheae or weakly tracheated. Distribution: Argentina, Bolivia, Chile, Colombia, Ecuador, Peru and Venezuela. Biology: This genus was collected in high altitude rivers and streams with rocky substrate and highly oxygenated water.

A

B

Plate 12. Andesiops sp., nymphal habitus. Figs. 12A-B. 12A, dorsal view; 12B, lateral view. Drawings by C. Molineri.

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KEY

TO

SOUTH AMERICAN ANDESIOPS

Nymphs 1 – 2(1) –

CLAVE

Head broader than long; antennae short (1.5 times length of head capsule); terminal filament 0.15 to 0.20 times length of cerci; tarsal claws with only a single denticle in the second row (Fig. 14H) ................................................. Andesiops torrens Head longer than broad; antennae long (3 times of head capsule); terminal filament 0.50 to 0.70 times length of cerci; tarsal claws with at least 4 or 5 denticles in the second row (Figs. 13L, 14D) .............................................................. 2 Second row of denticles of tarsal claws with nearly as many denticles as the first row (Fig. 14D); lingua apically with a pointed projection (Fig. 14B); terminal filament 0.65 to 0.70 times length of cerci (mature nymph) ...................... Andesiops peruvianus Second row of denticles of tarsal claws with about half as many denticles as the first row (Fig. 13L); lingua apically rounded (Fig. 13I); terminal filament 0.50 to 0.60 times length of cerci (mature nymph) ..........................................Andesiops ardua PARA LOS

ANDESIOPS

SUDAMERICANOS

Ninfa 1 – 2(1)

–

Cabeza más ancha que larga; antenas cortas (1,5 veces el ancho de la cabeza); filamento terminal 0,15 a 0,20 veces la longitud de los cercos; uñas tarsales con un solo dentículo en la segunda hilera (Fig. 14H) .................................. Andesiops torrens Cabeza más larga que ancha; antenas largas (3 veces el ancho de la cabeza); filamento terminal 0,50 a 0,70 veces la longitud de los cercos; uñas tarsales con la segunda hilera de dentículos con por lo menos 4 o 5 dentículos (Figs. 13L, 14D) .............................. 2 Segunda hilera de dentículos de las uñas tarsales con aproximadamente la misma cantidad de dentículos que la primera (Fig. 14D); lingua apicalmente terminada en una proyección aguda (Fig. 14B); filamento terminal 0,65 a 0,70 veces la longitud de los cercos (ninfas maduras) ........................................................ Andesiops peruvianus Segunda hilera de dentículos de las uñas tarsales con la mitad de dentículos que la primera (Fig. 13L); lingua apicalmente redondeada (Fig. 13I); filamento terminal 0,50 a 0,60 veces la longitud de los cercos (ninfa madura) ...............Andesiops ardua

Andesiops angolinus (NAVÁS) n. comb. Baetis angolinus NAVÁS, 1933a: 230. Deceptiviosa angolina; MCCAFFERTY, 2000b: 377.

Taxonomy: This species was originally described from a male subimago. MCCAFFERTY (2000b) examined the evidence and transferred it to Deceptiviosa which is a synonym of Andesiops and for that reason this species is transferred. However, the original description is not sufficient to distinguish this species from the others. Because the nymph is unknown this species is not included in the key. This species has: 1) Fore wings with long marginal intercalaries and intercostal area with six crossveins; 2) hind wings that are small and narrow, without cross veins. Distribution: Chile.

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D T

P

P

B SP

L

F

C

E

III II

I

H

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SP

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III II

L D S

J

I

K

Plate 13. Andesiops, nymphs. Figs. 13A-H, A. torrens, mouthparts: 13A, labrum, left d.v., right v.v.; 13B, labrum: setae on anterior margin; 13C, left mandible v.v.; 13D, left mandible: molars, enlarged; 13E, right mandible v.v.; 13F, hypopharynx v.v.; 13G, maxilla v.v.; 13H, labium, left d.v., right v.v. Figs. 13I-L, A. ardua: 13I, hypopharynx v.v.; 13J, maxilla; 13K, labium, left d.v., right v.v.; 13L, tarsal claw I. (D = denticles; L = lingua; P = prostheca; S = subapical setae; SP = superlingua; T = tooth; Roman numbers = segment numbers). Figs. 13A-L modified from NIETO, 2004c.

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Andesiops ardua (LUGO-ORTIZ & MCCAFFERTY) Deceptiviosa ardua LUGO-ORTIZ & MCCAFFERTY, 1999c: 95. Andesiops ardua; NIETO, 2004c: 99.

Taxonomy: This species, known from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Head longer than broad; 2) antennae long, 3 times of head capsule; 3) left mandible: teeth of molar without constrictions (as in Fig. 14A); 4) hypopharynx with lingua apically rounded (Fig. 13I); 5) labium not compact (Fig. 13K); 6) tarsal claws with second row of denticles with half as many denticles as first row (Fig. 13L); 7) posterior margins of abdominal terga with small spines (as in Fig. 14E); 8) paraprocts with posterior margins with large spines (as in Fig. 14F); 9) terminal filament 0.50-0.60 times length of cerci. Distribution: Chile. Andesiops peruvianus (ULMER) Baetis peruvianus ULMER, 1920a: 53. Baetis weiseri NAVÁS, 1922c: 385. Andesiops peruvianus; LUGO-ORTIZ & MCCAFFERTY, 1999c: 91; NIETO, 2004c: 99.

Taxonomy: This species, known from nymphs and adults, can be distinguished from the other species of the genus by the following combination of characters. In the male imago, 1) Turbinate eyes ellipsoidal, height of stalk half of eye diameter; 2) fore wings with paired marginal intercalaries (Fig. 11A); 3) fore femur 0.70 times length of tibia; tarsus subequal to tibia. In the nymph, 1) Head longer than broad; 2) antennae long, three times of head capsule; 3) left mandible: teeth of molar without constrictions (Fig. 14A); 4) lingua with a pointed projection apically (Fig. 14B); 5) labium not compact (as in Fig. 13K); 6) tarsal claws with second row of denticles with nearly as many denticles as the first row, but reduced in size (Fig. 14D); 7) posterior margins of abdominal terga with small spines (Fig. 14E); 8) paraprocts with posterior margin with large spines (Fig. 14F); 9) terminal filament 0.65-0.70 times length of cerci (mature nymph). Distribution and Biology: Argentina, Colombia and Peru. This species was collected in rivers and streams at altitude between 900 and 3000 meters. Most of the rivers where this species was found belong to the yungas area or the mountain rain forest. The rivers have moderate to fast currents and rocky substrates. Andesiops torrens (LUGO-ORTIZ & MCCAFFERTY) Deceptiviosa torrens LUGO-ORTIZ & MCCAFFERTY, 1999c: 98. Andesiops torrens; NIETO, 2004c: 99.

Taxonomy: This species, known from nymphs and adults, can be distinguished from the other species of the genus by the following combination of characters. In the male imago: 1) Turbinate eyes tubular, height of stalk equal to eye diameter; 2) fore wings with marginal intercalaries double or single (Fig. 11D); 3) fore femur 0.50 times length of tibia, tarsus longer than tibia. In the nymph, 1) Head broader than long; 2) antennae short, 1.5 times of head capsule; 3) left mandible: teeth of molar with constrictions (Fig. 13D); 4)

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SL

79

L

B

P

A

C

E

D D S

G

F

I S

J

D

H

Plate 14. Andesiops, nymphs. Figs. 14A-F, A. peruvianus: 14A, left mandible v.v.; 14B, hypopharynx v.v.; 14C, leg I; 14D, tarsal claw I; 14E, posterior margin of tergum IV; 14F, paraproct. Figs. 14G-J, A. torrens: 14G, leg I; 14H, tarsal claw I; 14I, posterior margin of tergum IV; 14J, paraproct. (D = denticles; L = lingua; P = prostheca; S = subapical setae; SP = superlingua). Figs. 14A-J modified from NIETO, 2004c.

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lingua with a rounded projection apically (Fig. 13F); 5) labium compact (Fig. 13H); 6) tarsal claws with a single denticle in second row location (Fig. 14H); 7) posterior margins of abdominal terga with rounded spines (Fig. 14I); 8) paraprocts with posterior margins with small spines (Fig. 14J); 9) terminal filament 0.15-0.20 times length of cerci. Distribution and Biology: Argentina, Chile. This species was collected at high latitude, in streams with fast and cool currents. Genus Apobaetis DAY Apobaetis DAY, 1955: 126; LUGO-ORTIZ & MCCAFFERTY, 1997: 243; MCCAFFERTY, 2000a: 265; SALLES et al., 2004: 2. (Type-species: Apobaetis indeprensus DAY, original designation).

Taxonomy: DAY (1955) described one species, Apobaetis indeprensus, from nymphs and adults collected in North America. In 1986, WALTZ & MCCAFFERTY transferred Pseudocloeon etowah (TRAVER, 1935) to Apobaetis. In 1997, LUGO-ORTIZ & MCCAFFERTY described a new species, A. signifer, from Brazil and Paraguay, thus establishing its presence in South America. Two additional species have been described, A. lakota MCCAFFERTY (2000a) from the United States and A. fiuzai SALLES & LUGO-ORTIZ (2002) from Brazil. Adult Characteristics: 1) Compound eyes of male elliptical; 2) fore wings with paired marginal intercalaries; 3) hind wings absent; 4) genitalia with styliger plate with a subtriangular projection (Fig. 16A). Nymphal Characteristics (Figs. 15A-B): 1) Labrum expanded laterally, anterior margin with a row of long spine-like setae (Fig. 16B); 2) mandibles with incisors deeply cleft and without setae between prostheca and mola (Figs. 16C-D); 3) right mandible with prostheca reduced to a long simple seta (Fig. 16D); 4) hypopharynx: lingua without projection apically and subequal in length to superlingua (Fig. 16E); 5) maxillae with palpi two-segmented, 1-1.5 times length of galea-lacinia (Fig. 16F); 6) labium with glossae and paraglossae subequal in length, glossae wider than paraglossae (Fig. 16H), segment II of palpi with a strong distomedial projection, segment III with 6-9 strong spine-like setae on inner margin (Fig. 16G); 7) tarsal claws long, 0.6-1.3 times length of tarsi and without denticles (Fig. 16I); 8) abdominal gills (Fig. 16J) on segments I-VII, long and slender, about 3 times length of each segment. Distribution: Brazil, Colombia and Paraguay. Biology: Nymphs have been collected from sandy substrate in swift current. The nymphs are relatively tolerant to poor water quality. KEY

TO

SOUTH AMERICAN APOBAETIS

Nymphs 1 –

Maxillary palp at least 1.5 times the length of galea-lacinia; anterior margin of labrum without row of short setae at middle (Figs. 17A-B); tarsal claws 0.65 times length of tarsus ........................................................................................... Apobaetis fiuzai Maxillary palp subequal in length to galea-lacinia; anterior margin of labrum with row of short setae at middle (Fig. 17E); tarsal claws 0.75 times length of tarsus ...................................................................................................................... Apobaetis signifer

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B A

Plate 15. Apobaetis sp. (undescribed). Figs. 15A-B, nymphal habitus: 15A, dorsal view; 15B, ventral view. Drawings by C. Molineri.

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SP

I P

B

A P

S

I

AS

L

C

SL

E D

III

II S I

H

G

F

I

J Plate 16. Apobaetis. Fig. 16A, A. indeprensus, male genitalia. Figs. 16B-J, Apobaetis sp., nymph. Figs. 16B-H, mouthparts: 16B, labrum, left d.v., right v.v.; 16C, left mandible v.v.; 16D, right mandible v.v.; 16E, hypopharynx v.v.; 16F, maxilla v.v.; 16G, labium: palps, left d.v., right v.v.; 16H, labium: glossae and paraglossae, left d.v., right v.v. 16I, leg I; 16J, gill IV. (AS = apical setae; I = incisors; L = lingua; P = prostheca; S = long spines; SL = superlingua; SP = styliger plate; Roman numbers = segment numbers). Fig. 16A modified from DAY, 1955.

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83

SUDAMERICANOS

Ninfas 1 –

Palpo maxilar por lo menos 1,5 veces la longitud de la galea-lacinia; margen anterior del labro sin setas cortas hacia la línea media (Figs. 17A-B); uñas tarsales 0,65 veces la longitud del tarso ............................................................... Apobaetis fiuzai Palpo maxilar subigual en longitud a la galea-lacinia; margen anterior del labro con setas cortas hacia la línea media (Fig. 17E); uñas tarsales 0,75 veces la longitud del tarso ............................................................................................................. Apobaetis signifer

Apobaetis fiuzai SALLES & LUGO-ORTIZ Apobaetis fiuzai SALLES & LUGO-ORTIZ, 2002: 2.

Taxonomy: This species, known from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Anterior margin of labrum without short setae (Figs. 17A-B); 2) palp of maxilla at least 1.5 length of galealacinia; 3) segment III of labial palp as in Fig. 17C; 4) color pattern as in Fig. 17D. Distribution: Brazil. A

III II

I

C

D

B III II I SS

E

F

G

Plate 17. Apobaetis, nymphs. Figs. 17A-D, A. fiuzai: 17A, labrum d.v.; 17B, labrum: apical margin d.v.; 17C, labium, left d.v., right v.v.; 17D, abdomen d.v. Figs. 17E-G, A. signifer: 17E, labrum d.v.; 17F, labium, left v.v., right d.v.; 17G, abdomen d.v. (SS = short setae; Roman numbers = segment numbers). Figs. 17A-D modified from SALLES & LUGO-ORTIZ, 2002; 17E-17G from LUGO-ORTIZ & MCCAFFERTY, 1997.

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Apobaetis signifer LUGO-ORTIZ & MCCAFFERTY Apobaetis signifer LUGO-ORTIZ & MCCAFFERTY, 1997: 245; DOMÍNGUEZ et al., 2002: 462.

Taxonomy: This species, known from nymph, can be distinguished from the other species of the genus by the following combination of characters: 1) Anterior margin of labrum with row of short setae at middle (Fig. 17E); 2) palp of maxilla subequal in length to galea-lacinia; 3) segment III of labial palp as in Fig. 17F; 4) color pattern as in Fig. 17G. Distribution: Brazil, Colombia and Paraguay. Genus Aturbina LUGO-ORTIZ & MCCAFFERTY Aturbina LUGO-ORTIZ & MCCAFFERTY, 1996d: 176; GILLIES, 2001: 331. (Type-species: Aturbina georgei LUGO-ORTIZ & MCCAFFERTY, original designation).

Taxonomy: Aturbina was established by LUGO-ORTIZ & MCCAFFERTY (1996d). They described one species, A. georgei, known from nymphs and adults from Brazil, French Guiana and Paraguay. GILLIES (2001) described a second species, A. beatrixae from Uruguay. Adult Characteristics: 1) Compound eyes of male imago not turbinate, with small facets dorsally and large facets laterally; 2) fore wing with marginal intercalaries paired (Figs. 18A, E); 3) hind wing with 2 longitudinal veins and a truncate costal projection (Figs. 18B-C, F-G); 4) genitalia: basal segment of forceps long, subequal in length to segments II and III combined and with a strong distomedial projection (Figs. 18D, H); 5) styliger plate slightly produced and truncate (Figs. 18D, H). Nymphal Characteristics: 1) Antennae long, 3 times of head capsule; 2) labrum (Figs. 19A-B): dorsal surface with a submarginal row of flattened spines that are basally broad and apically pointed, setae on anterior margin bifid; 2) mandibles with incisors fused apically (Figs. 19C-D); 3) left mandible with prostheca robust with denticles apically (Fig. 19C); 4) prostheca of right mandible with a transverse bipectinate seta (Fig. 19D); 5) hypopharynx with lingua rounded apically (Fig. 19E); 6) maxillae with palpi three-segmented, 1-1.5 times length of galea-lacinia (Fig. 19F); 7) labium: glossae subequal in length and half the width of paraglossae, segment II of labial palpi without projection, segment III apically broad (Fig. 19G); 8) tarsal claws with two rows of small denticles (Fig. 19H); 9) posterior margin of terga with long spines (Fig. 19I). Distribution: Argentina, Brazil, Colombia, French Guiana, Paraguay and Uruguay. Biology: The nymphs were collected from rivers in the middle of the forest. The adults were collected in the spring or summer. KEY

TO

SOUTH AMERICAN ATURBINA

Male Imago 1 –

Antennae short, subequal in length to head capsule; fore wings with a dark spot at junction of costal brace and subcosta (Fig. 18A); segment II of the forceps partially divided (Fig. 18D) ................................................................. Aturbina beatrixae Antennae long, 3 times the length of head capsule; fore wings without a spot at junction of costal brace and subcosta (Fig. 18E); segment II of the forceps without a division (Fig. 18H) ................................................................... Aturbina georgei

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Nymphs 1

Maxillae with palpi subequal in length to galea-lacinia (Fig. 19J); segment III of labial palpi apically rounded (Fig. 19K) ............................................ Aturbina beatrixae Maxillae with palpi 1.5 times the length of galea-lacinia (Fig. 19F); segment III of labial palpi apically truncate (Fig. 19G) ................................................. Aturbina georgei

–

CLAVE

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ATURBINA

SUDAMERICANOS

Machos Imagos 1

Antenas cortas, subigual al ancho de la cabeza; alas anteriores con una mancha oscura en la unión del arco costal y la vena subcosta (Fig. 18A); segundo segmento de los fórceps parcialmente dividido (Fig. 18D) ............................ Aturbina beatrixae Antenas largas, 3 veces el ancho de la cabeza; alas anteriores sin mancha en la unión del arco costal y la vena subcosta (Fig. 18E); segundo segmento de los fórceps no dividido (Fig. 18H) ................................................................ Aturbina georgei

–

S

III II

SP

I

D A IV

IV

CP

C

B

III II

E

SP

I IV

F

CP

G

H

Plate 18. Aturbina, male imagos. Figs. 18A-D, A. beatrixae: 18A, fore wing; 18B, hind wing; 18C, hind wing detail; 18D, genitalia. Figs. 18E-H, A. georgei: 18E, fore wing; 18F, hind wing; 18G, hind wing detail; 18H, genitalia (IV = intercalary veins; CP = costal process; S = spot; SP = Styliger Plate; Roman numbers = segment numbers).

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Ninfas 1 –

Palpo maxilar subigual en longitud a la galea-lacinia (Fig. 19J); segmento III del palpo labial redondeado apicalmente (Fig. 19K) ............................ Aturbina beatrixae Palpo maxilar 1,5 veces el largo de la galea-lacinia (Fig. 19F); segmento III del palpo labial romo apicalmente (Fig. 19G) ............................................ Aturbina georgei

Aturbina beatrixae GILLIES Aturbina beatrixae GILLIES, 2001: 332.

Taxonomy: This species was described by GILLIES (2001) based on nymphs and male and female imagos. This species can be distinguished from the other species of the genus by the following combination of characters. In the adults: 1) Antennae short, subequal in length to head capsule; 2) fore wings with a dark spot at junction of costal brace and subcosta (Fig. 18A); 3) segment II of forceps partially divided (Fig. 18D). In the nymphs, 1) Maxillae with palpi subequal in length to galea-lacinia (Fig. 19J); 2) segment III of labial palpi apically rounded (Fig. 19K). Distribution: Argentina and Uruguay. Specimens were collected from Paranaense area, in rocky substrate rivers. The adults were collected in November. Aturbina georgei LUGO-ORTIZ & MCCAFFERTY Aturbina georgei LUGO-ORTIZ & MCCAFFERTY, 1996d: 179; DOMÍNGUEZ et al., 2002: 462; SALLES et al., 2004: 2.

Taxonomy: This species was described by LUGO-ORTIZ & MCCAFFERTY (1996d) based on nymphs and male and female imagos. This species can be distinguished from the other species of the genus by the following combination of characters. In the adults: 1) Antennae long, 3 times length of head capsule; 2) fore wings without a spot at junction of costal brace and subcosta (Fig. 18E); 3) segment II of forceps not divided (Fig. 18H). In the nymphs, 1) Maxillae with palpi 1.5 times length of galea-lacinia (Fig. 19F); 2) segment II of labial palpi apically truncate (Fig. 19G). Distribution and Biology: Brazil, Colombia, French Guiana and Paraguay. Nymphs were collected from submerged vegetation, in rivers in the middle of the forest with sandy substrate and with high concentrations of tannines. Male imagos were collected in January. Genus Baetodes NEEDHAM & MURPHY Baetodes NEEDHAM & MURPHY, 1924: 55; TRAVER, 1944: 20; EDMUNDS, 1950a: 204; DEMOULIN, 1955a: 20; MAYO, 1968a: 251; 1972: 226; 1973b: 308; COHEN & ALLEN, 1972: 123; 1978: 253, KOSS, 1972: 93; FLOWERS, 1987: 1; MCCAFFERTY & PROVONSHA, 1993: 59; LUGO-ORTIZ & MCCAFFERTY, 1995b: 81, 1996b: 367; NIETO, 2004a: 63. (Type-species: Baetodes serratus NEEDHAM & MURPHY, original designation).

Taxonomy: Baetodes was originally described by NEEDHAM & MURPHY (1924) from South America. The original description was based on nymphs from Brazil. In 1943,

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TRAVER described the adults of the genus from Venezuela. Since then, many authors including EDMUNDS (1950a), DEMOULIN (1955a), MAYO (1968a, 1972 & 1973b), COHEN & ALLEN (1972, 1978), KOSS (1972), FLOWERS (1987), MCCAFFERTY & PROVONSHA (1993), LUGO-ORTIZ & MCCAFFERTY (1995b) and NIETO (2004a) have described new species for P

P TS

A

B L

C

D

SL

III

E II

H

D

I

F

G III II

I J

I

K

Plate 19. Aturbina, nymphs. Figs. 19A-I, A. georgei. Figs. 19A-G, mouthparts: 19A, labrum, left d.v., right v.v.; 19B, labrum: detail of dorsal submarginal row of flattened spines; 19C, left mandible v.v.; 19D, right mandible v.v.; 19E, hypopharynx v.v.; 19F, maxilla v.v.; 19G, labium, left d.v., right v.v. 19H, tarsal claw I; 19I, posterior margin of tergum IV. Figs. 19J-K, A. beatrixae: 19J, maxilla v.v.; 19K, labium. (D = denticles; L = lingua; P = prostheca; SL = superlingua; TS = transverse bipectinated seta; Roman numbers = segment numbers). Figs. 19J-K modified from GILLIES, 2001.

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the genus, sometimes redescribing some generic characteristics. At the present time, Baetodes includes thirty nine nominal species, distributed from Arizona to Argentina, making it one of the most species rich genera in the Neotropics. Twenty two species have been described from South America, distributed in Argentina, Bolivia, Brazil, Colombia, Ecuador, Peru, and Venezuela. Eighteen species are described from nymphs, two from adults, and two from both nymphs and adults. KOSS (1972) proposed new terms for structures that have been previously described as spines and his terminology is used in this paper. Adult Characteristics: 1) Fore wings with paired marginal intercalaries (Fig. 20B); 2) hind wings absent; 3) genitalia: forceps three-segmented (can appear to be four-segmented due to constriction of second segment in some species), basal segment with a strong distomedial projection, apical segment elongate (Figs. 20C, E).

A

B

IV

IV

P

D

P

C

E Plate 20. Baetodes, imagos. Figs. 20A-C, B. huaico: 20A, male l.v.; 20B, fore wing; 20C, genitalia v.v. Figs. 20D-20E, B. uruguai: 20D, male l.v.; 20E, genitalia v.v. (IV = intercalary veins; P = projection). Figs. 20A-E modified from NIETO, 2004a.

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Nymphal Characteristics: 1) Antennae short, 1.5-2 times of head capsule, and with a row of long fine setae on external margin of basal segments (Fig. 21A); 2) labrum: ventrally with heavily sclerotized lateral margins and a sublateral row of spine-like setae; dorsally the anterior margin with two kinds of bipectinate setae, fully branched setae at middle and apically branched setae laterally (Fig. 21B); 3) mandibles with incisors fused (Figs. 21D-E); 4) left mandible with prostheca robust with denticles apically (Fig. 21D); 5) right mandible with prostheca reduced to a spine-like seta (Fig. 21E); 6) hypopharynx: lingua subequal in length to superlingua (Fig. 21F); 7) maxillae with palpi two-segmented and subequal in length to galea-lacinia (Fig. 21G); 8) labium: glossae 0.5-0.75 times length and at least half the width of paraglossae, segment II of palpi without medial projection, segment III elongate (Fig. 21H); 9) coxae with high dorsal crest (Fig. 21I); 10) dorsal edge of femora with a compact row of fine setae intercalated with clavate setae (Fig. 21J); 11) tibiae anteriorly with a continuous longitudinal and transverse break on the dorsal and ventral surfaces (Fig. 21I); 12) tarsal claws with a row of denticles and with one stout subapical seta on lateral margin (Fig. 21K); 13) rudimentary hind wing pads; 14) gills (Fig. 21L) on abdominal segments I-V; 15) paraprocts without marginal spines; 16) terminal filament reduced, cerci with only a few inconspicuous setae. Distribution: North, Central, and South America. Biology: Species of Baetodes are found on the upper surfaces of rocks in fast-flowing water. They inhabit well-oxygenated streams and rivers, mainly located in the mountains. KEYS

TO

SOUTH AMERICAN BAETODES

Male Imagos 1 – 2(1) – 3(2) –

Abdominal tubercles present on segments II-V .................................. Baetodes spinifer Abdominal tubercles absent ............................................................................................. 2 Turbinate eyes large and erect, with upper surfaces contiguous (Fig. 20D) ........... ........................................................................................................................Baetodes uruguai Turbinate eyes moderately large and erect, upper surfaces not contiguous (Fig. 20A) ....................................................................................................................................... 3 Genitalia: segment II of forceps with strong constriction, giving the appearance of four-segmented forceps (as in Fig. 20E) ........................................ Baetodes arawak Genitalia: segment II of forceps without constriction (Fig. 20C) ..... Baetodes huaico

Nymphs 1 – 2(1) – 3(2) – 4(3) –

Ocelli absent .......................................................................................... Baetodes itatiayanus Ocelli present ...................................................................................................................... 2 Coxal gills present (Figs. 25F, L) ..................................................................................... 3 Coxal gills absent (Fig. 21I) ............................................................................................ 15 One finger-like gill on each coxa (Fig. 25F) ................................................................. 4 Two finger-like gills on each coxa (Fig. 25L) ............................................................... 6 Abdominal tubercles absent ......................................................... Baetodes sancticatarinae Abdominal tubercles present (Figs. 22L, 23C) ............................................................ 5

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Abdominal terga with clavate tubercles directed posteriorly (as in Fig. 23C); with two pectinate blade-like setae on tip of each glossa (as in Fig. 25K), and without non-pectinate blade-like setae on paraglossa .......................................Baetodes uruguai – Abdominal terga with tubercles prominent and pointed directed posteriorly (Fig. 23I); with one pectinate bladelike seta on tip of each glossa (Fig. 23H), and with one non-pectinate bladelike seta medially on each paraglossa ....................... ..................................................................................................................... Baetodes peniculus 6(3) Abdominal tubercles present (Fig. 23C) ....................................................................... 7 – Abdominal tubercles absent ........................................................................................... 13 7(6) First abdominal tubercle twice the length of the others (Figs. 22L, 24D) ............ 8 – First abdominal tubercle same size as the others (Fig. 23C) .................................... 9 8(7) Pronotum with tubercle (Fig. 22L); ventral edge of femora with scattered long spines (Fig. 22J) ............................................................................................ Baetodes gibbus – Pronotum without tubercle; ventral edge of femora with scattered short spines (Fig. 24B) ..............................................................................................Baetodes pseudogibbus 9(7) Pronotum with tubercle (Fig. 25B) .............................................................................. 10 – Pronotum without tubercle ............................................................................................ 11 10(9) Pronotum and mesoscutellum with double tubercles; abdominal tubercles large and erect (Fig. 25B); long spines on segment III of labial palpi ....... Baetodes spinae – Pronotum with a single median tubercle, mesoscutellum without tubercle; abdominal tubercles less prominent and directed posteriorly (Fig. 24A); short spines on segment III of labial palpi .................................................. Baetodes proiectus 11(9) Dorsal edge of femora with clavate spines slender and short, half the length of fine setae (Fig. 25M); segment III of labial palpi broadly rounded (Fig. 25K); apex of lingua rounded (Fig. 25J) .................................................................. Baetodes yuracare – Dorsal edge of femora with clavate spines long and slender, subequal to fine setae (Fig. 23B); segment III of labial palpi slender oval (as in Fig. 21H); apex of lingua broadly pointed (as in Fig. 21F) .................................................................................... 12 12(11) Dorsal edge of femora with 6 or 7 clavate spines; tarsal claws with 5 or 6 denticles ............................................................................................................................. Baetodes levis – Dorsal edge of femora with 10 or 11 clavate spines (Fig. 23A); tarsal claws with 10 or 11 denticles ...............................................................................................Baetodes huaico 13(6) Mandibles with 2 small lobes on the inner margin of incisors (Figs. 22A-B); labium with tips of glossae each with 2 short spines at base of one long blade-like spine (Fig. 22C) ................................................................................. Baetodes andamagensis – Mandibles without small lobes on the incisors (as in Figs. 21D-E); labium with glossae not as above ........................................................................................................ 14 14(13) Dorsal edge of femora with 6 or 7 clavate setae; labium with tips of glossae without pectinate blade-like setae (Fig. 22D) ....................................... Baetodes chilloni – Dorsal edge of femora with 10 or 11 clavate setae (Fig. 23E); labium with tips of glossae each with two pectinate blade-like setae (as in Fig. 25K) ..... Baetodes pehuenche 15(2) Abdominal tubercles present, although small (Fig. 22H) ........................................ 16 – Abdominal tubercles absent ........................................................................................... 18

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16(15) Labrum with a dorsal subapical row of subequal spine-like setae (Fig. 24F); lateral margin of mandibles with long setae (Figs. 24G-H) ........................... Baetodes rutilus – Labrum with a dorsal subapical row of spine-like setae varying in length, alternating one long, one short (as in Fig. 22F); lateral margin of mandibles without long setae (Figs. 21D-E) .................................................................................. 17 17(16) Tubercle present on the metanotum; segment I of labial palpi without a distomedial projection (as in Fig. 21H) ................................................ Baetodes serratus – Tubercle absent on the metanotum; segment I of labial palpi with a distomedial projection (Fig. 22G) ......................................................................... Baetodes cochunaensis 18(15) Terminal filament very short, less than half the length of abdominal segment X (Fig. 25E); labium with glossae very short, less than half the length of paraglossae (Fig. 25D) .................................................................................................... Baetodes traverae – Terminal filament at least as long as abdominal segment X; labium with glossae at least 3/4 length of paraglossae (Fig. 21H) ................................................................. 19 19(18) Labium with tips of glossae without pectinate blade-like seta; one wide nonpectinate blade-like seta distally near inner margin of each paraglossa (Fig. 25A); gills oblong (as in Fig. 24E) ......................................................................... Baetodes solus – Labium with tips of glossae with one pectinate blade-like setae (Fig. 21H); paraglossae with wide pectinate blade-like setae; gills lanceolate, wider at base, apically acute (Fig. 21L) ........................................................................... Baetodes copiosus CLAVES

PARA LOS

BAETODES

SUDAMERICANOS

Imagos Machos 1 – 2(1) – 3(2) –

Tubérculos abdominales presentes en los segmentos II-V ............... Baetodes spinifer Tubérculos abdominales ausentes .................................................................................. 2 Ojos compuestos altos y cónicos, superficies superiores contiguas (Fig. 20D) ..... ........................................................................................................................Baetodes uruguai Ojos compuestos moderadamente altos y cónicos, superficies superiores no contiguas (Fig. 20A) ........................................................................................................... 3 Genitalia (como en la Fig. 20E): segundo segmento de los fórceps con una fuerte constricción aparentando fórceps tetrasegmentados ......................... Baetodes arawak Genitalia (Fig. 20C): segundo segmento de los fórceps sin constricción ................ ..........................................................................................................................Baetodes huaico

Ninfas 1 – 2(1) – 3(2) – 4(3) –

Ocelos ausentes .................................................................................... Baetodes itatiayanus Ocelos presentes ................................................................................................................ 2 Branquias coxales presentes (Figs. 25F, L) ................................................................... 3 Branquias coxales ausentes (Fig. 21I) ......................................................................... 15 Cada coxa con una branquia digitiforme (Fig. 25F) ................................................... 4 Cada coxa con dos branquias digitiformes (Fig. 25L) ................................................ 6 Tubérculos abdominales ausentes .............................................. Baetodes sancticatarinae Tubérculos abdominales presentes (Figs. 22L, 23C) .................................................. 5

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Tergos abdominales con tubérculos pequeños y redondeados dirigidos posteriormente (como en la Fig. 23C); ápice de cada glosa con dos setas en forma de espinas pectinadas, paraglosas sin setas en forma de espinas pectinadas (como en la Fig. 25K) ............................................................................................Baetodes uruguai – Tergos abdominales con tubérculos prominentes y agudos dirigidos posteriormente (Fig. 23I); ápice de cada glosa con una seta en forma de espina pectinada (Fig. 23H), paraglosas con 1 seta en forma de espina no pectinada medialmente ............................................................................................. Baetodes peniculus 6(3) Tubérculos abdominales presentes (Fig. 23C) ............................................................. 7 – Tubérculos abdominales ausentes ................................................................................ 13 7(6) Primer tubérculo abdominal dos veces el tamaño de los otros (Figs. 22L, 24D) ....... 8 – Primer tubérculo abdominal de igual tamaño que los restantes (Fig. 23C) ........... 9 8(7) Pronoto con un tubérculo (Fig. 22L); margen ventral de los fémures con espinas clavadas largas (Fig. 22J) ............................................................................. Baetodes gibbus – Pronoto sin tubérculo; margen ventral de los fémures con espinas clavadas cortas (Fig. 24B) ..............................................................................................Baetodes pseudogibbus 9(7) Pronoto con tubérculo (Fig. 25B) ................................................................................ 10 – Pronoto sin tubérculo ..................................................................................................... 11 10(9) Pronoto y mesoescutelo con tubérculos dobles; tubérculos abdominales grandes y erectos (Fig. 25B); espinas largas en el segmento III del palpo labial ...................... .......................................................................................................................... Baetodes spinae – Pronoto con un tubérculo, mesoescutelo sin tubérculo; tubérculos abdominales menos prominentes y agudos dirigidos posteriormente (Fig. 24A); espinas cortas en el segmento III del palpo labial ....................................................... Baetodes proiectus 11(9) Margen dorsal de los fémures con espinas clavadas delgadas y cortas, la mitad de la longitud de las setas finas (Fig. 25M); segmento III del palpo labial agudo (Fig. 25K); ápice de la lingua redondeado (Fig. 25J) ................................. Baetodes yuracare – Margen dorsal de los fémures con espinas clavadas largas y delgadas, subiguales a las setas finas (Fig. 23B); segmento III del palpo labial ovalado (similar a la Fig. 21H); ápice de la lingua agudo (similar a la Fig. 21F) .............................................. 12 12(11) Margen dorsal de los fémures con 6 o 7 espinas clavadas; uñas tarsales con cincoseis dentículos .................................................................................................. Baetodes levis – Margen dorsal de los fémures con 10 o 11 espinas clavadas (Fig. 23A); uñas tarsales con 10-11 dentículos .....................................................................Baetodes huaico 13(6) Mandíbulas con 2 lóbulos pequeños en el margen interno de los incisivos (Figs. 22A-B); ápice de las glosas del labio con 2 espinas cortas en la base de una seta con forma de espina larga (Fig. 22C) ........................................... Baetodes andamagensis – Mandíbulas sin lóbulos en el margen interno de los incisivos (similar a las Figs. 21D-E); glosa del labio no como arriba ...................................................................... 14 14(13) Margen dorsal de los fémures con 6 o 7 espinas clavadas; ápice de las glosas del labio con setas no pectinadas (Fig. 22D) .............................................. Baetodes chilloni – Margen dorsal de los fémures con 10 o 11 espinas clavadas (Fig. 23E); ápice de las glosas del labio con dos setas con forma de espinas pectinadas (similar a la Fig. 25K) .................................................................................................................... Baetodes pehuenche

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15(2) Tubérculos abdominales presentes aunque pequeños (Fig. 22H) .......................... 16 – Tubérculos abdominales ausentes ................................................................................ 18 16(15) Labro con una hilera dorsal subapical de setas con forma de espinas subiguales (Fig. 24F); margen externo de las mandíbulas con setas largas (Figs. 24G-H) ...... ......................................................................................................................... Baetodes rutilus – Labro con una hilera dorsal subapical de setas con forma de espinas de longitud variable, alternando una larga y una corta (como en la Fig. 22F); margen externo de las mandíbulas sin setas largas (Figs. 21D-E) ....................................................... 17 17(16) Tubérculo presente en el metanoto; segmento I del palpo labial sin una proyección distomedial (similar a la Fig. 21H) ................................... Baetodes serratus – Tubérculo ausente en el Metanoto; segmento I del palpo labial con una proyección distomedial (Fig. 22G) ................................................. Baetodes cochunaensis 18(15) Filamento terminal muy corto, menos de la mitad de la longitud del segmento abdominal X (Fig. 25E); glosas del labio cortas, menos de la mitad de la longitud de las paraglosas (Fig. 25D) .................................................................... Baetodes traverae – Filamento terminal por lo menos tan largo como la longitud del segmento abdominal X; glosas del labio por lo menos 3/4 de la longitud de las paraglosas (Fig. 21H) ........................................................................................................................... 19 19(18) Ápice de las glosas del labio con setas con forma de espinas no pectinadas; margen interno de las paraglosas con una seta con forma de espina no pectinada (Fig. 25A); branquias oblongas (similar a la Fig. 24E) ........................... Baetodes solus – Ápice de las glosas del labio con una seta con forma de espina pectinada; paraglosas con setas con forma de espinas pectinadas (Fig. 21H); branquias lanceoladas, más anchas en la base, apicalmente agudas (Fig. 21L) ................................. Baetodes copiosus Baetodes andamagensis MAYO Baetodes andamagensis MAYO, 1972: 231 (nymph); NIETO, 2004a: 65.

Taxonomy: This species is known only from nymphs, which can be distinguished from the other species of the genus by the following combination of characters: 1) Two small lobes distally on incisors at inner margin of mandibles (Figs. 22A-B); 2) tip of each glossa with two short, blunt spines at base of one long blade-like spine (Fig. 22C); 3) two finger-like gills on each coxa; 4) abdominal tubercles absent; 5) abdominal gills smoky in basal half. Distribution: Peru. Baetodes arawak (TRAVER) Pseudocloeon arawak TRAVER, 1943: 90 (male). Baetodes arawak; FLOWERS, 1987: 8 (male); NIETO, 2004a: 65.

Taxonomy: This species is known only from male imagos, and can be distinguished from the other species of the genus by the following combination of characters: 1) Turbinate eyes moderately large, erect, upper surface almost round, not contiguous

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C A B P

SL

D

P

L

F

E PS III II

G

H

I

J

K D

I

SS

L

Plate 21. Baetodes copiosus, nymph. Figs. 21A-L. 21A, antenna; 21B, labrum, left d.v., right v.v.; 21C, labrum: subapical row of setae d. v.; 21D, left mandible v.v.; 21E, right mandible v.v.; 21F, hypopharynx v.v.; 21G, maxilla v.v.; 21H, labium, left d.v., right v.v.; 21I, leg I; 21J, dorsal edge of femur I; 21K, tarsal claw I; 21L, gill. (D = denticles; L = lingua; P = prostheca; PS = pectinate seta; SL = superlingua; SS = subapical seta; Roman numbers = segment numbers). Figs. 21A-L modified from NIETO, 2004a.

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apically (as in Fig. 20A); 2) forceps: second segment with a strong constriction, giving the appearance of two segments, basal segment with apical projection (as in Fig. 20E). Distribution: Venezuela. Baetodes chilloni MAYO Baetodes chilloni MAYO, 1972: 228 (nymph); NIETO, 2004a: 65.

Taxonomy: This species, known only from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Labrum with a dorsal subapical row of spine-like setae varying in length, alternating one long, one short (as in Fig. 22F); 2) two finger-like gills on each coxa; 3) dorsal edge of femora with 6-7 clavate setae; 4) tarsal claws with 5-6 denticles; 5) abdominal tubercles absent; 6) glossae with no pectinate setae (Fig. 22D). Distribution: Peru. Baetodes cochunaensis NIETO Baetodes cochunaensis NIETO, 2004a: 65.

Taxonomy: Baetodes cochunaensis, is known only from nymphs and can be distinguished from the other species of the genus Baetodes by the followings combination of characters: 1) Labrum expanded laterally, with a dorsal subapical row of spine-like setae varying in length, alternating one long, one short (Figs. 22E-F); 2) coxal gills absent; 3) dorsal edge of femora with 7-8 clavate setae; 4) tarsal claws with 11-12 denticles; 5) abdominal tubercles very small, only visible in dorsal view (Fig. 22H); 6) abdominal gills whitish, oblong and asymmetrical, wider at base, apically acute (Fig. 22I); 7) segment one of the labial palpi with a distomedial projection (Fig. 22G). Distribution: Argentina (Tucumán and Jujuy). Baetodes copiosus NIETO Baetodes copiosus NIETO, 2004a: 66.

Taxonomy: Baetodes copiosus is known only from nymphs and can be distinguished from the other species of Baetodes by the following combination of characters: 1) Labrum (Figs. 21B-C) expanded laterally, with a dorsal subapical row of subequal spine-like setae, and ventral surface with a row of pectinate spine-like setae; 2) coxal gills absent; 3) dorsal edge of femora with 7-8 short and slender clavate setae (Figs. 21I-J); 4) tarsal claws with 11-12 denticles (Fig. 21K); 5) abdominal tubercles absent; 6) abdominal gills whitish, lanceolate, wider at base, apically acute (Fig. 21L). Distribution and Biology: Argentina (Tucumán and Salta). This species was collected in rivers of rocky substrate, from mountain rain forests.

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A

B C

D

E

PS III II

F IP T

G

T

H I T

K T

J

CG

L

M

Plate 22. Baetodes, nymphs. Figs. 22A-C, B. andamagensis: 22A, left mandible v.v.; 22B, right mandible v.v.; 22C, tip of glossa. Fig. 22D, B. chilloni: labium v.v. Figs. 22E-I, B. cochunaensis: 22E, labrum, left d.v., right v.v.; 22F, labrum, subapical row of setae d. v.; 22G, labium, left d.v., right v.v.; 22H, abdominal segments d.v.; 22I, gill. Figs. 22J-M, B. gibbus: 22J, leg I; 22K, dorsal edge of femur; 22L, thorax and abdominal segments l.v.; 22M, gill. (CG = coxal gill; IP = internal projection; PS = pectinate seta; T = tubercle; Roman numbers = segment numbers). Fig. 22D modified from MAYO, 1972; 22E-M from NIETO, 2004a.

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Baetodes gibbus NIETO Baetodes gibbus NIETO, 2004a: 66.

Taxonomy: Baetodes gibbus is known only from nymphs and can be distinguished from the other species of Baetodes by the following combination of characters: 1) Labrum: dorsal surface with a subapical row of 6-7 spine-like setae near midline alternating one long, one short (as in Fig. 22F); 2) two finger-like gills on each coxa, gills at least 1.5 times length of coxa (Fig. 22J); 3) dorsal edge of femora with 7-8 clavate setae subequal to fine setae (Fig. 22K); 4) tarsal claws with 6 denticles; 5) abdominal tubercles present, first tubercle twice the size of the others (Fig. 22L); 6) abdominal gills whitish and oblong (Fig. 22M); 7) pronotum with a single median tubercle (Fig. 22L); 8) ventral edge of femora with scattered long spines (Fig. 22J). Distribution and Biology: Bolivia. This species was collected in rivers of rocky substrate, from mountain rain forests. Baetodes huaico NIETO Baetodes huaico NIETO, 2004a: 69.

Taxonomy: This species is known from nymphs and male imagos. Baetodes huaico can be distinguished from the other species of Baetodes by the following combination of characters. In the imago: 1) Turbinate eyes moderately large and erect, upper surface not contiguous apically (Fig. 20A); 2) genitalia (Fig. 20C): second segment without constriction. In the nymph: 1) Labrum: dorsal surface with a submarginal row of 8-9 spine-like setae near midline alternating one long, one short and 2-3 long seta near lateral margin (as in Fig. 22F); 2) two finger-like gills on each coxa, slightly longer than coxae (Fig. 23A); 3) dorsal edge of femora with 10-11 clavate setae (Fig. 23B); 4) tarsal claws with 11 denticles; 5) abdomen with small but distinct tubercles (Fig. 23C); 6) abdominal gills whitish and oblong (Fig. 23D); 7) abdominal color pattern with segments I-III, VII-IX dark and segments IV-VI, X light. Distribution and Biology: Argentina (Catamarca, Jujuy, Salta and Tucumán). This species was collected in many rivers, especially the rivers from mountain rain forests. Baetodes itatiayanus DEMOULIN Baetodes itatiayanus DEMOULIN, 1955a: 22 (nymph); MAYO, 1968a: 256; NIETO, 2004a: 70.

Taxonomy: This species is known only from nymphs. Baetodes itatiayanus can be distinguished by the following combination of characters: 1) Ocelli absent; 2) coxal gills absent; 3) abdominal tubercles present. Distribution: Brazil. Baetodes levis MAYO Baetodes levis MAYO, 1968a: 253 (nymph); NIETO, 2004a: 70.

Taxonomy: This species is known only from nymphs. Baetodes levis can be distinguished from other species of the genus by the following combination of characters: 1)

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Labrum with a subapical row of 10-11 spine-like setae near midline; 2) glossae robust, with two pectinate setae on each tip; 3) two finger-like gills on each coxa; 4) dorsal edge of femora with 6-7 clavate spines; 5) tarsal claws with 5-6 denticles; 6) abdominal tubercles directed posteriorly. Distribution: Colombia and Ecuador. Baetodes pehuenche NIETO Baetodes pehuenche NIETO, 2004a: 71.

Taxonomy: Baetodes pehuenche is known only from nymphs and can be distinguished from the other species of Baetodes by the following combination of characters: 1) Labrum with dorsal surface with a subapical row of 8-9 spine-like setae near midline, alternating one long and one short (as in Fig. 22F); 2) two finger-like gills on each coxa, at least 1.5 length of coxae (Fig. 23E); 3) dorsal edge of femora with 10-11 clavate setae as long as fine setae (Fig. 23F); 4) tarsal claws with 6-7 denticles; 5) abdominal tubercles absent; 6) abdominal gills oblong and asymmetrical (Fig. 23G); 7) abdominal color pattern with segments I-IX dark and segment X light. Distribution: Argentina (Neuquén). Baetodes peniculus MAYO Baetodes peniculus MAYO, 1973b: 310 (nymph); NIETO, 2004a: 72.

Taxonomy: Baetodes peniculus, known only from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Glossae each with one pectinate seta on tip (Fig. 23H) and paraglossae with one nonpectinate long seta; 2) one finger-like gill on each coxa; 3) each abdominal tergum with prominent, pointed tubercle directed posteriorly (Fig. 23I). Distribution: Venezuela. Baetodes proiectus MAYO Baetodes proiectus MAYO, 1973b: 308 (nymph); NIETO, 2004a: 72.

Taxonomy: Baetodes proiectus, known only from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Two fingerlike gills on each coxa; 2) pronotum, metanotum and abdominal terga with prominent tubercles (Fig. 24A); 3) sternum II of immature specimens with a projection from rim of sclerotized area around apodeme on each side forming a small, erect, round-tipped lobe. Distribution: Bolivia. Baetodes pseudogibbus NIETO Baetodes pseudogibbus NIETO, 2004a: 72.

Taxonomy: Baetodes pseudogibbus, known only from nymphs, can be distinguished from the other species of Baetodes by the following combination of characters: 1) Labrum:

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B

D

A CG T

T

C

G

E

F

H

T

CG

T

I

Plate 23. Baetodes, nymphs. Figs. 23A-D, B. huaico: 23A, leg I; 23B, dorsal edge of femur; 23C, metathoracic and abdominal segments l.v.; 23D, gill. Figs. 23E-G, B. pehuenche: 23E, leg I; 23F, dorsal edge of femur; 23G, gill. Figs. 23H-I, B. peniculus: 23H, tip of glossa; 23I, metanotum and abdominal terga l.v. (CG = coxal gills; T = tubercle). Figs. 23A-G modified from NIETO, 2004a; 23H-I from MAYO, 1973b.

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dorsal surface with a subapical row of 6-7 spine-like setae near midline alternating one long, one short (as in Fig. 22F); 2) two finger-like gills on each coxa, more than 1.5 length of coxae (Fig. 24B); 3) dorsal edge of femora with 6-7 clavate setae (Fig. 24C); 4) tarsal claws with 6-7 denticles; 5) abdominal tubercles present, first tubercle twice the size of others (Fig. 24D); 6) abdominal gills whitish, oblong and asymmetrical (Fig. 24E). Distribution: Peru. Baetodes rutilus NIETO Baetodes rutilus NIETO, 2004a: 73.

Taxonomy: Baetodes rutilus, known only from nymphs, can be distinguished from the other species of Baetodes by the following combination of characters: 1) Labrum (Fig. 24F) greatly expanded laterally, dorsal surface with a subapical row of spine-like setae; 2) coxal gills absent; 3) dorsal edge of femora with 10-11 short and robust clavate setae (Figs. 24J-K); 4) tarsal claws with 7-8 denticles; 5) abdominal spine-like tubercles directed posteriorly (Fig. 24L); 6) abdominal gills evenly rounded (Fig. 24M); 7) mandibles with lateral margin curved and with a row of long fine setae (Figs. 24G-H); 8) labium with glossae short, slightly longer than half the length of paraglossae and with one pectinate bladelike seta at tip (Fig. 24I). Distribution and Biology: Bolivia. This species was found in a waterfall, on the surfaces of vertical rocks where water flows, in the same habitat where a species of Mayobaetis was collected. Baetodes sancticatarinae MAYO Baetodes sancticatarinae MAYO, 1972: 240 (nymph); NIETO, 2004a: 74.

Taxonomy: Baetodes sancticatarinae, known only from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Glossae with pectinate bladelike setae and one non-pectinate blade-like seta on each paraglossa; 2) one finger-like gill on each coxa; 3) tarsal claws with 8 denticles; 4) abdominal tubercles absent; 5) abdominal gills oblong and asymmetrical. Distribution: Brazil. Baetodes serratus NEEDHAM & MURPHY Baetodes serratus NEEDHAM & MURPHY, 1924: 55 (nymph); TRAVER, 1944: 20, (nymph); MAYO, 1968a: 255 (nymph); NIETO, 2004a: 74.

Taxonomy: Baetodes serratus, known only from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Labrum with a subapical row of 8-9 spine-like setae near midline, alternating one long and one short (as in Fig. 22F); 2) coxal gills absent; 3) metanotum and abdominal terga with tubercles. Distribution: Brazil.

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A

T

101

C

T T

D

B CG P

E

G

P

H F PS III II I

K

J

I T

L

M

Plate 24. Baetodes, nymphs. Fig. 24A, B. proiectus: metanotum and abdominal terga l.v. Figs. 24B-E, B. pseudogibbus: 24B, leg I; 24C, dorsal edge of femur; 24D, metanotum and abdominal segments l.v.; 24E, gill. Figs. 24F-M, B. rutilus: 24F, labrum d.v.; 24G, left mandible v.v.; 24H, right mandible v.v.; 24I, labium, left d.v., right v.v.; 24J, leg I; 24K, dorsal edge of femur; 24L, metanotum and abdominal segments l.v.; 24M, gill. (CG = coxal gill; P = prostheca; PS = pectinate seta; T = tubercle; Roman numbers = segment numbers). Fig. 24A modified from MAYO, 1973b; 24B-M from NIETO, 2004a.

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Baetodes solus MAYO Baetodes solus MAYO, 1972: 231 (nymph); DOMÍNGUEZ et al., 2002: 462; NIETO, 2004a: 75.

Taxonomy: Baetodes solus, known only from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Glossae without pectinate blade-like setae, with one wide non-pectinate blade-like seta distally near inner margin of paraglossae (Fig. 25A); 2) coxal gills lacking; 3) abdominal tubercles absent. Distribution: Colombia and Peru. Baetodes spinae MAYO Baetodes spinae MAYO, 1968a: 251 (nymph); NIETO, 2004a: 75.

Taxonomy: This species, known only from nymphs can be distinguished by the following combination of characters: 1) Long spines on apical segment of labial palpi; 2) two finger-like gills on each coxa; 3) pronotum and mesonotum with a pair of tubercles, abdomen with prominent tubercles (Fig. 25B). Distribution: Colombia and Ecuador. Baetodes spinifer TRAVER Baetodes spiniferum TRAVER, 1943: 94 (male, female); NIETO, 2004a: 75. Baetodes spinifer; MAYO, 1972: 226; FLOWERS, 1987: 8.

Taxonomy: Baetodes spinifer, known only from the male imago and female subimago, and can be distinguished from other species of the genus by the following combination of characters: 1) Turbinate eyes very large, cylindrical, erect, apical surface round, eyes contiguous apically (as in Fig. 20D); 2) basal segment of forceps with a distomedial projection; 3) fore tibiae more than twice as long as femora; 4) abdominal tubercles on segments II-V. Distribution: Venezuela. Baetodes traverae MAYO Baetodes traverae MAYO, 1972: 230 (nymph); NIETO, 2004a: 75.

Taxonomy: Baetodes traverae, known only from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Labrum expanded laterally and with a group of spine-like setae behind apical margin (Fig. 25C); 2) coxal gills absent; 3) dorsal edge of femora with 7-8 clavate setae; 4) tarsal claws with 10 denticles; 5) abdominal tubercles absent; 6) abdominal gills oval; 6) glossae very short, less than half the length of the quadrate paraglossae (Fig. 25D); 7) terminal filament is extremely short (Fig. 25E). Distribution: Peru.

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T

B A C

G D

E

F

CG

H

SL L

M I PS

J

L III

CG

II I

K

N

Plate 25. Baetodes, nymphs. Fig. 25A, B. solus: labium v.v. Fig. 25B, B. spinae: nymph l.v. Figs. 25C-E, B. traverae: 25C, labrum d.v.; 25D, labium v.v.; 25E, abdominal segment X with cerci and terminal filament. Figs. 25F-H, B. uruguai: 25F, leg I; 25G, dorsal edge of femora; 25H, gill. Figs. 25I-N, B. yuracare: 25I, labrum, left d.v., right v.v.; 25J, hypopharynx v.v.; 25K, labium, left d.v., right v.v.; 25L, leg I; 25M, dorsal edge of femur; 25N, gill. (CG = coxal gill; L = lingua; PS = pectinate seta; SL = superlingua; T = tubercle; Roman numbers = segment numbers). Figs. 25A, C-E modified from MAYO, 1972; 25B from MAYO, 1968; 25F-N from NIETO, 2004a.

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Baetodes uruguai NIETO Baetodes uruguai NIETO, 2004a: 75.

Taxonomy: Baetodes uruguai, known from male imagos and nymphs, can be distinguished from the other species of Baetodes by the following combination of characters. In the imago: 1) Turbinate eyes large, cylindrical and erect; height of stalk almost twice length of eye diameter, eyes contiguous apically (Fig. 20D); 2) second segment of forceps with a constriction, appearing two-segmented (Fig. 20E). In the nymph: 1) Labrum, dorsal surface with a subapical row of 8-9 spine-like setae near midline alternating one long, one short (as in Fig. 22F); 2) one finger-like gill on each coxae (Fig. 25F); 3) dorsal edge of femora with 89 clavate setae slightly longer than fine setae (Fig. 25G); 4) tarsal claws with 9 denticles; 5) abdominal tubercles clavate and directed posteriorly; 6) gills whitish, transparent, basally wider, apically acute, anterior margin straight (Fig. 25H). Distribution and Biology: Argentina (Misiones). This species was found in rivers with rocky substrate. Baetodes yuracare NIETO Baetodes yuracare NIETO, 2004a: 76.

Taxonomy: Baetodes yuracare, known from male subimagos and nymphs, can be distinguished from the other species of Baetodes by the following combination of characters. In the nymphs: 1) Labrum twice as wide as long, dorsal surface with a subapical row of 6-7 spine-like setae near midline alternating one long and one short (Fig. 25I); 2) two finger-like gills on each coxa, at least twice length of coxae (Fig. 25L); 3) dorsal edge of femora with 67 short clavate setae, less than half the length of fine setae (Fig. 25M); 4) tarsal claws with 56 denticles; 5) small abdominal tubercles on each segment; 6) gills mostly whitish, with first third brownish, oblong with regular margin (Fig. 25N); 7) segment III of labial palpi broadly rounded (Fig. 25K); 8) apex of lingua rounded (Fig. 25J). Distribution: Bolivia. This species was collected in rivers of rocky substrate, in mountain rain forests. Genus Bernerius WALTZ & MCCAFFERTY Bernerius WALTZ & MCCAFFERTY, 1987a: 179. (Type-species: Bernerius incus WALTZ & MCCAFFERTY, original designation).

Taxonomy: The genus Bernerius was established by WALTZ & MCCAFFERTY (1987a). They included only one species, B. incus, originally described as Baetis (?) sp. B by BERNER (1980). This species, known only from nymphs, is the only species presently included in the genus. The type material was examined and the descriptions of some characters are modified from the original. Bernerius is in revision at this moment, and there are some doubts about its validity. Adults Characteristics: Unknown. Nymphal Characteristics: 1) Antennae long, at least 2 times of head capsule; 2) labrum (Fig. 26A), dorsally with a pair of long subapical setae, anterior margin with two

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P P

A

L

SL

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C

C

D III

J E

II

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I

Plate 26. Bernerius incus, nymph. Figs. 26A-J. Figs. 26A-F, mouthparts: 26A, labrum, left d.v., right v.v.; 26B, left mandible v.v.; 26C, right mandible v.v.; 26D, hypopharynx v.v.; 26E, maxilla v.v.; 26F, labium, left d.v., right v.v. 26G, leg I; 26H, posterior margin of abdominal tergum IV d.v.; 26I, gill IV; 26J, paraproct. (A = arc of long setae; C = constriction; L = lingua; P = prostheca; SL = superlingua; Roman numbers = segment numbers).

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kinds of bifid setae: basally bifid near midline, apically bifid near lateral margin; 3) left mandible (Fig. 26B) with incisors fused apically, prostheca robust and with denticles apically; 4) right mandible (Fig. 26C) with incisors cleft apically, prostheca slender and bifid; 5) hypopharynx (Fig. 26D): lingua with small rounded projection, lateral margin of superlingua with a row of short spines; 6) maxillae (Fig. 26E) with palpi two-segmented, segment II with a slight constriction apically; 7) labium (Fig. 26F): glossae subequal to paraglossae, segment II of palpi without lateral projection; 8) femora (Fig. 26G) with dorsal edge parallel to ventral edge and with a row of spines, tibia with a subproximal arc of long setae; 9) tarsal claws 0.3 times length of tarsi, without denticles (Fig. 26G); 10) hind wing pads present; 11) posterior margin of abdominal terga with spines (Fig. 26H), sterna with scales and fine setae, no setal tufts; 12) abdominal gills (Fig. 26I) present on segments I-VII, at least two times length of each segment; 13) posterior margin of paraprocts with spines (Fig. 26J); 14) terminal filament subequal in length to cerci. Distribution: Peru. Bernerius incus WALTZ & MCCAFFERTY Baetis (?) sp. BERNER, 1980: 190. Bernerius incus WALTZ & MCCAFFERTY, 1987a: 181.

Taxonomy: This species, known from nymphs, can be distinguished by the following combination of characters: 1) Maxillae with palpi 1.5 times length of galea-lacinia (Fig. 26E); 2) tibiae and tarsi with bipectinate spines (Fig. 26G); 3) gills elongate, pointed apically (Fig. 26I). Distribution: Discussed in the generic description. Genus Callibaetis EATON Callibaetis EATON, 1881: 196; GILLIES, 1990: 15. (Type-species: Callibaetis pictus EATON, original designation).

Taxonomy: This was one of the first genera described for the family Baetidae. It was established by EATON (1881) and included one species, Callibaetis pictus. Since then, many different species have been described, although some have been synonymized. This genus has a wide distribution from North America to Argentina, in South America. In most cases, the species are known only from adults, especially from female imagos. That is because the females have very characteristic color patterns in the wings and only sometimes are these patterns present in male imagos. Although this genus is very distinct with respect to other genera, the species are often difficult to identify. GILLIES (1990) reviewed all the species in Argentina and synonymized 19 species, with only 9 species remaining valid. In South America 14 species are known from adults and only in three cases (C. guttatus, C. radiatus and C. willineri) the associated nymphs are also known (DA-SILVA, 1991; SALLES et al., 2003b; TRAVER, 1944). Adult Characteristics: 1) Turbinate eyes of male oval, height of stalk half of eye diameter; 2) body with small reddish spots; 3) male wings hyaline or with spots, female wings always pigmented with a characteristic color pattern; 4) fore wings with marginal intercalary veins paired (Fig. 27E) or single (Fig. 27A); hind wings with three longitudinal veins, several

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CP V

V

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A IV

IV

D

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IV Plate 27. Callibaetis, imagos. Figs. 27A-C, C. radiatus . Female: 27A, fore wing; 27B, hind wing. Male: 27C, genitalia. Fig. 27D, C. camposi, female hind wing. Figs. 27E-H, C. dominguezi . Male: 27E, fore wing; 27F, hind wing. Female: 27G, fore wing; 27H, hind wing. Figs. 27I-J, C. fasciatus , female: 27I, fore wing; 27J, hind wing. Figs. 27K-L, C. gonzalezi, female: 27K, fore wing; 27L, hind wing. (CP = costal process; IV = intercalary veins; V = vitta). Figs. 27A-B, 27E-L modified from GILLIES, 1990; 27D modified from NAVÁS, 1930b.

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cross veins and costal process placed in basal third of wing (Figs. 27D, F); 5) genitalia with forceps three segmented, basal segment with a small distomedial bulge (Fig. 27C). Nymphal Characteristics: 1) Head longer than wide; 2) antennae long, 5-6 times head length; 3) labrum (Fig. 29A) wider than long, with a row of flattened spine-like setae along anterior margin (Fig. 29B); 4) mandibles (Figs. 29C-D) with incisors deeply cleft, inner set of incisors at oblique angle with outer set, with or without tuft of setae between prostheca and mola; 5) left mandible (Fig. 29C): prostheca with two denticles apically; 6) right mandible (Fig. 29D): prostheca reduced to a simple seta; 7) hypopharynx (Fig. 29E): lingua longer than superlingua and with a rounded projection apically, superlingua rounded apically; 8) maxillae (Fig. 29F) with long canines, a row of long setae on apex of galea-lacinia, palpi two-segmented, 1-1.5 times longer than galea-lacinia and with a combination of spinelike setae and simple setae; 9) labium (Fig. 29G) with glossae equal or shorter but always wider than paraglossae, glossae and paraglossae with a row of spine-like setae, segment II of palpi without projection and with a row of spine-like setae, segment III elongate and with a medial concavity; 10) legs (Fig. 29H) long and narrow, dorsal edge of femora parallel to ventral edge, dorsal edge with a row of short spines, tibiae longer subequal to tarsi and with bipectinate spines (Fig. 29J); 11) ventral margin of femur I with a long row of two/three pointed-spines (Fig. 29I); 12) tarsal claws with two rows of very long cylindrical denticles (Fig. 29K); 13) hind wing pads present; 14) posterior margin of abdominal terga with spines (Fig. 29L); 15) gills present on abdominal segments I-VII, elongate, trachea pigmented and with two lamellae narrowly connected to each other (Figs. 29M-O); 16) posterior margin of paraprocts with spines (Fig. 29P); 17) terminal filament subequal in length to cerci. Distribution: North, Central, and South America. Biology: Callibaetis can be found in many different habitats. The nymphs have been collected in rivers, streams or temporal waters with rocky or sandy substrate. Many species of this genus are ovoviviparous, with the female imagos maintaining the eggs in their abdomen until they deposit first instar nymphs in the water. KEYS

TO

Imagos 1 – 2(1) – 3(2) –

5

SOUTH AMERICAN CALLIBAETIS

5

Fore wings with single marginal intercalary veins (Fig. 27A), except sometimes double in 1 or 2 spaces at tip or base of wing ............................................................ 2 Fore wings with paired marginal intercalary veins (Fig. 27E) .................................. 5 Wings in both sexes completely hyaline (Fig. 28G) .......................... Callibaetis pollens Wings at least of females with dark bands ................................................................... 3 In females, vittae without clear windows around cross veins, pigment extending broadly across base of wings to anal margin (Fig. 27K); basal one third to half of hind wings tinted brown (Fig. 27L) ................................................. Callibaetis gonzalezi In females, vittae with numerous clear windows around cross veins, pigment never extending to anal margin (Fig. 27A); hind wings hyaline (Fig. 27B) ....................... 4

C. camposi, C. gregarius and C. nigrivenosus were not included in the key because the original descriptions do not have enough characters to distinguish these species from the others.

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–

10(9) –

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Hind wings with 7 to 9 cross veins and a subquadrangular costal process (Fig. 27B) .................................................................................................................. Callibaetis radiatus Hind wings with 12 to 15 cross veins and a broadly rounded costal process (Fig. 28L) .................................................................................................................. Callibaetis viviparus Fore wings, at least in females, with spots along posterior margin, sometimes extended forming bands (Figs. 28A-B) ............................................. Callibaetis guttatus Fore wings with bands or spots but not on the posterior margin (Figs. 27I, 28I) ...... 6 Fore wings of both sexes with conspicuous, oblique or transverse, pigmented bands, leaving broad, clear areas between them (Figs. 27I, 28I) ............................................. 7 Fore wings in both sexes without clearly defined bands, or if banding present, taking the form of lines of narrow windows in an otherwise lightly pigmented wing field (Figs. 27G, 28M) ............................................................................................ 8 Fore wings with 3 more or less discrete bands (Fig. 27I) ............ Callibaetis fasciatus Fore wings with 2 main bands, the outer band approximately parallel to posterior margin of the wing (Fig. 28I) ...............................................................Callibaetis sellacki Hind wings without cross veins (Fig. 27H); fore wings of females with vitta restricted to costal and subcostal areas, without windows (Fig. 27G); in fore wings of males, distal third of wings uniformly tinted brown (Fig. 27E) ................ Callibaetis dominguezi Hind wings with 6 to 20 cross-veins; fore wings of females with vitta extending at least partly to area posterior to vein R1, with numerous windows (Fig. 28E); male fore wings clear or with a small spot in stigma ........................................................... 9 Tibiae II and III with 2 dark spots in basal half in addition to a subapical spot; abdominal sterna with dark anterolateral streaks, but without paired submedian markings; female fore wing pigment limited to the vitta (Fig. 28P), male fore wings unpigmented except sometimes in stigma .................................................................. 10 Tibiae II and III may be dark, but not showing two distinct spots in basal half; abdominal sterna with paired, submedian dash marks in addition to anterolateral streaks; fore wings in both sexes with heavly pigmented vitta and diffuse light pigmentation throughout rest of wing, sometimes with lines of clear windows around cross-veins (Fig. 28E) ................................................................ Callibaetis jocosus In female, basal half of vittae of fore wings overlapping vein R1 to as far as R2 in less than 1/2 of its length (Fig. 28P); in male, stigma of fore wings often with a patch of smoky pigment; no median markings on abdominal sterna ............... Callibaetis zonalis In female, basal half of vittae of fore wings overlapping vein R1 to as far as R2 in 1/2 or more of its length (Fig. 28M); in male, fore wing entirely hyaline; a median, maroon dash usually present on sterna II-VIII .............................. Callibaetis willineri

Nymphs 1 – 2(1) –

Maxillary palpi shorter than galea-lacinia ........................................ Callibaetis willineri Maxillary palpi longer than galea-lacinia (Fig. 29F) .................................................... 2 Mandibles with a tuft of setae between prostheca and mola (Figs. 29C-D) ........... ................................................................................................................... Callibaetis radiatus Mandibles without setae between prostheca and mola (Figs. 29Q-R) ..................... .................................................................................................................... Callibaetis guttatus

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CLAVES

PARA LOS

Imagos 1 – 2(1) – 3(2)

– 4(3) – 5(1) – 6(5) – 7(6) – 8(6)

–

6

CALLIBAETIS

SUDAMERICANOS

6

Alas anteriores con venas intercalares marginales simples, excepto por 1 o 2 celdas en el ápice o base del ala (Fig. 27A) ............................................................................... 2 Alas anteriores con venas intercalares marginales pares (Fig. 27E) ........................ 5 Alas en ambos sexos completamente hialinas (Fig. 28G) ............... Callibaetis pollens Alas por lo menos en las hembras con bandas oscura .............................................. 3 En las hembras, vitta sin ventanas claras alrededor de las venas transversas, banda pigmentada extendiéndose hacia la base de las alas en el margen anal (Fig. 27K); mitad del tercio basal de las alas posteriores teñidas de castaño (Fig. 27L) ............ ................................................................................................................. Callibaetis gonzalezi En las hembras, vitta con numerosas ventanas claras; banda pigmentada no extendiéndose hacia el margen anal (Figs. 27A, 28K); alas posteriores hialinas (Fig. 27B) ............................................................................................................................. 4 Alas posteriores con 7 a 9 venas transversas y proyección costal subcuadrangular (Fig. 27B) ................................................................................................. Callibaetis radiatus Alas posteriores con 12 a 15 venas transversas y proyección costal roma (Fig. 28L) .................................................................................................................. Callibaetis viviparus Alas anteriores, por lo menos en las hembras, con manchas a lo largo del margen posterior de las alas, a veces se extiende formando bandas (Figs. 28A-B) .............. .................................................................................................................... Callibaetis guttatus Alas anteriores con manchas o bandas pero nunca en el margen posterior (Figs. 27I, 28I) ................................................................................................................................ 6 Alas anteriores en ambos sexos, con bandas pigmentadas conspicuas, oblicuas o transversas, dejando áreas claras entre ellas (Figs. 27I, 28I) ..................................... 7 Alas anteriores sin bandas claramente definidas, si existe algún bandeo se presenta como líneas de ventanas en una ala pigmentada levemente (Figs. 27G, 28M) ....... 8 Alas anteriores con 3 bandas más o menos discretas (Fig. 27I) ................................ ................................................................................................................... Callibaetis fasciatus Alas anteriores con 2 bandas principales, la externa aproximadamente paralela al margen posterior (Fig. 28I) ...................................................................Callibaetis sellacki Alas posteriores sin venas transversas (Fig. 27H); en las hembras alas anteriores con vitta restringida a las áreas costal y subcostal, sin ventanas (Fig. 27G); en los machos tercio distal de las alas anteriores uniformemente teñido de castaño (Fig. 27E) ... .............................................................................................................. Callibaetis dominguezi Alas posteriores con 6 a 20 venas transversas; alas anteriores de las hembras con vitta extendiéndose por lo menos parcialmente al área posterior de la vena R1, con numerosas ventanas (Fig. 28E); en los machos alas hialinas o con una pequeña mancha en el estigma ........................................................................................................ 9

C. camposi, C. gregarius y C. nigrivenosus no fueron incluidos en la clave porque las descripciones originales no presentan caracteres suficientes para separar estas especies de las restantes.

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I IV

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H M

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Plate 28. Callibaetis, female imagos. Figs. 28A-D, C. guttatus : 28A, fore wing; 28B, idem, variation; 28C, hind wing; 28D idem, variation. Figs. 28E-F, C. jocosus : 28E, fore wing; 28F, hind wing. Figs. 28G-H, C. pollens : 28G, fore wing; 28H, hind wing. Figs. 28I-J, C. sellacki: 28I, fore wing; 28J, hind wing. Figs. 28K-L, C. viviparus : 28K, fore wing; 28L, hind wing. Figs. 28M-O, C. willineri : 28M, fore wing; 28N, hind wing; 28O, hind wing, detail. Figs. 28P-Q, C. zonalis : 28P, fore wing; 28Q, hind wing. (IV = intercalary veins; V = vitta). Figs. 28A, C, E-F, I-J, P-Q modified from GILLIES, 1990; 28B-C modified from DA-SILVA, 1991; 28G-H, K-L modified from NEEDHAM & MURPHY, 1924.

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9(8)

–

10(9)

–

Tibias II y III con 2 manchas oscuras en la mitad basal, además de una mancha subapical; esternos abdominales con manchas oscuras anterolaterales y sin manchas pares submedianas; alas anteriores de las hembras con pigmentación limitada a la vitta (Fig. 28P), en los machos alas anteriores no pigmentadas excepto el estigma algunas veces .................................................................................................. 10 Tibias II y III pueden ser oscuras pero sin dos manchas discretas en la mitad basal; esternos abdominales con pares de manchas oscuras submedianas además de manchas oscuras anterolaterales; alas anteriores en ambos sexos con vitta claramente pigmentada y el resto con pigmentación difusa, algunas con líneas claras alrededor de las venas transversas (Fig. 28E) ......................... Callibaetis jocosus En las hembras, alas anteriores con la mitad basal de la vitta cubriendo la vena R1 y hasta la R2 en menos de la mitad de su longitud (Fig. 28P); en los machos estigma de las alas anteriores con un parche de pigmento; esternos abdominales sin manchas en el medio ........................................................................ Callibaetis zonalis En las hembras, alas anteriores con la mitad basal de la vitta cubriendo la vena R1 y hasta la R2 en la mitad o más de su longitud (Fig. 28M); en los machos alas anteriores completamente hialinas; esternos abdominales II-VIII usualmente con una raya mediana castaña .................................................................... Callibaetis willineri

Ninfas 1 – 2(1) –

Palpos maxilares más cortos que la galea-lacinia ........................... Callibaetis willineri Palpos maxilares más largos que la galea-lacinia (Fig. 29F) ...................................... 2 Mandíbulas con un mechón de setas entre la prosteca y la mola (Figs. 29C-D) ................................................................................................................... Callibaetis radiatus Mandíbulas sin setas entre la prosteca y la mola (Figs. 29Q-R) ..... Callibaetis guttatus

Callibaetis camposi NAVÁS Callibaetis camposi NAVÁS, 1930b: 18; GILLIES, 1990: 22.

Taxonomy: This species, known from female imagos, was originally described by NAVÁS (1930b). GILLIES (1990) reviewed the type material and although it was in poor condition he considered it a well defined species. This species can be distinguished from the other species of the genus by the following combination of characters: 1) Fore wing with costal, subcostal and R1 areas strongly pigmented, with well marked windows and with discrete pigmentation posterior to them; 2) fore wing with single marginal intercalary veins; 3) hind wings with 11-15 cross veins and with a pointed costal process (Fig. 27D). Distribution: Ecuador. Callibaetis dominguezi GILLIES Callibaetis dominguezi GILLIES, 1990: 22.

Taxonomy: This species, known from male and female imagos, can be distinguished from the other species of the genus by the following combination of characters: 1) Fore wing of male (Fig. 27E) pigmented in distal 1/3 and fore wing of female (Fig. 27G) with

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vitta along the anterior border of wing; 2) fore wings with paired marginal intercalary veins (Figs. 27E, G); 3) hind wing without cross veins (Figs. 27F, H). Distribution: Argentina and Chile. Callibaetis fasciatus (PICTET) Cloe fasciata PICTET, 1843: 262. Callibaetis trifasciatus ESBEN-PETERSEN, 1912: 339. Baetis gloriosus NAVÁS, 1923b: 2. Callibaetis gloriosus NAVÁS, 1930d: 360. Callibaetis fasciatus EATON, 1885: 197; GILLIES, 1990: 23.

Taxonomy: This species known from adults, can be distinguished from the other species of the genus by the following combination of characters: 1) Fore wing of both sexes with 3 or more conspicuous transverse, pigmented bands (Fig. 27I); 2) fore wing with marginal intercalary veins paired (Fig. 27I); 3) hind wing with several cross veins and a small pointed costal process (Fig. 27J); 4) abdominal sterna I-IX with a pair of median black marks. Distribution: Argentina, Brazil, Chile and Uruguay. Callibaetis gonzalezi (NAVÁS) Cloeon gonzalezi NAVÁS, 1934a: 27. Callibaetis gonzalezi; GILLIES, 1990: 24.

Taxonomy: This species, known from female imagos, can be distinguished from the other species of the genus by the following combination of characters: 1) Vittae without clear windows around cross-veins, pigment extending broadly across base of wings to anal margin (Fig. 27K); 2) fore wing with marginal intercalary veins single (Fig. 27K); 3) hind wings with 13-15 cross veins, basal one-third to half of hind wings tinted brown and with a broadly rounded costal process (Fig. 27L). Distribution: Argentina and Paraguay. Callibaetis gregarius NAVÁS Callibaetis gregarius NAVÁS, 1930c: 72; GILLIES, 1990: 24.

Taxonomy: This species was originally described from male imagos by NAVÁS (1930c). GILLIES (1990) examined part of the type material and determined that the examination of more specimens would be necessary to clarify the status of this taxon. Distribution: Brazil. Callibaetis guttatus NAVÁS Callibaetis Callibaetis Callibaetis Callibaetis

guttatus NAVÁS, 1915a: 120; GILLIES, 1990: 25; DA-SILVA, 1991: 346. apicatus NAVÁS, 1917: 189. bruchius NAVÁS, 1920e: 55. zonatus NAVÁS, 1929: 224.

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A

P S

C

L

B

F

S

P

SL

K

D

E III D II

H I

G

L

I J M N

O P

Q

P P

R

T

S

Plate 29. Callibaetis, nymphs. Figs. 29A-P, C. radiatus . Figs. 29A-G, mouthparts: 29A, labrum, left d.v., right v.v.; 29B, labrum: anterior margin d.v.; 29C, left mandible v.v.; 29D, right mandible v.v.; 29E, hypopharynx; 29F, maxilla v.v.; 29G, labium, left d.v., right v.v. 29H, leg I; 29I, femur I: ventral margin; 29J, tibia I: apical margin; 29K, tarsal claw I; 29L, posterior margin of tergum IV; 29M, gill I; 29N, gill II; 29O, gill IV; 29P, paraproct. Figs. 29Q-S, C. guttatus : 29Q, left mandible; 29R, right mandible; 29S, maxilla. (D = denticles; L = lingua; P = prostheca; S = setae; SL = superlingua; T = tooth (canine); Roman numbers = segment numbers). Figs. 29Q-S modified from DA-SILVA, 1991.

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Taxonomy: This species, known from female imagos, was originally described by NAVÁS (1915a). Later GILLIES (1990) examined type material of C. apicatus, C. bruchius and C. zonatus and synonymized these species with C. guttatus. Finally, DA-SILVA (1991) described the nymphs for the first time and noted additional variation in the adults. This species can be distinguished from the other species of the genus by the following combination of characters. In the imago: 1) Fore wings, at least in females (Figs. 28A-B), with spots along posterior margin, sometimes extended forming bands; 2) fore wings with paired marginal intercalary veins (Fig. 28A); 3) hind wings with 6-10 cross veins and with a pointed costal process (Figs. 28C-D). Also, this species has two dark spots in basal half of tibia I and II. This species shows some variation according to recent the authors. GILLIES (1990) noted that both sexes show fore wings with extensive spotting over general wing field, occasionally reduced to a series of spots along posterior margin (Fig. 28A), while DA-SILVA (1991) described the fore wings in the females (Fig. 28B) with seven transverse bands and males with fore wings hyaline. In the nymphs: 1) Mandibles without tuft of setae between mola and prostheca (Figs. 29O-R); 2) maxillary palpi longer than galea-lacinia (Fig. 29S). Distribution and Biology: Argentina and Brazil. This species was collected in streams or temporary waters containing macrophytes. Callibaetis jocosus NAVÁS Callibaetis Callibaetis Callibaetis Callibaetis Callibaetis Callibaetis

jocosa NAVÁS, 1912b: 195. stictogaster NAVÁS, 1915a: 121. jaffueli NAVÁS, 1918: 244. spegazzinus NAVÁS, 1920d: 36. rimatus NAVÁS, 1932a: 113. jocosus GILLIES, 1990: 26.

Taxonomy: This species, was originally described by NAVÁS (1912b). GILLIES examined type material of C. stictogaster, C. jaffueli, C. spegazzinus and C. rimatus and synonymized these species with C. jocosus. This species, known from male and female imagos, can be distinguished from the other species of the genus by the following combination of characters: 1) Fore wings with heavly pigmented vitta and diffuse light pigmentation throughout rest of wing, sometimes with lines of clear windows around cross veins (Fig. 28E); 2) fore wings with paired marginal intercalary veins (Fig. 28E); 3) hind wings with 15-20 cross veins and with a pointed costal process (Fig. 28F). Distribution: Argentina, Brazil, Chile and Paraguay. Callibaetis nigrivenosus BANKS Callibaetis nigrivenosa BANKS, 1918: 11. Callibaetis nigrivenosus; MCCAFFERTY, 1996: 231.

Taxonomy: This species was described by BANKS (1918). The original description does not present specific characteristics to distinguish this species from the others. Callibaetis nigrivenosus has: 1) Fore wings with a faint dark tinge, very deep and prominent on apical costal area, all veins very dark; 2) fore wings with paired marginal intercalary veins. Distribution: Ecuador.

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Callibaetis pollens NEEDHAM & MURPHY `Callibaetis pollens NEEDHAM & MURPHY, 1924: 51.

Taxonomy: This species, known from adults, was originally described by NEEDHAM & MURPHY (1924). It can be distinguished from the other species of the genus by the following combination of characters: 1) Fore wings completely hyaline (Fig. 28G); 2) fore wings with single marginal intercalary veins (Fig. 28G); 3) hind wings with 10-15 cross veins and with pointed costal process (Fig. 28H). Distribution: Brazil. Callibaetis radiatus NAVÁS Callibaetis radiatus NAVÁS, 1920a: 132; GILLIES, 1990: 26; SALLES et al., 2003b: 13. Callibaetis venulosus NAVÁS, 1932a: 114.

Taxonomy: This species was originally described from male and female imagos by NAVÁS (1920a). GILLIES (1990) examined type material of C. venulosus and synonymized this species with C. radiatus. In 2003, SALLES et al. redescribed this species including the description of the nymphal stage. This species can be distinguished from the other species of the genus by the following combination of characters. In the adults: 1) Fore wings with vittae with numerous clear windows, pigment overlapping extreme bases of main veins, not extending to inner margin (Fig. 27A); 2) fore wings with most of the marginal intercalary veins single (Fig. 27A); 3) hind wings with 7-9 cross veins and a subquadrangular costal process (Fig. 27B). In the nymphs: 1) Mandibles with a tuft of setae between mola and prostheca (Figs. 29C-D); 2) maxillary palpi 1.5 times longer than galea-lacinia (Fig. 29F). Distribution and Biology: Argentina, Brazil and Paraguay. This species was collected in many different kinds of rivers or temporary waters and even in swimming pools. This species is tolerant of low oxygen concentrations. Callibaetis sellacki (WEYENBERGH) Cloe sellacki WEYENBERGH, 1883: 164. Cloe Lorentzii WEYENBERGH, 1883: 167. Callibaetis sellacki; EATON, 1885: 198; GILLIES, 1990: 27. Callibaetis trifasciatus [partim] NAVÁS, 1915a: 120. Callibaetis fasciatus ULMER, 1921: 246. Callibaetis lineatus NAVÁS, 1932b: 82.

Taxonomy: This species, known from adults, can be distinguished from the other species of the genus by the following combination of characters: 1) Fore wings in both sexes with 2 main pigmented bands, the outer band approximately parallel to posterior margin of wings (Fig. 28I); 2) fore wings with paired marginal intercalary veins (Fig. 28I); 3) hind wings with 15-20 cross veins and a small pointed costal process (Fig. 28J). Distribution: Argentina.

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Callibaetis viviparus NEEDHAM & MURPHY Callibaetis viviparus NEEDHAM & MURPHY, 1924: 50.

Taxonomy: This species, known from female imagos, can be distinguished from the other species of the genus by the following combination of characters: 1) Fore wings with vitta having open windows in places where costal cross veins normally occur (Fig. 28K); 2) fore wings with single marginal intercalary veins (Fig. 28K); 3) hind wings with 12-15 cross veins and a broadly rounded costal process (Fig. 28L). Distribution: Brazil. Callibaetis willineri NAVÁS Callibaetis willineri NAVÁS, 1932a: 115; GILLIES, 1990: 30. Callibaetis alegre TRAVER, 1944: 46.3

Taxonomy: This species was originally described from adults by NAVÁS (1932a). Later, GILLIES (1990) synonymized C. alegre, known from nymphs and adults, with this species. C. willineri can be distinguished from the other species of the genus by the following combination of characters. In the adults: 1) Female fore wings (Fig. 28M) with basal half of vitta overlapping vein R1 to as far as R2 in 1/2 or more of its length, males with fore wings entirely hyaline; 2) fore wings with paired marginal intercalary veins (Fig. 28M); 3) hind wings with 6-10 cross veins and a large broadly rounded costal process (Figs. 28N-O). This species also has a median, maroon dash on sterna II-VIII. In the nymphs: 1) Mandibles with a tuft of setae between mola and prostheca; 2) maxillary palpi shorter than galea-lacinia. Distribution: Argentina, Brazil and Uruguay. Callibaetis zonalis NAVÁS Callibaetis zonalis NAVÁS, 1915b: 13; GILLIES, 1990: 31. Baetis opacus NAVÁS, 1915b: 12. Baetis virellus NAVÁS, 1915a: 119. Callibaetis vitreus NAVÁS, 1915a: 121; NAVÁS, 1919: 81. Callibaetis sobrius NAVÁS, 1916: 61. Callibaetis apertus NAVÁS, 1917: 190. Callibaetis depressus NAVÁS, 1922a: 59. Callibaetis amoenus NAVÁS, 1930e: 131.

Taxonomy: This species, was originally described from adults by NAVÁS (1915b). GILLIES examined type material of Baetis opacus, Baetis virellus, Callibaetis vitreus, C. sobrius, C. apertus, C. depressus and C. amoenus and synonymized these species with C. zonalis. This species, can be distinguished from the other species of the genus by the following combination of characters: 1) Female fore wings (Fig. 28P) with basal half of vitta overlapping vein R1 to as far as R2 in less than 1/2 of its length, male fore wings with a patch of smoky pigment; 2) fore wings with paired marginal intercalary veins (Fig. 28P); 3) hind wings with 15-17 cross veins and a small pointed costal process (Fig. 28Q). Distribution: Argentina, Brazil and Paraguay.

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Callibaetis abundans (NAVÁS) Nomen Dubium Baetis abundans NAVÁS, 1912b: 194. Callibaetis abundans; ULMER, 1943: 34; GILLIES, 1990: 20.

Taxonomy: This species was reviewed by GILLIES (1990) and he treated it as a nomen dubium. Distribution: Paraguay. Genus Camelobaetidius DEMOULIN Camelobaetidius DEMOULIN, 1966a: 9; MCCAFFERTY & WALTZ, 1990: 777; LUGO-ORTIZ & MCCAFFERTY, 1999a: 258; DOMINIQUE et al., 2000: 40; DOMINIQUE et al., 2001: 18; DOMINIQUE & THOMAS, 2001: 27; NIETO, 2002: 57; NIETO, 2003a: 233. (Type-species: Camelobaetidius leentvaari DEMOULIN, original designation). Dactylobaetis TRAVER & EDMUNDS, 1968: 629. (Type-species: Dactylobaetis warreni TRAVER & EDMUNDS, original designation).

Taxonomy: Camelobaetidius was established by DEMOULIN (1966a) and Dactylobaetis by TRAVER & EDMUNDS (1968). MCCAFFERTY & WALTZ (1990) synonymized Dactylobaetis with Camelobaetidius. This genus is widely distributed and is one of the most speciose genera among the Baetidae. South America has 24 known species with 16 known from nymphs, 4 from adults, and 4 from both nymphs and adults. Adult Characteristics: 1) Fore wings with paired marginal intercalary veins (Fig. 30A) and with anal margin rounded; 2) hind wings with two longitudinal veins and with a pointed costal process (Figs. 30B-C); 3) forceps three-segmented, segment III elongate (Fig. 30E). Nymphal Characteristics (Fig. 31A): 1) Head wider than long; 2) antennae 1.52.5 times of head capsule, scape subequal in length to pedicel; 3) labrum (Fig. 32A) anterior margin with two kinds of bipectinated setae: basally bifid near midline and apically bifid near lateral margin; 4) mandibles (Figs. 32B-C) with incisors fused apically, prosthecae robust with denticles apically; 5) hypopharynx (Fig. 32D) with lingua subequal in length to superlingua; 6) maxillae (Fig. 32E) with palpi twosegmented, shorter than or subequal in length to galea-lacinia; 7) labium (Fig. 32F): glossae subequal in length and half the width of paraglossae, glossae and paraglossae with a row of spine-like setae, segment III of palpi rounded (Fig. 32G); 8) legs robust (Figs. 32H, J), dorsal edge of femora with a row of long spine-like setae, ventral edge of tarsi with a row of setae, the most apical setae longer than the others; 9) tarsal claws spatulate, with a fan-shaped row of denticles; 10) foreleg with (Fig. 32H) or without (Fig. 34M) a single finger-like of coxal gill, between coxa I and prosternum; 11) hind wing pads present; 12) posterior margin of abdominal terga with rounded spines (Fig. 32K); 12) abdominal gills present on segments I-VII, gill I small; 13) terminal filament subequal or extremely short with respect to cerci. Distribution: North, Central and South America. Biology: Species of Camelobaetidius can be found in many different habitats. The nymphs were collected in rivers or streams with rocky or sandy substrate.

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H Plate 30. Camelobaetidius, male imagos. Figs. 30A-E, C. anubis: 30A, fore wing; 30B, hind wing; 30C, hind wing, detail; 30D, abdomen d.v.; 30E, genitalia v.v. Fig. 30F, C. penai, genitalia v.v. Figs. 30G-I, C. phaedrus: 30G, hind wing; 30H, abdomen d.v.; 30I, genitalia v.v. Figs. 30J-L, C. suapi: 30J, hind wing; 30K, abdomen d.v.; 30L, genitalia v.v. (CP = Costal Process; Roman numbers = segment numbers). Figs. 30A-I modified from NIETO, 2003a; 30J-L from NIETO, 2002.

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Male Imagos7 1 – 2(1) – 3(2) – 4(3) –

Genitalia: basal segment of forceps short (almost as long as wide) (Fig. 30E) ....... 2 Genitalia: basal segment of forceps long (1.5 times longer than wide) (Fig. 30F) ............................................................................................................... Camelobaetidius penai Genitalia: segment II of forceps strongly curved (Fig. 30E) ...... Camelobaetidius anubis Genitalia: segment II of forceps not as above (Fig. 30I) .......................................... 3 Abdominal color pattern as in Fig. 30H; hind wings with 3 longitudinal veins (Fig. 30G) ..................................................................................... Camelobaetidius phaedrus Abdominal color pattern not as above; hind wings with 2 longitudinal veins (Fig. 30J) .............................................................................................................................. 4 Abdominal segments with spots on segments V-VI; hind wings with posterior margin convex (as in Fig. 30G) ........................................................ Camelobaetidius billi Abdominal segments without spots (Fig. 30K); hind wings with posterior margin concave basally (Fig. 30J) ................................................................ Camelobaetidius suapi

Nymphs 1 – 2(1) – 3(2) – 4(1) – 5(4) – 6(5) – 7(5) – 7

Femur I with a tubercle on ventral margin at base (Fig. 35D); apical margin of tibia with an excavation and a patch of short spines (Fig. 35E) ............................. 2 Femur I without tubercles on ventral margin (Fig. 33D); apical margin of tibia without excavation or patch of spines .......................................................................... 4 Coxal gills although small present; prosternum with single, medial protuberance ........................................................................................................ Camelobaetidius leentvaari Coxal gills absent; prosternum without protuberance ............................................... 3 Tarsal claw with 25 denticles; abdominal gills with main branch of trachea and a few lateral branches darkened; general coloration pale yellow .................................. ............................................................................................................. Camelobaetidius mantis Tarsal claw with 30 or 31 denticles (Fig. 35F); abdominal gills whitish (Fig. 35I); general coloration dark brown ....................................................... Camelobaetidius ipaye Coxal gills present (Fig. 32H) .......................................................................................... 5 Coxal gills absent (Fig. 34M) ......................................................................................... 14 Terminal filament short, subequal in length to abdominal segment X (Fig. 33F) ...... 6 Terminal filament long, subequal in length to cerci ................................................... 7 Paraprocts with posterior margin smooth as in Fig. 33I; posterior margin of abdominal terga as in Fig. 33G ........................................................ Camelobaetidius apis Paraprocts with 4 or 5 small projections and five-six larger projections on posterior margin as in Fig. 33M; posterior margin of abdominal terga as in Fig. 33L ......................................................................................................... Camelobaetidius billi Pronotum with medial pair of tubercles ................................ Camelobaetidius tuberosus Pronotum without tubercles ............................................................................................ 8

C. alcyoneus, C. coveloae, C. dryops, and C. tantillus were not included in the key because the original descriptions do not have enough characters to distinguish these species from the others.

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Plate 31. Camelobaetidius phaedrus. Fig. 31A, nymphal habitus. Drawing by C. Molineri.

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Maxillae with segment II of palpi bifid (Figs. 35L-M) ............. Camelobaetidius janae Maxillae with segment II of palpi simple (Fig. 32E) .................................................. 9 Tarsal claws with less than 10 denticles ...................................................................... 10 Tarsal claws with more than 15 denticles ................................................................... 11 General abdominal color pattern as in Figs. 31A, 38H; mandibles with incisors fused, strongly extended, and apically pointed as in Figs. 38B-C; coxal gills long (Fig. 38F), subequal to coxa and trochanter combined; labrum (Fig. 38A), dorsally with a continuous subapical row of very long setae ........... Camelobaetidius phaedrus General abdominal color pattern not as above; mandibles with incisors normal (similar to Figs. 32B-C); coxal gills short (Fig. 37C), subequal in length to coxa; labrum (Fig. 37A), dorsally with only two-three long subapical setae placed laterally ........................................................................................................... Camelobaetidius patricki Tarsal claws with first denticle at least 1.3 times longer than the others (Fig. 39O) .............................................................................................................................................. 12 Tarsal claws with first denticle subequal to the others (Fig. 32I) .......................... 13 Tarsal claws with 16 to 18 denticles (Fig. 39O); segment II of labial palpi with pointed distomedial projection as in Fig. 39M; posterior margin of abdominal terga as in Fig. 39P ..................................................................... Camelobaetidius yacutinga Tarsal claws with 25 to 28 denticles (Fig. 36C); segment II of labial palpi with a rounded distomedial projection as in Fig. 36B; posterior margin of abdominal terga as in Fig. 36D ................................................................... Camelobaetidius mathuriae Tarsal claws with 20 to 28 denticles (Fig. 32I); coxal gills long, subequal to coxa and trochanter combined (Fig. 32H) .......................................... Camelobaetidius anubis Tarsal claws with 30 to 40 denticles; coxal gills shorter than coxa and trochanter combined .......................................................................................... Camelobaetidius serapis Tarsal claws with the first denticle distinctly longer than the others (Fig. 38O) .............................................................................................................................................. 15 Tarsal claws with the first denticle about equal to the others (Fig. 36N) ............ 16 Labium: segment II of palpi with a small medial digitiform projection (Fig. 38N); tarsal claw with the first denticle 1.4 times longer than the others (Fig. 38O); paraprocts as in Fig. 38R; abdominal gills with main branch of trachea darkened (Fig. 38Q) ............................................................................................ Camelobaetidius suapi Labium: segment II of palpi with a strong pointed distomedial projection (Fig. 34L); tarsal claw with the first denticle 1.2 times longer than the others (Fig. 34N); paraproct as in Fig. 34Q; abdominal gills whitish (Fig. 34P) ....................... ............................................................................................................ Camelobaetidius huarpe Labium: segment II of palpi with rounded distomedial projection (Figs. 36G, M) .............................................................................................................................................. 17 Labium: segment II of palpi with a strong pointed distomedial projection (Fig. 37I) ............................................................................................................................ 18 Tarsal claws with 13 to 18 denticles (Fig. 36H); segment II of labial palpi as in Fig. 36G; labrum, subapical setae as in Fig. 36F .............................. Camelobaetidius matilei Tarsal claws with 23 to 24 denticles (Fig. 36N); segment II of labial palpi as in Fig. 36M; labrum, subapical setae as in Fig. 36L ............................... Camelobaetidius ortizi

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Plate 32. Camelobaetidius anubis, nymph. Figs. 32A-M. Figs. 32A-G, mouthparts: 32A, labrum d.v.; 32B, left mandible v.v.; 32C, right mandible v.v.; 32D, hypopharynx v.v.; 32E, maxilla v.v.; 32F, labium, left d.v., right v.v.; 32G, labial palp detail. 32H, leg I; 32I, tarsal claw I; 32J, leg II; 32K, posterior margin of abdominal tergum IV; 32L, gill IV; 32M, paraproct. (CG = coxal gill; L = lingua; P = prostheca; S = apical seta; SL = superlingua; Roman numbers = segment numbers). Figs. 32A-M modified from NIETO, 2003a.

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18(16) Segment II of labial palpi with a strong rounded distomedial projection as in Fig. 34F; tarsal claws with 34 to 39 denticles (Fig. 34G) ............ Camelobaetidius edmundsi Segment II of labial palpi with a strong pointed distomedial projection as in Fig. 37I; tarsal claws with 30 or less denticles ................................................................... 19 19(18) Tarsal claws with 16 to 18 denticles (Fig. 34C); segment II of labial palpi as in Fig. 34B; posterior margin of paraprocts as in Fig. 34D ............ Camelobaetidius cayumba Tarsal claws with 20 to 30 denticles (Fig. 37J); segment II of labial palpi as in Fig. 37I; posterior margin of paraprocts as in Fig. 37M .................. Camelobaetidius penai CLAVES

PARA LOS

Imagos Machos 1 – 2(1) – 3(2) – 4(3) –

CAMELOBAETIDIUS

SUDAMERICANOS

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Genitalia: segmento I de los fórceps corto (casi tan largo como ancho) (Fig. 30E) ................................................................................................................................................ 2 Genitalia: segmento I de los fórceps largo (1,5 veces más largo que ancho) (Fig. 30F) ............................................................................................ Camelobaetidius penai Genitalia: segmento II de los fórceps fuertemente arqueado (Fig. 30E) ................. ............................................................................................................. Camelobaetidius anubis Genitalia: segmento II de los fórceps levemente arqueado (Fig. 30I) .................... 3 Patrón de coloración abdominal como en la Fig. 30H; alas posteriores con 3 venas longitudinales (Fig. 30G) ........................................................... Camelobaetidius phaedrus Patrón de coloración abdominal no como arriba; alas posteriores con 2 venas longitudinales (Fig. 30J) .................................................................................................... 4 Segmentos abdominales con manchas en los segmentos V-VI; margen posterior de las alas posteriores convexo (como en la Fig. 30G) ............... Camelobaetidius billi Segmentos abdominales sin manchas (Fig. 30K); margen posterior de las alas posteriores cóncavo basalmente (Fig. 30J) .................................. Camelobaetidius suapi

Ninfas 1 – 2(1) – 3(2) –

8

Fémur I con un tubérculo en el margen ventral (Fig. 35D); margen apical de la tibia con una hendidura y un parche de espinas (Fig. 35E) ...................................... 2 Fémur I sin tubérculo en el margen ventral (Fig. 33D); margen apical de la tibia sin hendidura ni parches de espinas ..................................................................................... 4 Branquias coxales, aunque pequeñas presentes; proesterno con una protuberancia medial ........................................................................................... Camelobaetidius leentvaari Branquias coxales ausentes; proesterno sin protuberancia ....................................... 3 Uñas tarsales con 25 dentículos; branquias abdominales con la tráquea principal y unas pocas secundarias oscurecidas; coloración general amarillo pálido ................. ............................................................................................................. Camelobaetidius mantis Uñas tarsales con 30 o 31 dentículos (Fig. 35F); branquias abdominales blancuzcas (Fig. 35I); coloración general castaño-oscura .............................. Camelobaetidius ipaye

C. alcyoneus, C. coveloae, C. dryops, C. tantillus no fueron incluidos en la clave porque las descripciones originales no presentan caracteres suficientes para separar estas especies de las restantes.

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K Plate 33. Camelobaetidius, nymphs. Figs. 33A-I, C. apis : 33A, labrum d.v.; 33B, labium, left d.v., right v.v.; 33C, labial palp detail; 33D, leg I; 33E, tarsal claw I; 33F, abdomen d.v.; 33G, posterior margin of abdominal tergum IV; 33H, gill IV; 33I, paraproct. Figs. 33J-M, C. billi: 33J, proximal part of the leg I; 33K, tarsal claw I; 33L, posterior margin of abdominal tergum IV; 33M, paraproct. (CG = coxal gill; TF = terminal filament; Roman numbers = segment numbers). Figs. 33A-I modified from NIETO, 2003a; 33J-M, from DOMINIQUE et al., 2000.

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Branquias coxales presentes (Fig. 32H) ......................................................................... 5 Branquias coxales ausentes (Fig. 34M) ........................................................................ 14 Filamento terminal corto, subigual a la longitud del X segmento abdominal (Fig. 33F) ............................................................................................................................. 6 – Filamento terminal largo, subigual a la longitud de los cercos ................................ 7 6(5) Márgenes posteriores de los paraproctos lisos como en la Fig. 33I; márgenes posteriores de los tergos abdominales como en la Fig. 33G ..... Camelobaetidius apis – Paraproctos con 4 o 5 espinas pequeñas y cinco-seis espinas más grandes como en la Fig. 33M; márgenes posteriores de los tergos abdominales como en la Fig. 33L ................................................................................................................. Camelobaetidius billi 7(5) Pronoto con un par de tubérculos mediales ......................... Camelobaetidius tuberosus – Pronoto sin tubérculos ...................................................................................................... 8 8(7) Maxilas con el segmento II de los palpos bífidos (Figs. 33L, 35M) ......................... ............................................................................................................... Camelobaetidius janae – Maxilas con el segmento II de los palpos simples (Fig. 32E) .................................. 9 9(8) Uñas tarsales con menos de 10 dentículos ................................................................. 10 – Uñas tarsales con más de 15 dentículos ...................................................................... 11 10(9) Patrón de coloración abdominal como en las Figs. 31A, 38H; mandíbulas con incisivos fusionados fuertemente extendidos y protuidos como en la Figs. 38B-C; branquias coxales largas (Fig. 38F), subiguales a la longitud de la coxa y el trocánter juntos; labro (Fig. 38A), dorsalmente con 1 hilera subapical continua de setas largas subapicales .............................................................. Camelobaetidius phaedrus – Patrón de coloración abdominal no como arriba; mandíbulas con incisivos normales (similar a las Figs. 32B-C); branquias coxales cortas (Fig. 37C), subiguales a la longitud de las coxas; labro (Fig. 37A), dorsalmente con 2-3 setas subapicales ubicadas lateralmente .............................................. Camelobaetidius patricki 11(9) Uñas tarsales con el primer dentículo por lo menos 1,3 veces más largo que los restantes (Fig. 39O) ......................................................................................................... 12 – Uñas tarsales con el primer dentículo subigual a los restantes (Fig. 32I) ............ 13 12(11) Uñas tarsales con 16 a 18 dentículos (Fig. 39O); segmento II del palpo labial con una proyección aguda distomedial como en la Fig. 39M; márgenes posteriores de los tergos abdominales como en la Fig. 39P ......................... Camelobaetidius yacutinga – Uñas tarsales con 25 a 28 dentículos (Fig. 36C); segmento II del palpo labial con una proyección distomedial redondeada (Fig. 36B); márgenes posteriores de los tergos abdominales como en la Fig. 36D ............................ Camelobaetidius mathuriae 13(11) Uñas tarsales con 20 a 28 dentículos (Fig. 32I); branquias coxales largas (Fig. 32H), subiguales a la longitud de la coxa y el trocánter juntos ........ Camelobaetidius anubis – Uñas tarsales con 30 a 40 dentículos; branquias coxales más cortas que la longitud de la coxa y el trocanter juntos .................................................... Camelobaetidius serapis 14(4) Uñas tarsales con el primer dentículo claramente más largo que los restantes (Fig. 38O) .......................................................................................................................... 15 – Uñas tarsales con el primer dentículo subigual a los restantes (Fig. 36N) .......... 16 15(14) Labio: segmento II del palpo con una proyección pequeña medial digitiforme (Fig. 38N); uñas tarsales con el primer dentículo 1,4 más largo que los restantes

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(Fig. 38O); paraproctos como en la Fig. 38R; branquias abdominales con la tráquea principal oscurecida (Fig. 38Q) ....................................... Camelobaetidius suapi Labio: segmento II del palpo con una fuerte proyección distomedial aguda (Fig. 34L); uñas tarsales con el primer dentículo 1,2 más largo que los restantes (Fig. 34N); paraproctos como en la Fig. 34Q; branquias abdominales blancuzcas (Fig. 34P) ................................................................... Camelobaetidius huarpe Labio: segmento II del palpo con una proyección distomedial redondeada (Figs. 36G, M) .............................................................................................................................. 17 Labio: segmento II del palpo con una fuerte proyección distomedial aguda (Fig. 37I) ............................................................................................................................ 18 Uñas tarsales con 13 a 18 dentículos (Fig. 36H); segmento II del palpo labial como en la Fig. 36G; labro con setas subapicales como en la Fig. 36F .............................. .............................................................................................................Camelobaetidius matilei Uñas tarsales con 23 a 24 dentículos (Fig. 36N); segmento II del palpo labial como en la Fig. 36M; labro con setas subapicales como en la Fig. 36L .............................. ...............................................................................................................Camelobaetidius ortizi Segmento II del palpo labial con una fuerte proyección distomedial redondeada como en la Fig. 34F; uñas tarsales con 34 a 39 dentículos (Fig. 34G) ..................... ......................................................................................................... Camelobaetidius edmundsi Segmento II del palpo labial con una fuerte proyección distomedial aguda como en la Fig. 37I; uñas tarsales con 30 o menos dentículos .......................................... 19 Uñas tarsales con 16 a 18 dentículos (Fig. 34C); segmento II del palpo labial como en la Fig. 34B; márgenes posteriores de los paraproctos como en la Fig. 34D ..... .......................................................................................................... Camelobaetidius cayumba Uñas tarsales con 20 a 30 dentículos (Fig. 37J); segmento II del palpo labial como en la Fig. 37I; márgenes posteriores de los paraproctos como en la Fig. 37M ...... ............................................................................................................... Camelobaetidius penai

Camelobaetidius alcyoneus (TRAVER) Baetis alcyoneus TRAVER, 1943: 85. Camelobaetidius alcyoneus; LUGO-ORTIZ & MCCAFFERTY, 1999a: 258.

Taxonomy: This species was originally described from adults by TRAVER (1943). After studying the hind wing illustration, LUGO-ORTIZ & MCCAFFERTY (1999a) transferred it to Camelobaetidius. Given that this shape of hind wing is not restricted to the genus, until this species is reared from a spatulate clawed nymph its position in Camelobaetidius will remain dubious. Distribution: Venezuela. Camelobaetidius anubis (TRAVER & EDMUNDS) Dactylobaetis anubis TRAVER & EDMUNDS, 1968: 666. Camelobaetidius anubis; MCCAFFERTY & WALTZ, 1990: 783; NIETO, 2003a: 244.

Taxonomy: This species was described from nymphs by TRAVER & EDMUNDS (1968). Later, NIETO (2003a) described the male imago. This species can be distinguished from the

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other species of the genus by the following combination of characters. In the adults: 1) Abdominal color pattern as in Fig. 30D; 2) basal segment of forceps short (Fig. 30E), as long as wide, segment II strongly curved; 3) turbinate eyes with dark margin. In the nymphs: 1) Labrum (Fig. 32A), dorsally with a pair of long subapical setae near midline and 2-3 long setae near lateral margin; 2) segment II of labial palpi with a strong pointed distomedial projection (Fig. 32G); 3) coxal gills long, subequal to coxa and trochanter combined (Fig. 32H); 4) tarsal claws with 20-28 denticles (Fig. 32I); 5) posterior margin of abdominal terga as in Fig. 32K; 6) abdominal gills with main branch of trachea darkened (Fig. 32L); 7) paraprocts as in Fig. 32M; 8) terminal filament subequal in length to cerci. Distribution: Argentina (Misiones), Brazil (Paraná). Camelobaetidius apis NIETO Camelobaetidius apis NIETO, 2003a: 237.

Taxonomy: This species, known from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Labrum (Fig. 33A), dorsally with a subapical row of long setae; 2) segment II of labial palpi rounded medially (Figs. 33B-C); 3) coxal gills shorter than coxae (Fig. 33D); 4) tarsal claws with 30-31 denticles (Fig. 33E); 5) posterior margin of abdominal terga as in Fig. 33G; 6) abdominal gills whitish (Fig. 33H); 7) paraprocts as in Fig. 33I; 8) terminal filament subequal in length to abdominal segment X (Fig. 33F). Distribution and Biology: Argentina: Misiones. This species was found in the Paranaense area, in rivers of rocky substrate. Camelobaetidius billi DOMINIQUE, THOMAS, ORTH & DAUTA Camelobaetidius billi DOMINIQUE et al., 2000: 40; SALLES & DIAS, 2004: 269.

Taxonomy: This species was originally described from nymphs by DOMINIQUE et al. (2000). The adults were later described by SALLES & DIAS (2004). This species can be distinguished from the other species of the genus by the following combination of characters. In the adults: 1) Abdominal color pattern with spots frequently on segments V-VI; 2) basal segment of forceps short, as long as wide, segment II slightly curved; 3) hind wings with posterior margin convex and with two longitudinal veins (Fig. 30G). In the nymphs: 1) Labrum, dorsally with a subapical row of long setae (similar to Fig. 33A); 2) segment II of labial palpi rounded medially (similar to Fig. 33C); 3) coxal gills shorter than coxa (Fig. 33J); 4) tarsal claws with 30-31 denticles (Fig. 33K); 5) posterior margin of abdominal terga as in Fig. 33L; 6) abdominal gills whitish; 7) paraprocts as in Fig. 33M; 8) terminal filament subequal in length to abdominal segment X (similar to Fig. 33F). Distribution: Brazil and French Guiana. Camelobaetidius cayumba (TRAVER & EDMUNDS) Dactylobaetis cayumba TRAVER & EDMUNDS, 1968: 669. Camelobaetidius cayumba; THOMAS et al., 2001a: 118; DOMÍNGUEZ et al., 2002: 462.

Taxonomy: This species was originally described from nymphs by TRAVER & EDMUNDS (1968). Later THOMAS et al. (2001a) redescribed this species. Camelobaetidius cayumba can be

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distinguished from the other species of the genus by the following combination of characters: 1) Labrum (Fig. 34A), dorsally with a pair of subapical setae and 2-3 setae near lateral margin; 2) segment II of labial palpi with a pointed distomedial projection (Fig. 34B); 3) coxal gills absent; 4) tarsal claws with 16-18 denticles (Fig. 34C); 5) abdominal gills with main branch of trachea darkened; 6) paraprocts as in Fig. 34D; 7) terminal filament subequal in length to cerci. Distribution: Colombia, French Guiana and Peru. Camelobaetidius coveloae (TRAVER) Baetis coveloae TRAVER, 1971: 61. Camelobaetidius coveloae; LUGO-ORTIZ & MCCAFFERTY, 1999a: 258.

Taxonomy: This species was described from male imagos by TRAVER (1971). After studying the illustrations of the hind wing, LUGO-ORTIZ & MCCAFFERTY (1999a) transferred this species to Camelobaetidius. Given that this shape of hind wing is not restricted to this genus, until this species is reared its position in Camelobaetidius will remain dubious. Distribution: Uruguay. Camelobaetidius dryops (NEEDHAM & MURPHY) Baetis dryops NEEDHAM & MURPHY, 1924: 53. Camelobaetidius dryops; LUGO-ORTIZ & MCCAFFERTY, 1999a: 258.

Taxonomy: This species was originally described from adults by NEEDHAM & MURPHY (1924). LUGO-ORTIZ & MCCAFFERTY (1999a) examined the holotype and transferred this species to Camelobaetidius. The characters used as the basis for the transfer were the shape of hind wings and the elongated terminal forceps, characters which are not restricted to this genus. For this reason, until this species is reared its position in Camelobaetidius will remain dubious. Distribution: Peru. Camelobaetidius edmundsi DOMINIQUE, MATHURIAU & THOMAS Camelobaetidius edmundsi DOMINIQUE et al., 2001: 19.

Taxonomy: This species, known from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Labrum (Fig. 34E), dorsally with a pair of subapical setae near midline and 2-3 setae near lateral margin; 2) segment II of labial palpi with a strong rounded distomedial blunt projection (Fig. 34F); 3) coxal gills absent; 4) tarsal claws with 34-39 denticles (Fig. 34G); 5) posterior margin of abdominal terga as in Fig. 34H; 6) abdominal gills whitish; 7) paraprocts as in Fig. 34I; 8) terminal filament subequal in length to cerci. Distribution: Colombia. Camelobaetidius huarpe NIETO Camelobaetidius huarpe NIETO, 2003a: 249.

Taxonomy: This species, known from nymphs and the female imago, can be distinguished from the other species of the genus by the following combination of

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Plate 34. Camelobaetidius, nymphs. Figs. 34A-D, C. cayumba: 34A, labrum d.v.; 34B, labium, left d.v., right v.v.; 34C, tarsal claw I; 34D, paraproct. Figs. 34E-I, C. edmundsi: 34E, labrum d.v.; 34F, labium, left d.v., right v.v.; 34G, tarsal claw I; 34H, posterior margin of abdominal tergum IV; 34I, paraproct. Figs. 34J-Q, C. huarpe: 34J, labrum d.v.; 34K, labium, left d.v., right v.v.; 34L, labial palp detail; 34M, leg I; 34N, tarsal claw I; 34O, posterior margin of abdominal tergum IV; 34P, gill IV; 34Q, paraproct. (Roman numbers = segment numbers). Figs. 34A-D modified from THOMAS et al., 2001a; 34E-F, from DOMINIQUE et al., 2001; 34J-Q from NIETO, 2003a.

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characters. In the nymphs: 1) Labrum (Fig. 34J), dorsally with a pair of long subapical setae near midline and 2-3 long setae near lateral margin; 2) segment II of labial palpi with a strong pointed distomedial projection (Figs. 34K-L); 3) coxal gills absent (Fig. 34M); 4) tarsal claws with 20-21 denticles, the first denticle 1.2 times longer than the others (Fig. 34N); 5) posterior margin of abdominal terga as in Fig. 34O; 6) abdominal gills whitish (Fig. 34P); 7) paraprocts as in Fig. 34Q; 8) terminal filament subequal in length to cerci. Distribution: Argentina (San Juan). Camelobaetidius ipaye NIETO Camelobaetidius ipaye NIETO, 2003a: 235.

Taxonomy: This species known from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Labrum (Fig. 35A), dorsally with a subapical row of setae; 2) segment II of labial palpi with a rounded projection (Figs. 35B-C); 3) coxal gills absent; 4) tarsal claws with 30-31 denticles (Fig. 35F); 5) posterior margin of abdominal terga as in Fig. 35H; 6) abdominal gills whitish (Fig. 35I); 7) paraprocts as in Fig. 35J; 8) terminal filament subequal in length to abdominal segment X; 9) femur I with a tubercle on ventral margin at base (Fig. 35D); 10) apical margin of tibiae with an excavation and a patch of short spines (Fig. 35E); 11) general coloration dark brown (Fig. 35G). Distribution and Biology: Argentina (Misiones). This species was found in the Paranaense area, in rivers of rocky substrate. Camelobaetidius janae DOMINIQUE, THOMAS, ORTH & DAUTA Camelobaetidius janae DOMINIQUE et al., 2000: 44; SALLES et al., 2004: 5.

Taxonomy: This species, known from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Labrum (Fig. 35K), dorsally with 2 subapical setae near midline and 3-4 setae near lateral margin; 2) segment II of labial palpi with a broadly rounded distomedial projection (Figs. 35N-O); 3) coxal gills absent; 4) tarsal claws with 21 denticles (Fig. 35P); 5) posterior margin of abdominal terga as in Fig. 35Q; 6) abdominal gills whitish; 7) paraprocts as in Fig. 35R; 8) terminal filament subequal in length to cerci; 9) maxillae with segment II of palpi bifid (Figs. 35L-M). Distribution: French Guiana and Brazil. Camelobaetidius leentvaari DEMOULIN Camelobaetidius leentvaari DEMOULIN, 1966a: 9; SALLES et al., 2005: 70.

Taxonomy: This species was described from nymphs by DEMOULIN (1966a). Later, SALLES et al. (2005) examined the type material and redescribed this species. This species can be distinguished from the other species of the genus by the following combination of characters: 1) Labrum, dorsally with a subapical row of setae (similar to Fig. 35A); 2) segment II of labial palpi medially rounded (similar to Fig. 35C); 3) coxal gills although small, present; 4) tarsal claws with 17-23 denticles; 5) abdominal gills whitish; 6) terminal

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Plate 35. Camelobaetidius, nymphs. Figs. 35A-J, C. ipaye: 35A, labrum d.v.; 35B, labium, left d.v., right v.v.; 35C, labial palp detail; 35D, leg I; 35E, apical margin of tibia I; 35F, tarsal claw I; 35G, abdomen d.v.; 35H, posterior margin of abdominal tergum IV; 35I, gill IV; 35J, paraproct. Figs. 35K-R, C. janae: 35K, labrum d.v.; 35L, maxilla v.v.; 35M, maxillary palp detail; 35N, labium v.v.; 35O, labial palp detail; 35P, tarsal claw I; 35Q, posterior margin of abdominal tergum IV; 35R, paraproct. (E = excavation; P = spine patch; T = tubercle; Roman numbers = segment numbers). Figs. 35A-J modified from NIETO, 2003a; 35K-R from DOMINIQUE et al., 2000.

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filament subequal in length to abdominal segment X; 7) femur I with a tubercle on ventral margin at base (similar to Fig. 35D); 8) apical margin of tibia with an excavation and a patch of short spines (similar to Fig. 35E); 9) prosternum with single, medial protuberance. Distribution: Brazil and Surinam. Camelobaetidius mantis TRAVER & EDMUNDS Camelobaetidius mantis TRAVER & EDMUNDS, 1968: 675; DOMÍNGUEZ et al., 2002: 462.

Taxonomy: This species, known from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Labrum, dorsally with a subapical row of setae (similar to Fig. 35A); 2) segment II of labial palp medially rounded (similar to Fig. 35C); 3) coxal gills absent; 4) tarsal claws with 25 denticles; 5) abdominal gills with main trachea and a few lateral branches darkened; 6) terminal filament half length of abdominal segment X; 7) femur I with a tubercle on ventral margin at base (similar to Fig. 35D); 8) apical margin of tibia with an excavation and with a patch of short spines (similar to Fig. 35E); 9) general coloration pale yellow. Distribution: Brazil and Colombia. Camelobaetidius mathuriae DOMINIQUE & THOMAS Camelobaetidius mathuriae DOMINIQUE & THOMAS (in DOMINIQUE et al., 2001: 26).

Taxonomy: This species, known from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Labrum (Fig. 36A), dorsally with a pair of subapical setae near midline and 3-4 setae near lateral margin; 2) segment II of labial palpi with a bluntly rounded distomedial projection as in Fig. 36B; 3) coxal gills short; 4) tarsal claws with 25-28 denticles (Fig. 36C); 5) posterior margin of abdominal terga as in Fig. 36D; 6) abdominal gills with trachea scarcely darkened; 7) paraprocts as in Fig. 36E; 8) terminal filament subequal in length to cerci. Distribution: Colombia. Camelobaetidius matilei THOMAS, PERU & HOREAU Camelobaetidius matilei THOMAS et al., 2001b: 124.

Taxonomy: This species, known from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Labrum (Fig. 36F), dorsally with 2 subapical setae near lateral margin; 2) segment II of labial palpi with small rounded distomedial projection (Fig. 36G); 3) coxal gills absent; 4) tarsal claws with 13-15 denticles (Fig. 36H); 5) posterior margin of abdominal terga as in Fig. 36I; 6) abdominal gills with main branch of trachea with some dark pigment (Fig. 36J); 7) paraproct as in Fig. 36K; 8) terminal filament subequal in length to cerci. Distribution: French Guiana.

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Plate 36. Camelobaetidius, nymphs. Figs. 36A-E, C. mathuriae: 36A, labrum d.v.; 36B, labium v.v.; 36C, tarsal claw I; 36D, posterior margin of abdominal tergum IV; 36E, paraproct. Figs. 36F-K, C. matilei: 36F, labrum d.v.; 36G, labium v.v.; 36H, tarsal claw I; 36I, posterior margin of abdominal tergum IV; 36J, gill IV; 36K, paraproct. Figs. 36L-P, C. ortizi: 36L, labrum d.v.; 36M, labium v.v.; 36N, tarsal claw I; 36O, posterior margin of abdominal tergum V; 36P, paraproct. (Roman numbers = segment numbers). Figs. 36A-E modified from DOMINIQUE et al., 2001; 36F-K from THOMAS et al., 2001; 36LP from DOMINIQUE & THOMAS, 2001.

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Camelobaetidius ortizi DOMINIQUE & THOMAS Camelobaetidius ortizi DOMINIQUE & THOMAS, 2001: 28.

Taxonomy: This species, known from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Labrum (Fig. 36L), dorsally with 2 subapical setae near midline and 2-3 setae near lateral margin; 2) segment II of labial palpi with a large broadly rounded distomedial projection (Fig. 36M); 3) coxal gills absent; 4) tarsal claws with 23-24 denticles (Fig. 36N); 5) posterior margin of abdominal terga as in Fig. 36O; 6) abdominal gills with main branch of trachea darkened; 7) paraprocts as in Fig. 36P; 8) terminal filament subequal in length to cerci. Distribution: French Guiana. Camelobaetidius patricki DOMINIQUE & THOMAS Camelobaetidius patricki DOMINIQUE & THOMAS (in DOMINIQUE et al., 2001: 23).

Taxonomy: This species, known from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Labrum (Fig. 37A), dorsally with a pair of subapical setae near midline and 3-4 setae laterally; 2) segment II of labial palpi with rounded distomedial projection (Fig. 37B); 3) coxal gills present and short, subequal in length to coxa (Fig. 37C); 4) tarsal claws with 9 denticles (Fig. 37D); 5) posterior margin of abdominal terga as in Fig. 37E; 6) abdominal gills with main branch of trachea and lateral branches darkened; 7) paraprocts as in Fig. 37F; 8) terminal filament subequal in length to cerci. Distribution: Colombia. Camelobaetidius penai (TRAVER & EDMUNDS) Dactylobaetis penai TRAVER & EDMUNDS, 1968: 654 (male, female, nymph); HUBBARD, 1982a: 260; HUBBARD et al., 1992: 203. Camelobaetidius penai; MCCAFFERTY & WALTZ, 1990: 783; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 27; NIETO, 2003a: 252.

Taxonomy: This species, known from nymphs and imagos, can be distinguished from the other species by the following combination of characters. In the adults: 1) Basal segment of forceps long (1.5 times longer than wide) (Fig. 30F). In the nymphs: 1) Labrum (Fig. 37G), dorsally with a subapical pair of setae near midline and 5-6 setae near lateral margin; 2) segment II of labial palpi with a strong pointed distomedial projection (Figs. 37H-I); 3) coxal gills absent; 4) tarsal claws with 20-30 denticles (Fig. 37J); 5) posterior margin of abdominal terga as in Fig. 37K; 6) abdominal gills whitish or main branch of trachea and lateral branches darkened (Fig. 37L); 7) paraproct as in Fig. 37M; 8) terminal filament subequal in length to cerci. Distribution and Biology: Argentina and Bolivia. This species is widely distributed and has been collected in many different kinds of rivers, both mountain and lowland, with sandy or rocky substrate.

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Plate 37. Camelobaetidius, nymphs. Figs. 37A-F, C. patricki: 37A, labrum d.v.; 37B, labium v.v.; 37C, proximal part of leg I; 37D, tarsal claw I; 37E, posterior margin of abdominal tergum IV; 37F, paraproct. Figs. 37G-M, C. penai: 37G, labrum d.v.; 37H, labium, left d.v., right v.v.; 37I, labial palp detail; 37J, tarsal claw I; 37K, posterior margin of abdominal tergum IV; 37L, gill IV; 37M, paraproct. (CG = coxal gill; Roman numbers = segment numbers). Figs. 37A-F, modified from DOMINIQUE et al., 2001; 37G-M from NIETO, 2003a.

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Camelobaetidius phaedrus (TRAVER & EDMUNDS) Dactylobaetis phaedrus TRAVER & EDMUNDS, 1968: 667. Camelobaetidius phaedrus; MCCAFFERTY & WALTZ, 1990: 783; NIETO, 2003a: 239.

Taxonomy: This species was described from nymphs by TRAVER & EDMUNDS (1968). The adults were later described by NIETO (2003a). This species can be distinguished from the other species of the genus by the following combination of characters. In the adults: 1) Abdominal color pattern as in Fig. 30H; 2) basal segment of forceps short, as long as wide, segment II slightly curved (Fig. 30I); 3) hind wing with posterior margin strongly convex and with three longitudinal veins (Fig. 30G). In the nymphs (Fig. 31A): 1) Labrum (Fig. 38A), with a row of long subapical setae dorsally; 2) segment II of labial palpi medially rounded (Figs. 38D-E); 3) coxal gills long, equal in length to coxa and trochanter combined (Fig. 38F); 4) tarsal claws with 7-11 denticles (Fig. 38G); 5) posterior margin of abdominal terga as in Fig. 38I; 6) abdominal gills with main branch of trachea and few a lateral branches darkened (Fig. 38J); 7) paraprocts as in Fig. 38K; 8) terminal filament subequal in length to cerci; 9) mandibles with incisors fused, strongly projected and apically pointed (Figs. 38B-C); 10) abdominal color pattern as in Fig. 38H. Distribution and Biology: Argentina (Misiones) and Brazil (Ariranha river; Rio Grande do Sul, Arroio Irapua). This species was collected in the Paranense area, in rivers with rocky substrate. Camelobaetidius serapis (TRAVER & EDMUNDS) Dactylobaetis serapis TRAVER & EDMUNDS, 1968: 668. Camelobaetidius serapis; DOMINIQUE, MATHURIAU & THOMAS, 2001: 31.

Taxonomy: This species, known from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Labrum, dorsally with 2 subapical setae near midline and 3-4 setae near lateral margin; 2) segment II of labial palpi with a pointed distomedial projection; 3) coxal gills short; 4) tarsal claws with 30-40 denticles; 5) terminal filament subequal in length to cerci. Distribution: Brazil. Camelobaetidius suapi NIETO Camelobaetidius suapi NIETO, 2002: 57.

Taxonomy: This species, known from nymphs and adults, can be distinguished from the other species of the genus by the following combination of characters. In the adults: 1) Abdominal color pattern as in Fig. 30K; 2) basal segment of forceps short, as long as wide (Fig. 30L); 3) hind wing with posterior margin concave basally (Fig. 30J). In the nymphs: 1) Labrum (Fig. 38L), dorsally with a pair of long subapical setae near midline and 2-3 long setae near lateral margin; 2) segment II of labial palp with a small digitiform projection (Figs. 38M-N); 3) coxal gills absent; 4) tarsal claws with 20-21 denticles, first one 1.4 times longer than others (Fig. 38O); 5) posterior margin of abdominal terga as in Fig. 38P; 6) abdominal gills with main branch of trachea darkened (Fig. 38Q); 7) paraproct as in Fig. 38R; 8) terminal filament subequal in length to cerci.

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Plate 38. Camelobaetidius, nymphs. Figs. 38A-E, C. phaedrus: 38A, labrum d.v.; 38B, left mandible v.v.; 38C, right mandible v.v.; 38D, labium, left d.v., right v.v.; 38E, labial palp detail; 38F, leg I; 38G, tarsal claw I; 38H, abdomen d.v.; 38I, posterior margin of abdominal tergum IV; 38J, gill IV; 38K, paraproct. Figs. 38L-R, C. suapi: 38L, labrum d.v.; 38M, labium, left d.v., right v.v.; 38N, labial palp detail; 38O, tarsal claw I; 38P, posterior margin of abdominal tergum IV; 38Q, gill IV; 38R, paraproct. (CG = coxal gill; IC = incisors; P = prostheca; Roman numbers = segment numbers). Figs. 38A-J modified from NIETO, 2003a; 38L-R from NIETO, 2002.

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Distribution and Biology: Bolivia. This species was collected in the Andean mountain rain forests, between 1000 and 1200 m, in waters with rocky substrate. Camelobaetidius tantillus (NEEDHAM & MURPHY) Baetis tantillus NEEDHAM & MURPHY, 1924: 53. Camelobaetidius tantillus; LUGO-ORTIZ & MCCAFFERTY, 1999a: 259; DOMÍNGUEZ et al., 2002: 462.

Taxonomy: This species was originally described from female imagos by NEEDHAM & MURPHY (1924). After studying the holotype, LUGO-ORTIZ & MCCAFFERTY (1999a) transferred this species to Camelobaetidius. However, the shape of the hind wing, the character on which the authors based the transfer, is not restricted to this genus. For this reason the position of this species in Camelobaetidius remains dubious. Distribution: Colombia and Peru. Camelobaetidius tuberosus LUGO-ORTIZ & MCCAFFERTY Camelobaetidius tuberosus LUGO-ORTIZ & MCCAFFERTY, 1999b: 222; NIETO, 2003a: 242.

Taxonomy: This species, known from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Labrum (Fig. 39A), dorsally with a row of long subapical setae; 2) segment II of labial palpi medially rounded (Figs. 39B-C); 3) coxal gills present and long, equal in length to coxa and trochanter combined (Fig. 39D); 4) tarsal claws with 15-17 denticles (Fig. 39E); 5) posterior margin of abdominal terga as in Fig. 39G; 6) abdominal gills with tracheae darkened (Fig. 39H); 7) paraprocts as in Fig. 39I; 8) terminal filament subequal in length to cerci; 9) pronotum with medial pair of erect tubercles; 10) abdominal color pattern as in Fig. 39F. Distribution and Biology: Argentina (Misiones) and Paraguay (Depto. Cordillera). This species was found in rivers with rocky substrate. Camelobaetidius yacutinga NIETO Camelobaetidius yacutinga NIETO, 2003a: 247.

Taxonomy: This species, known from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Labrum (Fig. 39J), dorsally with a pair of long subapical setae near midline and 2-3 long setae near lateral margin; 2) segment II of labial palpi with a pointed distomedial projection (Figs. 39L-M); 3) coxal gills present and long, subequal to coxa and trochanter combined (Fig. 39N); 4) tarsal claws with 16-18 denticles, the first denticle much longer than the others (Fig. 39O); 5) posterior margin of abdominal terga as in Fig. 39P; 6) abdominal gills with the main branch of trachea darkened (Fig. 39Q); 7) paraprocts as in Fig. 39R; 8) terminal filament subequal in length to cerci; 9) hypopharynx with the lingua rounded apically (Fig. 39K). Distribution and Biology: Argentina: Misiones. This species was found in rivers with rocky substrate.

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A III

C II

B

I

G

D

F H

CG

E

I L

J

SL

L K

III

O

P

II

N

I

M

CG

Q

R

Plate 39. Camelobaetidius, nymphs. Figs. 39A-I, C. tuberosus: 39A, labrum d.v.; 39B, labium, left d.v., right v.v.; 39C, labial palp detail; 39D, leg I; 39E, tarsal claw I; 39F, abdomen d.v.; 39G, posterior margin of abdominal tergum IV; 39H, gill IV; 39I, paraproct. Figs. 39J-R, C. yacutinga: 39J, labrum d.v.; 39K, hypopharynx; 39L, labium, left d.v., right v.v.; 39M, labial palp detail; 39N, leg I; 39O, tarsal claw I; 39P, posterior margin of abdominal tergum IV; 39Q, gill IV; 39R, paraproct. (CG = coxal gill; L = lingua; SL = superlingua; Roman numbers = segment numbers). Figs. 39A-R modified from NIETO, 2003a.

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Genus Chane NIETO Chane NIETO, 2003b: 155. (Type-species: Chane baure NIETO, original designation).

Taxonomy: The genus Chane was established by NIETO (2003b). The description was based on nymphs and adults. This genus is currently known only from Bolivia. Adult Characteristics: 1) Turbinate eyes oval (Fig. 40A); 2) fore wing with paired marginal intercalary veins (Fig. 40B); 3) hind wings absent; 4) genitalia (Fig. 40C): second segment of the forceps with an internal projection. Nymphal Characteristics: 1) General view as in Fig. 41A; 2) labrum (Fig. 41B) with a wide U-shaped indentation on anterior margin, numerous long fine setae dorsally and a row of long spines on ventral surface at anterior margin; 3) mandibles (Figs. 41C-D): setae present between prosthecae and molae, prosthecae robust with apical denticles inserted in a concavity; 4) hypopharynx (Fig. 41E) lingua with pointed medial process; 5) maxillae (Figs. 41F-G) with 4 large spine-like teeth, 2 spine-like setae and a row of long setae on galea-lacinia, galea with a tuft of long fine setae on ventral side and a row of 6-7 long and strong setae on dorsal side, base of palpi articulated at base of galea-lacinia, palpi segment II with two rows of extremely long, fine simple spine-like setae on dorsal side; 6) labium (Fig. 41H): dorsal surface of glossae B

A

IV IV

C

Plate 40. Chane baure, male imago. Figs. 40A-C. 40A, head, thorax and first abdominal segments (wings omitted), l.v.; 40B, fore wing; 40C, male genitalia, v.v. (IV = intercalary veins). Figs. 40A-C modified from NIETO, 2003b.

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with short fine setae, dorsal surface of paraglossae with long fine setae, segments II and III of palpi with an extremely enlarged projection extended anteriorly and dorsal surfaces with two rows of very long, fine, simple setae; 7) tarsal claws (Fig. 41I) with one row of denticles, the subapical denticle much larger than others; 8) gills on abdominal segments II-VII. Distribution: Eastern Bolivia. Biology: The nymphs were collected in a river with sandy substrate. The adults were collected in June. A B

P

S

S

P

MP

E F

C

D H

III

G

II

J

I

K

SD

D

I

Plate 41. Chane baure, male nymph. Figs. 41A-K. 41A, head, thorax and abdominal segments I-III, l. v.; 41B, labrum, left d.v., right v.v.; 41C, left mandible, d.v.; 41D, right mandible d.v.; 41E, hypopharynx v.v.; 41F, maxilla, v.v.; 41G, maxilla, d.v.; 41H, labium, left d.v., right v.v.; 41I, tarsal claw; 41J, posterior margin of abdominal tergum IV; 41K, paraproct. (D = denticles; MP = medial process of lingua; P = prostheca; S = setae; SD = subapical denticles; Roman numbers = segment numbers). Figs. 41A-K modified from NIETO, 2003b.

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Chane baure NIETO Chane baure NIETO, 2003b: 155.

Taxonomy: This species was described from the nymph, and male and female imagos. Adults were associated with the nymphs based on color patterns. Chane baure can be characterized as follows. In the adult: 1) General color patterns: orange-brown, dorsal portion of turbinate eyes brownish, margin dark brown. In the nymphs: 1) Posterior margin of abdominal terga with spines in central part (Fig. 41J); 2) gills whitish; 3) posterior margin of paraprocts with small spines (Fig. 41K). Distribution and Biology: Discussed in the generic description. Genus Cloeodes TRAVER Cloeodes TRAVER, 1938: 32; WALTZ & MCCAFFERTY, 1987a: 177; 1987c: 192; MCCAFFERTY & LUGOORTIZ, 1995: 33. (Type-species: Cloeodes maculipes TRAVER, original designation). Notobaetis MORIHARA & EDMUNDS, 1980: 606; MCCAFFERTY & LUGO-ORTIZ, 1995: 33. (Type-species: Notobaetis penai MORIHARA & EDMUNDS, original designation). Centroptella BRAASCH & SOLDÁN, 1980: 123 [partim].

Taxonomy: Cloeodes was established by TRAVER in 1938. The original description included nymphs and imagos from Puerto Rico. Three species were described in the paper although only one, Cloeodes maculipes, remained in this genus. WALTZ & MCCAFFERTY (1987a, c) redescribed some generic characters, described new species and transferred species placed in other genera. Additionally, they synonymized Centroptella (BRAASCH & SOLDÁN, 1980) with Cloeodes and placed Notobaetis as a subgenus of Cloeodes. Later, MCCAFFERTY & LUGO-ORTIZ (1995) described a new species and removed the subgenera established previously. In South America there are presently 13 species known, with seven known from adults, four from nymphs and two from both adults and nymphs. Adult Characteristics: 1) Fore wings with paired (Fig. 42A) or single marginal intercalary veins (all South American species described have paired marginal intercalary veins); 2) hind wings present or absent, if present with two or three longitudinal veins (Figs. 42C, F); 3) genitalia (Figs. 42D, K): forceps three-segmented, segment II may be partially divided, bases of forceps close together. Nymphal Characteristics (Fig. 43A): 1) Antennae 1.5-3 times of head capsule, scape subequal in length to pedicel (Fig. 44A); 2) labrum (Fig. 44B), dorsally with a pair of subpapical setae near midline, anterior margin with two kinds of bifid setae: basally bifid near midline, apically bifid near lateral margin; 3) mandibles with or without setae between prostheca and mola, all South American species without setae; 4) left mandible (Fig. 44C) with incisors fused apically, prostheca robust with denticles apically; 5) right mandible (Fig. 44D) with incisors strongly cleft, prostheca slender and branched; 6) hypopharynx (Fig. 44E): apicolateral margin of superlingua with a row of short spines; 7) maxillae (Fig. 44F) with palpi two-segmented, segment II with an apical constriction; 8) labium (Fig. 44G): glossae subequal to paraglossae, segment II of palpi without projection; 9) femora (Fig. 44H) with dorsal edge parallel to ventral edge and with a row of spines, tibia with a subproximal arc of long setae; 10) tarsal claws 0.3-

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0.6 times length of tarsi, without denticles (Fig. 44J); 11) hind wing pads present or absent; 12) posterior margin of abdominal terga with spines (Fig. 44K), and abdominal sterna II-VI each with two small rows of long fine setae (Fig. 44L); 13) abdominal gills present on segments IVII, at least two times length of each segment; 14) posterior margin of paraprocts with spines (Fig. 44N); 15) terminal filament subequal in length to cerci. Distribution: Africa, Madagascar, North and South America, South of Asia and Australia. Biology: This genus has been collected in a wide diversity of habitats. The nymphs can be found in rivers that are well-oxygenated as well as those that are poorly-oxygenated. KEYS

TO

SOUTH AMERICAN CLOEODES

Male Imagos9 1 – 2(1) – 3(2) – 4(1) –

Hind wings present ........................................................................................................... 2 Hind wings absent ............................................................................................................. 4 Hind wings with costal projection in the center of anterior margin (Fig. 42C) ..... ........................................................................................................................... Cloeodes penai Hind wings with costal projection in the first third of anterior margin (Fig. 42F) ................................................................................................................................................ 3 Segment II of forceps with a basal constriction (Fig. 42K) ........... Cloeodes hydation Segment II of forceps without a basal constriction (Fig. 42G) ................................. ........................................................................................................................ Cloeodes aymara Segment II of forceps with a basal constriction; apical margin of subgenital plate without spines (Fig. 42E) ......................................................................... Cloeodes anduzei Segment II of forceps without a constriction; apical margin of subgenital plate with a short spine (Fig. 42M) .......................................................... Cloeodes venezuelensis

Nymphs 1 – 2(1) – 3(1) – 4(3) –

9

Hind wing pads absent ...................................................................................................... 2 Hind wing pads present .................................................................................................... 3 Maxillae with palpi subequal or shorter than galea-lacinia (similar to Fig. 44F); labium with segment III of palpi conical (similar to Fig. 44G) ......... Cloeodes redactus Maxillae with palpi longer than galea-lacinia (Fig. 45A); labium with segment III of palpi truncate (Fig. 45B) ......................................................................... Cloeodes auwe Antennae short, 1.5 times of head capsule; tarsal claw at least 0.5 times the length of tarsi ............................................................................................................Cloeodes irvingi Antennae long, at least 2 times of head capsule; tarsal claw less than 0.4 times the length of tarsi ..................................................................................................................... 4 Femora with a distal projection (Figs. 45H-J), with two spatulate spines apically (Fig. 45H) ............................................................................................................................. 5 Femora rounded distally (Fig. 44I), with 2 or 3 pointed spines apically (Fig. 44I) ........................................................................................................................... Cloeodes penai

C. binocularis, C. nocturnus, C. stelzneri and C. turbinops were not included in the key because the original descriptions do not have enough characters to distinguish these species from the others.

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A

C A1

B III

IV

IV

F

II

E D C

G

I

I H S

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L

M

Plate 42. Cloeodes, male imagos. Figs. 42A-D, C. penai: 42A, fore wing; 42B, hind wing; 42C, hind wing, detail; 42D, genitalia. Fig. 42E, C. anduzei, genitalia. Figs. 42F-G, C. aymara: 42F, hind wing; 42G, genitalia. Fig. 42H, C. binocularis, turbinate eyes. Figs. 42I-K, C. hydation: 42I, fore wing; 42J, hind wing; 42K, genitalia. Fig. 42L, C. turbinops, genital forcep. Fig. 42M, C. venezuelensis, genitalia. (C = constriction; IV = intercalary veins; S = spine; Roman numbers = segment numbers). Figs. 42E, M modified from TRAVER, 1943; 42F-G modified from TRAVER, 1971; 42H, L modified from NEEDHAM & MURPHY, 1924; 42I-K modified from MCCAFFERTY & LUGO-ORTIZ, 1995.

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5(4) –

CLAVES

General color pattern of head, thorax, and segments I-V of abdomen dark (Fig. 45G); gills apically rounded (similar to Fig. 44M) ............................... Cloeodes jaragua General color not like before (Fig. 45E); gills apically pointed (Fig. 45E) .............. ...................................................................................................................... Cloeodes hydation PARA LOS

Imagos Machos 1 – 2(1) – 3(2) – 4(1) –

CLOEODES

SUDAMERICANOS

10

Alas posteriores presentes ................................................................................................ 2 Alas posteriores ausentes .................................................................................................. 4 Alas posteriores con proyección costal en la mitad del margen anterior del ala (Fig. 42C) ........................................................................................................ Cloeodes penai Alas posteriores con proyección costal en el primer tercio del margen anterior del ala (Fig. 42F) ........................................................................................................................ 3 Segmento II de los fórceps con una constricción basal (Fig. 42K) .... Cloeodes hydation Segmento II de los fórceps sin una constricción basal (Fig. 42G)........ Cloeodes aymara Segmento II de los fórceps con una constricción basal; margen apical de la placa subgenital sin espinas (Fig. 42E) ............................................................ Cloeodes anduzei Segmento II de los fórceps sin constricción; margen apical de la placa subgenital con una espina corta (Fig. 42M) ..................................................... Cloeodes venezuelensis

Ninfas 1 – 2(1) – 3(1) – 4(3) – 5(4) –

10

Pterotecas posteriores ausentes ....................................................................................... 2 Pterotecas posteriores presentes ..................................................................................... 3 Maxilas con palpo subigual o más corto que la galea-lacinia (similar a la Fig. 44F); labio con el segmento III del palpo cónico (similar a la Fig. 44G) ............... Cloeodes redactus Maxilas con el palpo más largo que la galea-lacinia (Fig. 45A); labio con el segmento III del palpo truncado (Fig. 45B) ............................................. Cloeodes auwe Antenas más cortas, 1,5 veces el ancho de la cabeza; uñas tarsales por lo menos 0,5 la longitud del tarso ..................................................................................... Cloeodes irvingi Antenas más largas, por lo menos 2 veces el ancho de la cabeza; uñas tarsales menos de 0,4 la longitud del tarso ................................................................................. 4 Fémures con una proyección distal (Figs. 45H-J), con 2 espinas espatuladas apicalmente (Fig. 45H) ...................................................................................................... 5 Fémures redondeados distalmente (Fig. 44I), con 2 o 3 espinas agudas (Fig. 44I) ........................................................................................................................... Cloeodes penai Coloración general: cabeza, tórax y segmentos I-V del abdomen oscuros (Fig. 45G); branquias redondeadas apicalmente (similar a la Fig. 44M) ................ Cloeodes jaragua Coloración general no como arriba (Fig. 45E); branquias agudas apicalmente (Fig. 45E) ................................................................................................... Cloeodes hydation

C. binocularis, C. nocturnus, C. stelzneri y C. turbinops no fueron incluidos en la clave porque las descripciones originales carecen de caracteres suficientes como para separar estas especies de las restantes.

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A Plate 43. Cloeodes sp. Fig. 43A, nymphal habitus. Drawing by C. Molineri.

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Cloeodes anduzei (TRAVER) Pseudocloeon anduzei TRAVER, 1943: 88. Cloeodes (Cloeodes) anduzei; WALTZ & MCCAFFERTY, 1987c: 197.

Taxonomy: Cloeodes anduzei, known from adults, was originally placed in Pseudocloeon. WALTZ & MCCAFFERTY (1987c) transferred this species to Cloeodes. This species can be distinguished from the other species of the genus by the following combination of characters: 1) Turbinate eyes large; 2) hind wings absent; 3) genitalia: segment II of forceps with a basal constriction (Fig. 42E). Distriution: Venezuela. Cloeodes auwe SALLES & BATISTA Cloeodes auwe SALLES & BATISTA, in SALLES et al., 2004: 5.

Taxonomy: This species, known from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Antennae long, more than two times of head capsule; 2) maxillae with palpi longer than galea-lacinia (Fig. 45A); 3) hind wing pads absent; 4) femora with two spatulate spines apically (Fig. 45C); 5) tarsal claws 0.5 times length of tarsi; 6) abdominal gills rounded apically (Fig. 45D); 7) labium with segment III of palpi truncate (Fig. 45B). Distribution and Biology: Brazil. This species was collected in backwaters among roots and marginal vegetation. Cloeodes aymara (TRAVER) Baetis aymara TRAVER, 1971: 60. Cloeodes (Cloeodes) aymara; WALTZ & MCCAFFERTY, 1987c: 197.

Taxonomy: This species, known from adults, was originally placed in Baetis. Later, WALTZ & MCCAFFERTY examined the type material and transferred this species to Cloeodes. This species can be distinguished from the other species of the genus by the following combination of characters: 1) Turbinate eyes quite large; 2) hind wings present, costal projection in the first third of anterior margin (Fig. 42F); 3) genitalia (Fig. 42G): segment II of forceps without constriction; 4) males of this species have anterior abdominal segments whitish, the middle segments translucent and the posterior segments pale reddish-brown. Distribution: Uruguay. Cloeodes binocularis (NEEDHAM & MURPHY) Pseudocloeon binocularis NEEDHAM & MURPHY, 1924: 57. Cloeodes binocularis; MCCAFFERTY, 2000b: 378.

Taxonomy: This species, known from adults, was originally placed in Pseudocloeon. In 2000, MCCAFFERTY apparently transferred this species to Cloeodes, but the basis for the transfer is not readily apparent in the paper. This species has: 1) Turbinate eyes cylindrincal and erect (Fig. 42H); 2) hind wings absent. Distribution: Peru.

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Cloeodes hydation MCCAFFERTY & LUGO-ORTIZ Cloeodes hydation MCCAFFERTY& LUGO-ORTIZ, 1995: 30.

Taxonomy: This species, known from nymphs, can be distinguished from the other species of the genus by the following combination of characters. In the male imago: 1) Turbinate eyes oval, relatively short; 2) hind wings small, 0.1 times length of fore wings (Figs. 42I-J), costal projection in the first third of anterior margin; 3) genitalia (Fig. 42K): segment II of forceps with a basal constriction. In the nymphs: 1) Antennae long, more than two times of head capsule; 2) maxillae with palpi slightly longer than galea-lacinia; 3) hind wing pads present; 4) femora with a distal projection, with two spatulate spines apically (Fig. 45F); 5) tarsal claws less than 0.4 times length of tarsi; 6) abdominal gills apically pointed (Fig. 45E). Distribution and Biology: Brazil. This species was collected in small temporary rock pools, in an intermittent lowland tropical river. This species is ecologically distinctive in its tolerance to short periods of habitat desiccation (NOLTE et al., 1996). Cloeodes irvingi WALTZ & MCCAFFERTY Cloeodes (Cloeodes) irvingi WALTZ & MCCAFFERTY, 1987c: 200.

Taxonomy: This species, known from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Antennae short, 1.5 times of head capsule; 2) maxillae with palpi subequal in length to galea-lacinia; 3) hind wing pads present; 4) femora with two spatulate spines apically (similar to Fig. 45F); 5) tarsal claws long, more than 0.5 times length of tarsi. Distribution: Paraguay. Cloeodes jaragua SALLES & LUGO-ORTIZ Cloeodes jaragua SALLES & LUGO-ORTIZ, 2003b: 449.

Taxonomy: This species, known from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Antennae long, 2.7 times of head capsule; 2) maxillae with palpi slightly longer than galea-lacinia; 3) hind wing pads present; 4) femora with a distal projection, with two spatulate spines apically (Figs. 45H-J); 5) tarsal claws 0.3 times length of tarsi; 6) abdominal gills apically rounded; 7) color pattern of head, thorax, and segments I-V of the abdomen dark (Fig. 45G). Distribution and Biology: Brazil. This species was collected in small temporary rock pools, in rivers at 1000 m of altitude. Cloeodes nocturnus (NAVÁS) Baetis nocturnus NAVÁS, 1922b: 199. Cloeodes nocturnus; LUGO-ORTIZ & MCCAFFERTY, 1999a: 259.

Taxonomy: This species, known from adults, was originally placed in the genus Baetis. After examination of the original description, LUGO-ORTIZ & MCCAFFERTY (1999a) concluded that this species belonged to Cloeodes. However, the shape of the hind wings, the

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character used as the bases for transfer of this species, is not restricted to Cloeodes. For this reason the position of this species in Cloeodes is maintained with doubt. This species has the hind wings with a small costal process in the basal third of the wing (similar to Fig. 42F). Distribution: Argentina. P

A

P

C

D

SL

L

B C

G

E

III

S

I

II AS

F I

K

H LS

J

L M

N

Plate 44. Cloeodes penai, nymph. Figs. 44A-N: 44A, antenna; 44B, labrum, left d.v., right v.v.; 44C, left mandible v.v.; 44D, right mandible v.v.; 44E, hypopharynx v.v.; 44F, maxilla v.v.; 44G, labium, left d.v., right v.v.; 44H, leg I; 44I, femur I, apical margin; 44J, tarsal claw I; 44K, posterior margin of abdominal tergum IV; 44L, ventrolateral view of abdominal sternum IV; 44M, gill IV; 44N, paraproct. (AS = arc of setae; C = constriction; L = lingua; LS = long setae; P = prostheca; S = spines; SL = superlingua; Roman numbers = segment numbers).

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Cloeodes penai (MORIHARA & EDMUNDS) Notobaetis penai MORIHARA & EDMUNDS, 1980: 606. Cloeodes (Notobaetis) penai; WALTZ & MCCAFFERTY, 1987c: 196.

Taxonomy: This species, known from nymphs and adults, can be distinguished from the other species of the genus by the following combination of characters. In the male imago: 1) Turbinate eyes oval, two times longer than wide; 2) hind wings small, 0.1 times length of fore wings (Figs. 42A-B), costal projection in the center of anterior margin (Fig. 42C); 3) genitalia (Fig. 42D): segment II of forceps without constriction. In the nymphs: 1) Antennae long, more than two times of head capsule; 2) maxillae with palpi subequal in length to galea-lacinia (Fig. 44F); 3) hind wing pads present; 4) femora rounded distally (Fig. 44H) and with 2-3 pointed spines apically (Fig. 44I); 5) tarsal claws less than 0.4 times length of tarsi; 6) abdominal gills apically rounded (Fig. 44M). Distribution and Biology: Argentina. Cloeodes redactus WALTZ & MCCAFFERTY Genus poss. Cloeodes; ROBACK, 1966: 133. Cloeodes (Cloeodes) redactus WALTZ & MCCAFFERTY, 1987c: 204. Cloeodes redactus, DOMÍNGUEZ et al., 2002: 462.

Taxonomy: This species, known from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Antennae longer than head capsule; 2) maxillae with palpi shorter or subequal in length to galea-lacinia; 3) hind wing pads absent; 4) femora with 2-3 spatulate spines apically (similar to Fig. 45F); 5) tarsal claws 0.5 times length of tarsi. Distribution: Colombia and Peru. Cloeodes stelzneri (WEYENBERGH) Cloe stelzneri WEYENBERGH, 1883: 170. Baetis stelzneri; EATON, 1885: 171. Cloeodes stelzneri; LUGO-ORTIZ & MCCAFFERTY, 1999a: 259.

Taxonomy: This species, known from adults, was originally placed in the genus Cloe. The type material is apparently lost (LUGO-ORTIZ & MCCAFFERTY, 1999a). After examination of the original description, LUGO-ORTIZ & MCCAFFERTY transferred this species to Cloeodes and suggested a possible synonymy between this species and Cloeodes penai. This species has a costal projection in center of anterior margin of the hind wings. Distribution: Argentina. Cloeodes turbinops (NEEDHAM & MURPHY) Pseudocloeon turbinops NEEDHAM & MURPHY, 1924: 57. Cloeodes turbinops; LUGO-ORTIZ & MCCAFFERTY, 1999a: 261.

Taxonomy: This species, known from adults, was originally placed in Pseudocloeon. After examination of the holotype, LUGO-ORTIZ & MCCAFFERTY (1999a) transferred this

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III

C II

A I

B

AS

D

F

H P

E

I

P

G

J Plate 45. Cloeodes, nymphs. Figs. 45A-D, C. auwe: 45A, maxilla; 45B, labium, left d.v., right v.v.; 45C, femur I, apical margin; 45D, gill IV. Figs. 45E-F, C. hydation: 45E, nymph d.v.; 45F, femur I, apical margin. Figs. 45G-J, C. jaragua: 45G, nymph d.v.; 45H, femur I, apical margin; 45I, femur II, apical margin; 45J, femur III, apical margin. (AS = arc of setae; P = projection; Roman numbers = segment numbers). Figs. 45A-D modified from Salles et al., 2004; 45E-F modified from MCCAFFERTY & LUGO-ORTIZ, 1995; 45G-J modified from SALLES & LUGO-ORTIZ, 2000b.

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species to Cloeodes. However, these authors concluded that it will be difficult to associate future specimens with this species because of uncertainties regarding its specific characteristics. This species has: 1) Fore wings relatively narrow-elongate and somewhat pointed apically; 2) hind wings absent; 3) genitalia (Fig. 42L): forceps relatively erect with a broad segment II and relatively elongated and ovoid segment III. Distribution: Guyana. Cloeodes venezuelensis (TRAVER) Pseudocloeon venezuelensis TRAVER, 1943: 92. Cloeodes venezuelensis; LUGO-ORTIZ & MCCAFFERTY, 1999a: 261.

Taxonomy: This species, known from adults, was originally placed in Pseudocloeon. After examination of the original description, LUGO-ORTIZ & MCCAFFERTY (1999a) transferred this species to Cloeodes. This species has: 1) Male turbinate eyes erect, cylindrical and with a slightly larger diameter apically than basally; 2) hind wings absent; 3) genitalia (Fig. 42M): with a short spine on apical margin of the subgenital plate. Distribution: Venezuela. Genus Cryptonympha LUGO-ORTIZ & MCCAFFERTY ”Genus 2 nr. Pseudocloeon KLAPALEK” ROBACK, 1966: 135. Cryptonympha LUGO-ORTIZ & MCCAFFERTY, 1998: 58; ORTH et al., 2000: 35; SALLES & FRANCISCHETTI, 2004: 213. (Type-species: Cryptonympha copiosa LUGO-ORTIZ & MCCAFFERTY, original designation).

Taxonomy: This genus was established by LUGO-ORTIZ & MCCAFFERTY in 1998. They described one species, Cryptonympha copiosa, from nymphs. Later, SALLES & FRANCISCHETTI (2004) described another species, C. dasilvai, known only from nymphs as well. Adult Characteristics: Unknown. Nymphal Characteristics: 1) Antennae long, 2.5 times of head capsule; 2) labrum (Fig. 46A) with bipectinate setae along anterior margin (Fig. 46B); 3) left mandible (Fig. 46C) with incisors fused, prostheca robust with denticles apically; 4) right mandible with incisors fused (Fig. 46D), setae between prostheca and mola, prostheca robust with denticles apically and with a bipectinate transverse seta (Fig. 46E); 5) hypopharynx (Fig. 46F): lingua subequal in length to superlingua and with a rounded projection apically; 6) maxillae (Fig. 46G) with palpi two-segmented, 1.5 times length of galea-lacinia; 7) labium (Fig. 46H) with glossae subequal to paraglossae, acute apically and with a row of spine-like setae, segment II of palpi with a distomedial projection, segment III conical; 8) tarsal claws with one or two rows of denticles (Fig. 46J); 9) abdominal gills on segments I-VII; 10) posterior margin of abdominal terga with spines (Fig. 46K); 11) posterior margin of paraprocts with spines (Fig. 46M); 12) terminal filament subequal in length to cerci. Distribution: Brazil, Colombia and French Guiana. Biology: Unknown.

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Labrum (Fig. 46A) dorsally with 1 pair of long subapical setae; left mandible (Fig. 46C) with a tuft of setae between prostheca and mola; segment II of labial palpi with a thumblike distomedial projection (Fig. 46H); hind wing pads absent ....... Cryptonympha copiosa Labrum (Fig. 46N) dorsally with 2 pairs of long subapical setae; left mandible (Fig. 46O) without a tuft of setae between prostheca and mola; segment II of labial palpi with a weak distomedial projection (Fig. 46P); hind wing pads rudimentary ............ .............................................................................................................. Cryptonympha dasilvai PARA LOS

CRYPTONYMPHA

SUDAMERICANOS

Ninfa 1

–

Labro (Fig. 46A), dorsalmente con 1 par de setas largas subapicales; mandíbula izquierda (Fig. 46C) con un mechón de setas entre la prosteca y la mola; segmento II del palpo labial con una proyección distomedial digitiforme (Fig. 46H); pterotecas posteriores ausentes ...................................................... Cryptonympha copiosa Labro (Fig. 46N), dorsalmente con 2 pares de setas largas subapicales; mandíbula izquierda (Fig. 46O) sin setas entre la prosteca y la mola; segmento II del palpo labial con una débil proyección distomedial (Fig. 46P); pterotecas posteriores rudimentarias .................................................................................... Cryptonympha dasilvai

Cryptonympha copiosa LUGO-ORTIZ & MCCAFFERTY Cryptonympha copiosa LUGO-ORTIZ & MCCAFFERTY, 1998: 60; DOMÍNGUEZ et al., 2002: 462.

Taxonomy: This species, known from nymphs, can be distinguished by the following combination of characters: 1) Labrum (Fig. 46A) with a pair of long subapical setae on dorsal surface; 2) left mandible (as in Fig. 46C) with a tuft of setae between prostheca and mola and with a bipectinate transverse seta; 3) segment II of labial palpi with a thumb-like distomedial projection (as in Fig. 46H); 4) hind wing pads absent; 5) tarsal claws with one row of denticles; 6) gill I strongly narrowed basally (as in Fig. 46L). Distribution: Brazil and Colombia. Cryptonympha dasilvai SALLES & FRANCISCHETTI Cryptonympha dasilvai SALLES & FRANCISCHETTI, 2004: 213.

Taxonomy: This species, known from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Labrum (Fig. 46N) dorsally with 2 pairs of long subapical setae; 2) left mandible (Fig. 46O) without a tuft of setae between prostheca and mola; 3) segment II of labial palpi with a weak distomedial projection (Fig. 46P); 4) hind wing pads rudimentary; 5) tarsal claws with two rows of denticles (as in Fig. 46J); 6) gill I not narrowed basally (Fig. 46L). Distribution: Brazil.

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P Plate 46. Cryptonympha, nymphs. Figs. 46A-M, Cryptonympha sp. Figs. 46A-H, mouthparts: 46A, labrum, left d.v., right v.v.; 46B: labrum, setae of anterior margin; 46C, left mandible v.v.; 46D, right mandible v.v.; 46E, prostheca of right mandible; 46F, hypopharynx v.v.; 46G, maxilla v.v.; 46H, labium, left d.v., right v.v. 46I, leg I; 46J, tarsal claw I v.v.; 46K, posterior margin of abdominal tergum IV; 46L, gill I; 46M, paraproct. Figs. 46N-P, C. dasilvai: 46N, labrum d.v.; 46O, left mandible v.v.; 46P, labium, left d.v., right v.v. (D = denticles; L = lingua; P = prostheca; S = setae; SL = superlingua; TS = transverse seta; Roman numbers = segment numbers). Figs. 46N-P modified from SALLES & FRANCISCHETTI, 2004.

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Genus Fallceon WALTZ & MCCAFFERTY Fallceon WALTZ & MCCAFFERTY, 1987c: 668; LUGO-ORTIZ & MCCAFFERTY, 1999a: 259. (Type-species: Baetis quilleri DODDS, original designation).

Taxonomy: This genus was established by WALTZ & MCCAFFERTY (1987c). They originally included species placed in the genus Baetis. The species initially included in this genus had distributions restricted to North and Central America. Later, LUGO-ORTIZ & MCCAFFERTY (1999a) transferred three species, previously placed in Baetis, to this genus: Fallceon inops (NAVÁS), Fallceon murphyae (HUBBARD) and Fallceon yaro (TRAVER). The three species, known from adults, are distributed in Argentina, Paraguay, Peru and Uruguay. Later, MCCAFFERTY (2000b) proposed Fallceon inops as nomen dubium. The shape of the hind wing was the character used by WALTZ & MCCAFFERTY as the basis for the transfer of these species. However, a hind wing with two longitudinal veins and a hooked costal process is not restricted to Fallceon. For this reason and until the nymphs of these species are reared we include this genus in the keys to South American genera but a discussion of its species is not included. Genus Guajirolus FLOWERS Genus 3 nr. Pseudocloeon ROBACK, 1966: 135. Guajirolus FLOWERS, 1985: 27; LUGO-ORTIZ & MCCAFFERTY, 1995c: 68; NIETO, 2003b: 153. (Type-species: Guajirolus ektrapeloglossa FLOWERS, original designation).

Taxonomy: Guajirolus was established in 1985 by FLOWERS, who described G. ektrapeloglossa from nymphs and adults and placed ROBACK’s (1966) “Genus 3 nr. Pseudocloeon KLAPALEK” into Guajirolus. LUGO-ORTIZ & MCCAFFERTY (1995c) described G. nanus from nymph and NIETO (2003b) described G. queremba from the nymph and female subimago.

IP

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B Plate 47. Guajirolus, imagos. Fig. 47A, G. queremba , female fore wing. 47B, G. ektrapeloglossa, male genitalia v.v. (IP = internal projection; IV = intercalary veins; P = projection). Fig. 47A modified from NIETO, 2003b; 47B from FLOWERS, 1985.

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Adult Characteristics: 1) Fore wings with paired marginal intercalary veins (Fig. 47A); 2) hind wings absent; 3) strong medial projection on second segment of genital forceps and a small projection between forceps (Fig. 47B). Nymphal Characteristics: 1) Anterior margin of labrum with a U-shaped emargination, ventral surface with a row of spines along anterior margin (Fig. 48A); 2) mandibles (Figs. 48B-C) with incisors fused apically, simple setae between prosthecae and molae, prosthecae robust with denticles apically and inserted in a concavity; 3) hypopharynx (Fig. 48D): lingua subequal in length to superlingua and with three lobes apically; 4) maxillae (Fig. 48E) with a tuft of long setae on galea apically, palpi three-segmented, two times length of galea-lacinia; 5) labium (Fig. 48F): glossae subequal to paraglossae; dorsal surface of glossae with large spine-like setae; ventral surface of paraglossae with three long simple setae, palpi three-segmented, segment II greatly enlarged anteromedially; 6) tarsal claws with subapical denticle larger than others (Figs. 48I-J); 7) abdominal gills present on segments II-VII (Fig. 48K); 8) posterior margin of abdominal terga with spines (Fig. 48L). Distribution: Argentina, Bolivia, Colombia, French Guiana, Costa Rica and Panama. Biology: The nymphs have been found on rocky substrates. KEY

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CLAVE

Paraprocts with unorganized spination (Fig. 48H); lingua with medial lobe pointed (Fig. 48G) ...................................................................................... Guajirolus ektrapeloglossa Paraprocts with organized spination (Fig. 48M); lingua with medial lobe rounded (Figs. 48D) ............................................................................................ Guajirolus queremba PARA LOS

GUAJIROLUS

SUDAMERICANOS

Ninfa 1 –

Paraproctos con una hilera de puntas desordenadas (Fig. 48H); lingua con el lóbulo medio agudo (Fig. 48G) ................................................ Guajirolus ektrapeloglossa Paraproctos con una hilera de puntas ordenadas (Fig. 48M); lingua con el lóbulo medio redondeado (Figs. 48D) ........................................................ Guajirolus queremba

Guajirolus ektrapeloglossa FLOWERS Guajirolus ektrapeloglossa FLOWERS, 1985: 29.

Taxonomy: FLOWERS (1985) described this species based on the nymph and male and female imagos. Adults were associated with nymphs based on color patterns. Guajirolus ektrapeloglossa can be distinguished from the other species of the genus by the following combination of characters: 1) Lingua: medial lobe pointed (Fig. 48G); 2) paraprocts with unorganized spination (Fig. 48H). Distribution and Biology: Panama and Colombia. The nymphs have been collected from both medium sized rivers and small streams. Water temperatures at collection localities ranged from 22-24°C. Gut contents consisted of diatoms, algae cells, and plant tissue.

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Plate 48. Guajirolus, nymphs. Figs. 48A-F, G. queremba. Figs. 48A-F, mouthparts: 48A, labrum, left d.v., right v.v.; 48B, left mandible v.v.; 48C, right mandible v.v.; 48D, hypopharynx v.v.; 48E, maxilla v.v.; 48F, labium, left d.v., right v.v. Figs. 48G-H, G. ektrapeloglossa: 48G, hypopharynx v.v.; 48H, paraproct. Figs. 48I-M, G. queremba: 48I, leg I; 48J, tarsal claw I; 48K, abdominal segments d.v.; 48L, posterior margin of tergum IV; 48M, paraproct. (D = denticles; L = lingua; P = prostheca; S = setae; SD = subapical denticle; SL = superlingua; Roman numbers = segment numbers). Figs. 48A-F, I-M modified from NIETO, 2003b; 48G-H from FLOWERS, 1987.

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Guajirolus queremba NIETO Guajirolus queremba NIETO, 2003b: 65.

Taxonomy: This species was described based on the nymph and female subimago. Guajirolus queremba can be distinguished from the other species of the genus by the following combination of characters: 1) Lingua with medial lobe rounded (Fig. 48D); 2) paraprocts with organized spination (Fig. 48M). Distribution: Argentina. Genus Harpagobaetis MOL Harpagobaetis MOL, 1986: 63; ORTH et al., 2000: 35; SALLES & LUGO-ORTIZ, 2002: 155. (Type-species: Harpagobaetis gulosus MOL, original designation).

Taxonomy: Harpagobaetis was established by MOL in 1986. This is the only carnivorous baetid genus in South America. Harpagobaetis gulosus, known from nymphs, is the only species described in this genus. Adult Characteristics: Unknown. Nymphal Characteristcs: 1) Antennae short, 1.5 times of head capsule; 2) labrum (Fig. 49A) laterally expanded, width 3 times length, anteriorly with W-shaped and with short spines on margin at middle (Fig. 49B), laterally with palmately branched setae (Fig. 49C); 3) mandibles (Figs. 49D-G) with 4 cleft incisors, molars pointed and strongly protruding; 3) prostheca of left mandible with 2-3 denticles apically (Fig. 49E), right prostheca reduced to a simple seta (Fig. 49G); 4) hypopharynx: lingua large and broadly rounded apically, superlinguae laterally projected (Figs. 49H-I); 5) maxillae (Figs. 49J-K): canines elongate, palpi three segmented, shorter than galea-lacinia; 6) labium (Figs. 49LM) with glossae subequal to paraglossae, both slender elongate with a row of long spinelike setae on external margin, segment II of palpi without projection, segment III conical; 7) tarsal claws with 2 rows of denticles and with a pair subapical setae (Fig. 49P); 8) hind wing pads present; 9) terminal filament subequal in length to cerci. Distribution: Brazil, French Guiana, Surinam. Biology: Harpagobaetis feeds on larvae of Diptera. Specimens of this genus were collected in the lower part of Double Steps Falls. Harpagobaetis gulosus MOL Harpagobaetis gulosus MOL, 1986: 65; SALLES & LUGO-ORTIZ, 2002: 155.

Taxonomy: This species, known only from nymphs, can be distinguished by the following combination of characters: 1) Maxillae with 4 long canines, second canine half length of others (Fig. 49K); 2) ventral margin of femora with rounded spines (Fig. 49N) and tibia with serrated spines (Fig. 49O); 3) abdomen with dark spots on anterior margin of terga II-VI. Distribution and Biology: Brazil and Surinam. Biology discussed in the generic description.

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Plate 49. Harpagobaetis gulosus, nymph. Figs. 49A-P. Figs. 49A-M, mouthparts: 49A, labrum, d.v.; 49B, labrum, apical setae detail; 49C, labrum, lateral setae detail; 49D, left mandible v.v.; 49E, distal portion of left mandible; 49F, right mandible v.v.; 49G, distal portion of right mandible; 49H, hypopharynx v.v.; 49I, hypopharynx posterior v.; 49J, maxilla v.v.; 49K, distal portion of maxilla; 49L, labium, right half, v.v.; 49M, detail of glossa and paraglossa v.v.; 49N, leg I; 49O, apical portion of tibia; 49P, tarsal claw I. (D = denticles; I = incisors; L = lingua; P = prostheca; PS = serrate spines; S = rounded spines; SL = superlingua; SS = subapical seta; Roman numbers = segment numbers). Figs. 49A-P modified from MOL, 1986.

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Genus Iguaira SALLES & LUGO-ORTIZ Iguaira SALLES & LUGO-ORTIZ, 2003a: 202. (Type-species: Iguaira poranga SALLES & LUGO-ORTIZ, original designation).

Taxonomy: This genus was established by SALLES & LUGO-ORTIZ in 2003. They included only one species, Iguaira poranga, known from nymphs. Iguaira is in revision at this moment, and there are some doubts about its validity. Adult Characteristics: Unknown. Nymphal Characteristics: 1) Labrum (Fig. 50A) wider than long, anterior margin with bifid setae (Fig. 50B); 2) left mandible (Fig. 50C) with incisors cleft at mid-length, prostheca with 2-3 denticles apically; 3) right mandible (Fig. 50D) with incisors cleft to base, prostheca slender and bifid; 4) hypopharynx (Fig. 50E): lingua slightly longer than superlingua; 5) maxillae (Fig. 50F) palpi two-segmented, longer than galea-lacinia; 6) labium (Fig. 50G): glossae slightly shorter and wider than paraglossae and with short spines marginally, paraglossae with a row of long spine-like setae, segment II of palpi with a strong distomedial projection, segment III conical, both segments with long setae; 7) leg (Fig. 50H): dorsal edge of femora with short spines; 8) tarsal claws (Fig. 50I) 0.5 times length of tarsi and with minute denticles; 9) hind wing pads present; 10) posterior margin of abdominal terga with spines (Fig. 50J); 11) gills (Fig. 50K) present on segments I-VII, elongate, two times length of each segment; 12) posterior margin of paraprocts with spines (Fig. 50L); 13) terminal filament subequal in length to cerci. Distribution: Brazil (Minas Gerais). Iguaira poranga SALLES & LUGO-ORTIZ Iguaira poranga SALLES & LUGO-ORTIZ, 2003a: 203.

Taxonomy: This species, known from nymphs, can be distinguished by the following combination of characters: 1) Labrum, dorsally with scattered short setae (Fig. 50A); 2) hypopharynx: lingua with a pointed projection apically (Fig. 50E); 3) maxillae with palp at least two times length of galea-lacinia (Fig. 50F). Distribution: Discussed in the generic description. Genus Mayobaetis WALTZ & MCCAFFERTY Moribaetis (Mayobaetis) WALTZ & MCCAFFERTY, 1985: 240. Mayobaetis; LUGO-ORTIZ & MCCAFFERTY, 1996b: 369. (Type-species: Baetis ellenae MAYO, original designation).

Taxonomy: This genus was originally placed as subgenus of Moribaetis by WALTZ & MCCAFFERTY (1985). Only one species was included, Mayobaetis ellenae, known from nymphs and adults, described as Baetis ellenae by MAYO (1973a). LUGO-ORTIZ & MCCAFFERTY (1996b) assigned generic rank to Mayobaetis and placed it in the Baetodes complex. Adult Characteristics: 1) Compound eyes tubular, height of stalk subequal to diameter; 2) fore wings with paired marginal intercalary veins, vein A1 strongly curved (Fig. 51A); 3) hind wings with two longitudinal veins, costal process pointed and in basal third of wing margin (Figs. 51B-C); 4) genitalia: forceps three-segmented (Fig. 51D).

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Nymphal Characteristcs: 1) Antennae short, 1.5 times of head capsule, scape 3 times length of pedicel, scape and pedicel dorsoventrally flattened, with scale-like tubercle and with a row of fine setae (Fig. 52A); 2) labrum with two different kinds of bipectinated setae along anterior margin: basally bifid near the midline and apically bifid in the lateral margin, ventrally with lateral margin sclerotized (Fig. 52B); 3) mandibles with incisors I

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Plate 50. Iguaira poranga, nymph. Figs. 50A-L. Figs. 50A-G, mouthparts: 50A, labrum, left d.v., right v.v.; 50B, labrum, apical setae; 50C, left mandible v.v.; 50D, right mandible v.v.; 50E, hypopharynx v.v.; 50F, maxilla v.v.; 50G, labium, left d.v., right v.v.; 50H, leg I; 50I, tarsal claw I; 50J, posterior margin of abdominal tergum IV; 50K, gill IV; 50L, paraproct. (D = denticles; I = incisors; L = lingua; P = prostheca; SL = superlingua; Roman numbers = segment numbers).

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B C Plate 51. Mayobaetis, imagos. Figs. 51A-C, Mayobaetis sp.: 51A, fore wing; 51B, hind wing; 51C, hind wing, detail. Fig. 51D, M. ellenae, male genitalia. (IV = intercalary veins; P = projection). Fig. 51D modified from LUGO-ORTIZ & MCCAFFERTY, 1996b.

fused apically (Figs. 52C-D); 4) prostheca of left mandible robust with denticles apically (Fig. 52C), right prostheca reduced and apically bifid (Fig. 52D); 5) hypopharynx: lingua longer than superlinguae and with strong apical projection (Fig. 52E); 6) maxillae: palpi two-segmented and subequal in length to galea-lacinia, segment II with an apical constriction (Fig. 52F); 7) labium: glossae subequal in length and half width of paraglossae, glossae with 2-3 rows and paraglossae with 3 rows of pectinate setae, segment II of palpi without projection, segment III ovoid (Fig. 52G); 8) femora with a row of long setae and stout spines alternated (Fig. 52H); 9) tarsal claws with one row of denticles and with a subapical strong seta on external surface (Fig. 52I); 10) posterior margin of abdominal terga smooth; 11) posterior margin of paraprocts smooth; 12) terminal filament 0.3 times length of cerci, cerci with short spines on external margin (Fig. 52J). Distribution: Argentina, Bolivia, Colombia, Costa Rica, Ecuador, Peru. Biology: Mayobaetis inhabits small waterfalls (Fig. 233A). The nymphs are found on the surfaces of rocky slopes in the area (Fig. 233B), or in the splash zones. Mayobaetis ellenae MAYO Baetis sp. 1 ROBACK, 1966: 137. Baetis ellenae MAYO, 1973a: 285; BERNER, 1980: 190. Moribaetis (Mayobaetis) ellenae; WALTZ & MCCAFFERTY, 1985: 240. Mayobaetis ellenae; LUGO-ORTIZ & MCCAFFERTY, 1996: 369.

Taxonomy: This species, known from nymphs and adults, can be distinguished by the following combination of characters. In the male imago: 1) Genitalia: basal segment of forceps with a small distomedial projection (Fig. 51D). In the nymphs: 1) Color pattern of abdomen with dark band at the midline of terga II-VII or VII (Fig. 52K). Distribution and Biology: Costa Rica, Ecuador and Peru. Biology discussed in the generic characteristics.

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Plate 52. Mayobaetis, nymphs. Figs. 52A-J, Mayobaetis sp.: 52A, antenna: scape pedicel and first segments of flagellum. Figs. 52B-G, mouthparts: 52B, labrum, left d.v., right v.v.; 52C, left mandible v.v.; 52D, right mandible v.v; 52E, hypopharynx v.v.; 52F, maxilla v.v.; 52G, labium, left d.v., right v.v. 52H, leg I; 52I, tarsal claw I; 52J, cercus. Fig. 52K, M. ellenae, abdomen d.v. (C = constriction; D = denticles; L = lingua; MS = marginal setae; P = prostheca; S = scale; SL = superlingua; SP = spines; SS = subapical setae; Roman numbers = segment numbers).

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Genus Moribaetis WALTZ & MCCAFFERTY Moribaetis (Moribaetis) WALTZ & MCCAFFERTY, 1985: 243. Moribaetis; LUGO-ORTIZ & MCCAFFERTY, 1996b: 368; LUGO-ORTIZ & MCCAFFERTY, 1999a: 260. (Type-species: Baetis maculipennis FLOWERS, original designation).

Taxonomy: This genus was established by WALTZ & MCCAFFERTY (1985). They originally included species placed in the genus Baetis. Later, several species were included in this genus, all distributed in Central America. LUGO-ORTIZ & MCCAFFERTY (1999a) transferred three species, previously placed in Baetis, to this genus: Moribaetis aneto (TRAVER); Moribaetis comes (NAVÁS) and Moribaetis socius (NEEDHAM & MURPHY). These species, known from adults, are distributed in Brazil, Peru and Uruguay. Later, MCCAFFERTY (2000b) proposed Moribaetis socius as nomen dubium. The shape of the hind wing was the character used by WALTZ & MCCAFFERTY as the basis for the transfer of these species. However, a hind wing with three longitudinal veins (the second vein bifurcate) and a pointed costal projection is not restricted to Moribaetis. For that reason and until the nymphs of these species are reared we include this genus in the key to South American genera but a discussion of its species is not included. Genus Nanomis LUGO-ORTIZ & MCCAFFERTY Nanomis LUGO-ORTIZ & MCCAFFERTY, 1999c: 100. (Type-species: Nanomis galera LUGO-ORTIZ & MCCAFFERTY, original designation).

Taxonomy: This genus was established by LUGO-ORTIZ & MCCAFFERTY (1999c). The original description included only one species Nanomis galera, known from nymphs. Adult Characteristics: Unknown. Nymphal Characteristics: 1) Antennae long, 2.5 times of head capsule, scape and pedicel flattened dorsoventrally; 2) labrum (Fig. 53A) with two kinds of bipectinate setae on anterior margin: basally bifid near the midline and apically branched on lateral margin (Fig. 53B); 3) left mandible (Figs. 53C) with incisors fused and prostheca robust with denticles apically (Fig. 53C); 4) right mandible with incisors cleft and prostheca slender and bifid (Fig. 53D); 5) hypopharynx: lingua subequal in length to superlingua and with small rounded projection apically; 6) maxillae (Fig. 53E) with palpi two-segmented and subequal in length to galea-lacinia; 7) labium (Fig. 53F): glossae subequal to paraglossae, segment II with small distomedial projection, segment III oval; 8) legs (Fig. 53G): dorsal edge of femora with a row of long apically rounded spines (Fig. 53H), tibiae with pectinate spines (Fig. 53I), tibia III with a subapical paddle-shaped seta; 9) tarsal claws with one row of denticles (Fig. 53J); 10) hind wing pads present; 11) posterior margin of paraprocts with spines (Fig. 53L); 12) terminal filament 0.80 times the length of cerci. Distribution: Argentina, Bolivia, Colombia, Ecuador and Peru. Biology: Specimens of this genus have been collected in many rivers, especially from mountain rain forests.

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Plate 53. Nanomis galera, nymph. Figs. 53A-L. Figs. 53A-F, mouthparts: 53A, labrum, left d.v., right v.v.; 53B, labrum, apical margin; 53C, left mandible v.v.; 53D, right mandible; 53E, maxilla v.v.; 53F, labium, left d.v., right v.v. 53G, leg I; 53H, dorsal edge of femur I; 53I, apical margin of tibia I; 53J, tarsal claw I; 53K, gill IV; 53L, paraproct. (D = denticles; P = prostheca; S = spine; Roman numbers = segment numbers).

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Nanomis galera LUGO-ORTIZ & MCCAFFERTY Nanomis galera LUGO-ORTIZ & MCCAFFERTY, 1999c: 100.

Taxonomy: This species, known from nymphs, can be distinguished by the following combination of characters: 1) Labrum (Fig. 53A) dorsally with 1 pair of subapical setae near the midline and 5-6 subapical setae laterally; 2) labium (Fig. 53F) with segment II of palp with slight distomedial projection; 3) posterior margin of abdominal terga I-V smooth and terga VI-X with spines; 4) gills on segments I-VII, asymmetrical, tracheae pigmented (Fig. 53K). Distribution and Biology: Discussed in the generic description. Genus Paracloeodes DAY Paracloeodes DAY, 1955: 121; LUGO-ORTIZ & MCCAFFERTY, 1996c: 2. (Type-species: Paracloeodes abditus DAY, original designation); (=Pseudocloeon minutus DAGGY).

Taxonomy: Paracloeodes was established by DAY in 1955. Originally this genus included one species, Paracloeodes abditus, known from nymphs and adults from the USA. However, MCCAFFERTY & WALTZ (1990) synonymized this species with P. minutus (DAGGY, 1945). Seven species are currently described in Paracloeodes, four of them known from South America. Adult Characteristics: 1) Compound eyes oval, height of stalk half of eye diameter (Figs. 54A-B); 2) fore wings with paired marginal intercalary veins (Fig. 54D); 3) hind wings with two longitudinal veins, costal process quadrangulate (Figs. 54E-F); 4) genitalia: forceps three-segmented, segment III elongate (Fig. 54C). III II

C A I

B

E CP

F

D IV

IV

Plate 54. Paracloeodes, imagos. Figs. 54A-C, P. minutus, male: 54A, head l.v.; 54B, head d.v.; 54C, genitalia v.v. Figs. 54D-F. P. eurybranchus, female: 54D, fore wing; 54E, hind wing; 54F, hind wing detail. (CP = costal process; IV = intercalary veins; Roman numbers = segment numbers). Figs. 54A-C modified from DAY, 1955; 54D-E from LUGO-ORTIZ & MCCAFFERTY, 1996c.

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Nymphal Characteristics: 1) Antennae variable, 2-4 times of head capsule; 2) labrum (Fig. 55A) wider than long, anterior margin with bifid setae (Fig. 55B); 3) left mandible (Figs. 55C, 56B) with incisors cleft at mid length, prostheca with 2-3 denticles apically; 4) right mandible (Figs. 55D, 56C) with incisors cleft basally, prostheca slender and bifid; 5) hypopharynx (Fig. 55E): lingua subequal in length to superlingua; 6) maxillae (Figs. 55F, 56E): palpi two-segmented, longer than galea-lacinia; 7) labium (Figs. 55G, 56F): glossae slightly shorter and equal in width to paraglossae and with short spines marginally, paraglossae with a row of long spine-like setae, segment II of palp with a strong distomedial projection, segment III conical, both segments with long setae; 8) legs (Figs. 55H, 56G) robust, dorsal edge of femora with short spines; 9) tarsal claws 0.5 times length of tarsi and with minute denticles; 10) hind wing pads present or absent; 11) posterior margin of abdominal terga with spines (Fig. 55I); 12) gills (Fig. 55J) present on segments I-VII, elongate, two times length of each segment; 13) posterior margin of paraprocts with spines (Fig. 55K); 14) terminal filament subequal in length to cerci. Distribution: North, Central and South America. Biology: Specimens of this genus have been collected in many different habitats. The nymphs are often found in rivers with rocky substrate. KEY

TO

SOUTH AMERICAN PARACLOEODES

Nymphs 1

Hind wing pads absent; prostheca of right mandible elongate and pectinate (Fig. 55M); 2 small tubercles between fore wing pads (Figs. 55O-P) .................... Paracloeodes binodulus Hind wing pads present; prostheca of right mandible slender and bifid (Figs. 55D, 56C); without tubercles between fore wing pads ........................................................ 2 Antennae short, 2.0 to 2.5 times of head capsule; labrum (Fig. 55A), dorsally with a pair of long subapical setae near midline and three-four setae laterally ........... 3 Antennae long, 3.5 to 4.0 times of head capsule; labrum (Fig. 56A), dorsally without isolated pair of long subapical setae near midline, only scattered short setae present ............................................................................................. Paracloeodes ibicui Nymphs small, 3.9 to 4.1 mm; gills slender as in Fig. 56J; hypopharynx similar to Fig. 55N; femora with medial brown marking on anterior face ................................ ......................................................................................................... Paracloeodes leptobranchus Nymphs large, 4.5 to 5.5 mm; gills broader in midregion (Fig. 55J); hypopharynx as in Fig. 55E; femora without medium brown marking on anterior face (Fig. 55H) ......................................................................................................... Paracloeodes eurybranchus

– 2(1) – 3(2) –

CLAVE

PARA LOS

PARACLOEODES

SUDAMERICANOS

Ninfas 1 –

Pterotecas posteriores ausentes; prosteca de la mandíbula derecha alargada y pectinada (Fig. 55M); 2 tubérculos pequeños entre las pterotecas anteriores (Figs. 55O-P) ................................................................................................. Paracloeodes binodulus Pterotecas posteriores presentes; prosteca de la mandíbula derecha delgada y bífida (Figs. 55D, 56C); sin tubérculos entre las pterotecas anteriores .................. 2

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Plate 55. Paracloeodes, nymphs. Figs. 55A-K. P. eurybranchus. Figs. 55A-G, mouthparts: 55A, labrum, left d.v., right v.v.; 55B, labrum, apical setae; 55C, left mandible v.v.; 55D, right mandible v.v.; 55E, hypopharynx; 55F, maxilla v.v.; 55G, labium, left d.v., right v.v. 55H, leg I; 55I, posterior margin of abdominal tergum IV; 55J, gill IV; 55K, paraproct. Figs. 55L-R, P. binodulus: 55L, labrum d.v.; 55M, left mandible; 55N, hypopharynx; 55O, thorax d.v.; 55P, thorax l.v.; 55Q, tarsal claw I; 55R, gill IV. (D = denticles; I = incisors; L = lingua; P = prostheca; SL = superlingua; T = tubercle; Roman numbers = segment numbers). Figs. 55L-R modified from LUGO-ORTIZ & MCCAFFERTY, 1996c.

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Antenas cortas, 2,0 a 2,5 veces el ancho de la cabeza; labro (Fig. 55A), dorsalmente con un par de setas largas subapicales hacia la línea media y tres-cuatro setas hacia laterales ................................................................................................................................. 3 Antenas largas, 3,5 a 4,0 veces el ancho de la cabeza; labro (Fig. 56A), dorsalmente sin par de setas largas subapicales aisladas, solo con setas cortas distribuidas aleatoriamente .......................................................................................... Paracloeodes ibicui Ninfas pequeñas, 3,9 a 4,1 mm; branquias alargadas como en la Fig. 56J; hipofaringe similar a la Fig. 55N; fémures con una mancha castaña en el medio de la superficie anterior ................................................................... Paracloeodes leptobranchus Ninfas grandes, 4,5 a 5,5 mm; branquias más anchas en el medio (Fig. 55J); hipofaringe como en la Fig. 55E; fémures sin manchas (Fig. 55H) .......................... ......................................................................................................... Paracloeodes eurybranchus

Paracloeodes binodulus LUGO-ORTIZ & MCCAFFERTY Paracloeodes binodulus LUGO-ORTIZ & MCCAFFERTY, 1996c: 163; DOMÍNGUEZ et al., 2002: 462; SALLES et al., 2004: 6.

Taxonomy: This species, known from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Length of body, 2.73.4 mm; 2) antennae two times of head capsule; 3) labrum (Fig. 55L), dorsally with scattered long fine setae; 4) prostheca of right mandible slender and pectinate (Fig. 55M); 5) hypopharynx as in Fig. 55N; 6) hind wing pads absent; 7) tarsal claws with a row of 1012 minute denticles (Fig. 55Q); 8) abdominal gills as in Fig. 55R; 9) two small tubercles between fore wing pads (Figs. 55O-P). Distribution: Brazil and Colombia. Paracloeodes eurybranchus LUGO-ORTIZ & MCCAFFERTY Paracloeodes eurybranchus LUGO-ORTIZ & MCCAFFERTY, 1996c: 166.

Taxonomy: This species, known from nymphs and the female imago, can be distinguished from the other species of the genus by the following combination of characters. In the nymphs: 1) Length of body 4.5-5.5 mm; 2) antennae 2-2.5 times of head capsule; 3) labrum (Fig. 55A), dorsally with a pair of subapical setae near midline and 3-4 setae laterally; 4) prostheca of right mandible slender and apically bifid (Fig. 55D); 5) hypopharynx as in Fig. 55E; 6) hind wing pads present; 7) tarsal claws with minute denticles basally and four to six longer denticles in midregion (Fig. 55H); 8) abdominal gills as in Fig. 55J. Distribution and Biology: Argentina and Brazil. The nymphs were collected in rivers with rocky substrate. Paracloeodes ibicui LUGO-ORTIZ & MCCAFFERTY Paracloeodes ibicui LUGO-ORTIZ & MCCAFFERTY, 1996c: 167.

Taxonomy: This species, known from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Length of body, 3.0

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I

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Plate 56. Paracloeodes, nymphs. Figs. 56A-I, P. ibicui. Figs.56A-F, mouthparts: 56A, labrum, left d.v., right v.v.; 56B, left mandible v.v.; 56C, right mandible v.v.; 56D, hypopharynx; 56E, maxilla v.v.; 56F, labium, left d.v., right v.v. 56G, leg I; 56H, abdomen d.v.; 56I, gill IV. Fig. 56J, P. leptobranchus, gill IV. (D = denticles; I = incisors; L = lingua; P = prostheca; SL = superlingua; Roman numbers = segment numbers). Fig. 56J modified from LUGO-ORTIZ & MCCAFFERTY, 1996c.

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mm; 2) antennae 3.5-4 times of head capsule; 3) labrum (Fig. 56A), dorsally with scattered short setae; 4) prostheca of right mandible slender and basally bifid (Fig. 56C); 5) hypopharynx as in Fig. 56D; 6) hind wing pads present; 7) tarsal claws with minute denticles basally and four to six larger denticles in midregion (Fig. 56G); 8) abdominal gills as in Fig. 56I; 9) abdominal terga I-IX with longitudinal median dark stripe on. Distribution and Biology: Argentina and Paraguay. The nymphs were collected in December. They were found in the Paranaense area, in a river with rocky substrate. Paracloeodes leptobranchus LUGO-ORTIZ & MCCAFFERTY Paracloeodes leptobranchus LUGO-ORTIZ & MCCAFFERTY, 1996c: 168.

Taxonomy: This species, known from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Length of body, 3.94.1 mm; 2) antennae 2.5 times of head capsule; 3) labrum, dorsally with a pair of subapical setae near midline and 3-4 near lateral margin (similar to Fig. 55A); 4) prostheca of right mandible bifid; 5) hypopharynx similar to Fig. 55N; 6) hind wing pads present; 7) tarsal claws with minute denticles basally and 4-6 larger denticles in midregion; 8) abdominal gills as in Fig. 56J. Distribution: Brazil and Paraguay. Genus Prebaetodes LUGO-ORTIZ & MCCAFFERTY Prebaetodes LUGO-ORTIZ & MCCAFFERTY, 1996b: 371. (Type-species: Prebaetodes sitesi LUGO-ORTIZ & MCCAFFERTY, original designation).

Taxonomy: This genus was established by LUGO-ORTIZ & MCCAFFERTY (1996b). They included one species, Prebaetodes sitesi, known from nymphs. In the same paper they stated that this genus belongs to the Baetodes complex. Adult Characteristics: Unknown. Nymphal Characteristics: 1) Antennae (Fig. 57A) long, 3 times of head capsule, scape 1.5 times length of pedicel, only pedicel with a row of long setae on external margin and with scale-like tubercle; 2) labrum (Fig. 57B), width 3 times length, anterior margin with two kinds of bipectinated setae (Fig. 57C): basally bifid setae near midline and apically bifid setae at lateral margin; ventrally with a row of branched setae; 3) mandibles (Figs. 57D-E) with incisors fused apically; 4) left mandible (Fig. 57D) with prostheca with denticles apically, right prostheca reduced to a short apine (Fig. 57E); 5) hypopharynx (Fig. 57F): lingua shorter than superlinguae and with a pair of tufts of setae antero-laterally (Fig. 57F); 6) maxillae (Fig. 57G) robust, palpi two-segmented, slightly longer than galealacinia; 7) labium (Fig. 57H): paraglossae 1.3 times longer and 2.5 times wider than glossae, glossae with 2-3 rows and paraglossae with 3 rows of long pectinate setae, segment II of palpi slender and without projection; 8) legs (Fig. 57I): dorsal edge of femora with a row of long setae alternating with stout spines, tarsal claws with one row of denticles and with a subapical strong seta (Fig. 57J); 9) hind wing pads rudimentary; 10) paraproct with posterior margin smooth; 11) terminal filament 0.25-0.30 times the length of cerci. Distribution: Colombia, Ecuador and Venezuela.

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I

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P P

D

E

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I

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D SS

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Plate 57. Prebaetodes sitesi, nymph. Figs. 57A-K. 57A, antenna: scape, pedicel and first segments of flagellum. Figs. 57B-H, mouthparts: 57B, labrum, left d.v., right v.v.; 57C, labrum: setae of apical margin, left d.v., right v.v.; 57D, left mandible v.v.; 57E, right mandible v.v.; 57F, hypopharynx v.v.; 57G, maxilla v.v.; 57H, labium, left d.v., right v.v. 57I, leg I; 57J, tarsal claw I; 57K, posterior margin of tergum IV. (D = denticles; I = incisors; L = lingua; P = prostheca; SL = superlingua; SS = subapical seta; T = tuft of setae; Roman numbers = segment numbers).

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Prebaetodes sitesi LUGO-ORTIZ & MCCAFFERTY Prebaetodes sitesi LUGO-ORTIZ & MCCAFFERTY, 1996b: 371.

Taxonomy: This species, known from nymphs, can be distinguished by the following combination of characters: 1) Labrum (Fig. 57B), dorsally with a row of long subapical setae; 2) tarsal claws with a row of 7-10 denticles (Fig. 57J); 3) posterior margin of abdominal terga with rounded spines (Fig. 57K). Distribution: Discussed in the generic description. Genus Rivudiva LUGO-ORTIZ & MCCAFFERTY Rivudiva LUGO-ORTIZ & MCCAFFERTY, 1998: 61; ORTH et al., 2000: 36. (Type-species: Rivudiva minantenna LUGO-ORTIZ & MCCAFFERTY, original designation).

Taxonomy: This genus was established by LUGO-ORTIZ & MCCAFFERTY in 1998. They included two species, Rivudiva minantenna (the type species) and R. trichobasis. Both species are known only from nymphs. After examination of the type material of both species we add some characters to the original description. Adult Characteristics: Unknown. T

B A

III

II

I

D

C

Plate 58. Rivudiva minantenna, nymph. Figs. 58A-D. Figs. 58A-D, mouthparts: 58A, labrum, left d.v., right v.v.; 58B, hypopharynx v.v.; 58C, maxilla v.v.; 58D, labium, left d.v., right v.v. (T = tuft of setae; Roman numbers = segment numbers).

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Nymphal Characteristics: 1) Antennae short, 1.5 times of head capsule; 2) labrum (Figs. 58A, 59A) subrectangular, anterior margin with branched setae (Fig. 59B); 3) mandibles (Figs. 59C, E) with incisors cleft in two sets; 4) left mandible (Fig. 59C) with prostheca with two denticles apically, molars with constrictions (Fig. 59D), prostheca of right mandible slender and bifid (Fig. 59E); 5) hypopharynx (Fig. 58B): lingua broad twice width of superlinguae; 6) maxillae (Figs. 58C, 59F) with palpi two segmented and at least 1.5 times length of galea-lacinia; 7) labium (Figs. 58D, 59G-H): glossae subequal in length I P

P

A E C

B

D

III II

I

F

G

H

I

K J D

Plate 59. Rivudiva trichobasis, nymph. Figs. 59A-K. Figs. 59A-H, mouthparts: 59A, labrum, left d.v., right v.v.; 59B, labrum: apical margin; 59C, left mandible v.v.; 59D, left mandible, molar detail; 59E, right mandible; 59F, maxilla v.v.; 59G, labium, palpi, left d.v., right v.v.; 59H, labium: glossae and paraglossae, left d.v., right v.v.; 59I, leg I; 59J, tarsal claw I; 59K, paraproct. (D = denticles; I = incisors; P = prostheca; Roman numbers = segment numbers).

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but wider than paraglossae, segment II of palpi broadly rounded, segment III reduced and conical; 8) leg (Fig. 59I): femora with dorsal and ventral edge with a row of long spine-like setae; 9) tarsal claws (Fig. 59J) with two rows of denticles; 10) posterior margin of abdominal terga with spines; 11) posterior margin of paraprocts with spines (Fig. 59K). Distribution: Brazil and French Guiana. KEY

TO

SOUTH AMERICAN RIVUDIVA

Nymphs 1

Hind wing pads present; antennae: scape subequal in length to pedicel, both without setae; lingua with a tuft of setae apically (Fig. 58B); paraglossae with three rows of spine-like setae (Fig. 58D) ................................................ Rivudiva minantenna Hind wing pads absent; antennae: scape 1.5 times the length of pedicel, both with spine-like setae; lingua without a tuft of setae apically; paraglossae with a tuft of spine-like setae (Fig. 59H) .................................................................. Rivudiva trichobasis

–

CLAVE

PARA LOS

RIVUDIVA

SUDAMERICANOS

Ninfas 1 –

Pterotecas posteriores presentes; antenas: escapo subigual en longitud al pedicelo y ambos sin setas; lingua con un mechón de setas apicales (Fig. 58B); paraglosas con tres hileras de setas como espinas (Fig. 58D) ...................... Rivudiva minantenna Pterotecas posteriores ausentes; antenas: escapo 1,5 la longitud del pedicelo, ambos cubiertos con setas como espinas; lingua sin un mechón de setas apicales; paraglosas con 1 mechón de setas como espinas (Fig. 59H) ............................ Rivudiva trichobasis

Rivudiva minantenna LUGO-ORTIZ & MCCAFFERTY Rivudiva minantenna LUGO-ORTIZ & MCCAFFERTY, 1998: 63.

Taxonomy: Rivudiva minantenna, known only from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Antennae: scape subequal in length to pedicel and both without setae; 2) lingua with a tuft of setae apically (Fig. 58B); 3) paraglossae with 3 rows of spine-like setae (Fig. 58D); 4) hind wing pads present. Distribution: Brazil. Rivudiva trichobasis LUGO-ORTIZ & MCCAFFERTY Rivudiva trichobasis LUGO-ORTIZ & MCCAFFERTY, 1998: 64.

Taxonomy: Rivudiva trichobasis known only from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Antennae: scape 1.5 times length of pedicel and both with spine-like setae; 2) lingua without a tuft of setae apically; 3) paraglossae with a tuft of spine-like setae (Fig. 59H); 4) hind wing pads absent. Distribution: Brazil.

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Genus Spiritiops LUGO-ORTIZ & MCCAFFERTY Spiritiops LUGO-ORTIZ & MCCAFFERTY, 1998: 64; ORTH et al., 2000: 36. (Type-species: Spiritiops silvudus, LUGO-ORTIZ & MCCAFFERTY, original designation).

Taxonomy: Spiritiops was established by LUGO-ORTIZ & MCCAFFERTY (1998). They described one species in this genus, S. silvudus, known from nymphs. Although this genus has been reported from other country, no other species have been described. After examination of the type material we have some generic characters. Adult Characteristics: Unknown. Nymphal Characteristics: 1) Antennae long, 3 times of head capsule; 2) labrum (Fig. 60A) wider than long, anterior margin with bifid pectinate setae (Fig. 60B); 3) mandibles (Figs. 60C-D) with incisors fused apically; 4) left mandible (Fig. 60C) with prostheca robust and with denticles apically; 5) right mandible (Fig. 60D) with setae between prostheca and mola, prostheca robust with a small transverse bipectinate seta; 6) hypopharynx (Fig. 60E): lingua subequal in length to superlinguae and with a small rounded projection apically; 7) maxillae (Fig. 60F) with palpi two-segmented, segment II with a constriction; 8) labium (Fig. 60G) with glossae subequal to paraglossae, glossae with a row and paraglossae with 3 rows of spine-like setae, segment II of palpi with slight distomedial projection, segment III small and conical; 9) legs (Fig. 60H) robust, dorsal edge of femora with a row of spine-like setae; 10) tarsal claws (Fig. 60I) with 2 rows of denticles and with a strong subapical seta; 11) hind wing pads present; 12) terminal filament 0.80 times the length of cerci. Distribution and Biology: Brazil, Colombia and French Guiana. This genus was collected in a waterfall, on the surfaces of vertical rocks where water flows. Spiritiops silvudus LUGO-ORTIZ & MCCAFFERTY Spiritiops silvudus LUGO-ORTIZ & MCCAFFERTY, 1998: 66; DOMÍNGUEZ et al., 2002: 462; SALLES et al., 2004: 6.

Taxonomy: Spiritiops silvudus, known from nymphs, can be distinguished by the following combination of characters: 1) Labrum (Fig. 60A), dorsally with two pairs of long subapical setae; 2) posterior margin of abdominal terga with rounded spines (Fig. 60J); 3) posterior margin of paraprocts with spines (Fig. 60K). Distribution: Brazil and Colombia. Genus Tomedontus LUGO-ORTIZ & MCCAFFERTY Tomedontus LUGO-ORTIZ & MCCAFFERTY, 1995a: 237. (Type-species: Tomedontus primus LUGO-ORTIZ & MCCAFFERTY, original designation).

Taxonomy: This genus was established by LUGO-ORTIZ & MCCAFFERTY (1995a). They included one species, T. primus, known from nymphs. Since the original description no other species have been described. Adult Characteristics: Unknown.

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Nymphal Characteristics: 1) Frontal keel present; 2) labrum (Fig. 61A) trapezoidal, with a row of flattened setae subapically and dorsally; 3) mandibles (Figs. 61B-C) with incisors fused apically, setae between prostheca and mola, base of incisors with a tuft of setae, prosthecae robust with 3-4 denticles apically; 4) right mandible with a row of short setae on inner margin of incisors (Fig. 61C); 5) hypopharynx (Fig. 61D): superlinguae longer than lingua, lingua with a pair of pointed projections laterally, superlinguae with

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Plate 60. Spiritiops silvudus, nymph. Figs. 60A-K. Figs. 60A-G, mouthparts: 60A, labrum, left d.v., right v.v.; 60B, labrum: apical margin; 60C, left mandible v.v.; 60D, right mandible; 60E, hypopharynx; 60F, maxilla v.v.; 60G, labium, left d.v., right v.v. 60H, leg I; 60I, tarsal claw; 60J, posterior margin of abdominal tergum IV; 60K, paraproct. (C = constriction; D = denticles; L = lingua; P = prostheca; S = setae; SS = subapical seta; SL = superlingua; TS = transverse seta; Roman numbers = segment numbers).

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pointed apical projections; 6) maxillae (Fig. 61E) robust with palpi two segmented, longer than galea-lacinia; 7) labium (Fig. 61F): glossae 2/3 length of paraglossae, glossae with a row and paraglossae with 3 rows of long setae, segment II of palpi with a digitiform distomedial projection, segment III conical; 8) leg (Fig. 61G) robust, dorsal edge of femora with a row of short spines; 9) tarsal claws with one row of 5-7 small denticles; 10) hind wing pads absent; 11) abdominal gills present on abdominal segments I-VI. Distribution: Brazil (Amazonas) and Colombia. P

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Plate 61. Tomedontus primus, nymph. Figs. 61A-J. Figs. 61A-F, mouthparts: 61A, labrum d.v.; 61B, left mandible v.v.; 61C, right mandible v.v.; 61D, hypopharynx v.v.; 61E, maxilla v.v.; 61F, labium, left d.v., right v.v. 61G, leg I; 61H, posterior margin of abdominal tergum IV; 61I, gill V; 61J, paraproct. (L = lingua; P = prostheca; S = setae; SL = superlingua; Roman numbers = segment numbers). Figs. 61AJ modified from LUGO-ORTIZ & MCCAFFERTY, 1995a.

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Tomedontus primus LUGO-ORTIZ & MCCAFFERTY Tomedontus primus LUGO-ORTIZ & MCCAFFERTY, 1995a: 239; DOMÍNGUEZ et al., 2002: 462.

Taxonomy: This species, known from nymphs, can be distinguished by the following combination of characters: 1) Posterior margin of abdominal terga with spines (Fig. 61H); 2) gills with tracheae weakly tinged with pigment (Fig. 61I); 3) posterior margin of paraprocts with irregular spines (Fig. 61J). Distribution: Discussed in the generic description. Genus Tupiara SALLES, LUGO-ORTIZ, DA-SILVA & FRANCISCHETTI Tupiara SALLES et al., 2003a: 24. (Type-species: Tupiara ibirapitanga SALLES et al., original designation).

Taxonomy: This genus was established by SALLES et al. (2003a). They included one species, Tupiara ibirapitanga, known from nymphs and adults. Adult Characteristics: 1) Compound eyes oval, height of stalk half of eye diameter (Fig. 62A); 2) fore wings with paired marginal intercalary veins (Fig. 62B); 2) hind wing with two longitudinal veins, costal process acute, located in basal third of anterior margin (Fig. 62C); 3) genitalia (Fig. 62D): forceps three-segmented, basal segments with a small projection apically, apical segments elongate. A

D

A

B

C IV Plate 62. Tupiara ibirapitanga, male imago. Figs. 62A-D. 62A, head l.v.; 62B, fore wing; 62C, hind wing; 62D, genitalia v.v. (IV = intercalary veins). Figs. 62A-D modified from SALLES et al., 2003a.

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Nymphal Characteristics: 1) Frontal keel present; 2) antennae long, 4-5 times of head capsule; 3) labrum (Fig. 63A): as long as wide, anterior margin with two kind of setae: simple near midline and branched laterally (Fig. 63B); 4) left mandible (Fig. 63C) with incisors partially cleft in two sets, prostheca robust with 2-3 denticles apically, molars with constrictions; 5) right mandible (Fig. 63D) with incisors cleft in two sets, prostheca slender and bifid; 6) hypopharynx (Fig. 63E): lingua slightly longer than superlinguae and with a pointed medial projection, superlinguae rounded apically; 7) maxillae (Fig. 63F) with palpi two-segmented and longer than galea-lacinia; 8) labium (Fig. 63G): glossae 0.7

P

A

B

C

SL

L

E

P

D III

II

F

H I

G

J D

I

L

K

Plate 63. Tupiara ibirapitanga, nymph. Figs. 63A-L. Figs. 63A-G, mouthparts: 63A, labrum, left d.v., right v.v.; 63B, labrum, setae of apical margin; 63C, left mandible v.v.; 63D, right mandible v.v.; 63E, hypopharynx v.v.; 63F, maxilla v.v.; 63G, labium, left d.v., right v.v. 63H, leg I; 63I, tarsal claw I; 63J, posterior margin of abdominal tergum IV; 63K, gill IV; 63L, paraproct. (D = denticles; L = lingua; P = prostheca; SL = superlingua; Roman numbers = segment numbers).

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times length and half the width of paraglossae, paraglossae with 3 rows of spine-like setae, segment II of palpi medially rounded, segment III conical; 9) legs (Fig. 63H) robust, dorsal edge of femora with a row of short spines, tibiae 1.5 times length of tarsi; 10) tarsal claws (Fig. 63I) with 2 rows of denticles, one longer than the other; 11) terminal filament subequal in length to cerci. Distribution: Brazil (Amazonas, Minas Gerais, Rio de Janeiro). Tupiara ibirapitanga SALLES, LUGO-ORTIZ, DA-SILVA & FRANCISCHETTI Tupiara ibirapitanga SALLES et al., 2003a: 25.

Taxonomy: This species, known from nymphs and adults, can be distinguished by the following combination of characters. In the male imago: 1) compound eyes oval (Fig. 62A). In the nymphs: 1) Labrum (Fig. 63A), dorsally with a pair of subapical setae near midline and 2-3 setae laterally; 2) posterior margin of abdominal terga with spines (Fig. 63J); 3) abdominal gills present on segments I-VII, rounded and 1.5 times length of each segment (Fig. 63K); 4) posterior margin of paraprocts with spines (Fig. 63L). Distribution: Discussed in the generic description. Genus Varipes LUGO-ORTIZ & MCCAFFERTY Varipes LUGO-ORTIZ & MCCAFFERTY, 1998: 67-70; SALLES & BATISTA, 2004: 1; NIETO, 2004b. (Type-species: Varipes lasiobrachius LUGO-ORTIZ & MCCAFFERTY, original designation).

Taxonomy: Varipes was established for a single species, V. lasiobrachius, described from nymphs from Ecuador and Colombia. Recently, Varipes helenae from Brazil was described by SALLES & BATISTA (2004) based also on nymphs. These authors noticed some generic character variation between their new species and the type species. In the same year, after examining the type material of V. lasiobrachius, NIETO (2004b) redescribed the genus, and describing three new species. Two of the new species described were from Argentina, the other from Bolivia. In the same paper the adult was described for the first time. Adult Characteristics: 1) Turbinate eyes oval (Fig. 64A); 2) fore wings with paired marginal intercalary veins (Fig. 64B); 3) hind wings absent; 4) forceps three-segmented; basal segment with internal projection in middle of segment (Fig. 64C), hind margin of styliger plate with a small projection on both sides (Fig. 64C). Nymphal Characteristics: 1) Antennae: scape and pedicel with scale-like tubercles (Fig. 65A); 2) labrum with bifid marginal setae (Figs. 65B, 66A, G, 67A); 3) left mandible with incisors fused apically, prostheca with 2-3 denticles apically (Fig. 65C), molars with constrictions (Fig. 65D); 4) right mandible with prostheca reduced to simple (Fig. 65E) or bifid (Figs. 66H, 67B) long fine seta, but never robust with denticles apically; 5) hypopharynx (Figs. 65F, 67C): lingua subequal in length to superlinguae; 6) maxillae with palpi two-segmented, 1-1.5 times length of galea-lacinia (Figs. 65G, 66C, E, I, 67D); 7) labium: glossae subequal to paraglossae, segment II of palpi with a distomedial projection (Figs. 65H, 66J, 67E); 8) femora with abundant long spine-like setae on dorsal and ventral margins (Figs. 65I, K, 66K, M, 67F, H); 9) femora II and III with a transverse row of long

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spine-like setae on proximal third (Figs. 65K; 66M; 67H); 10) tarsal claws wide (Figs. 65J, 66L) or elongate (Fig. 67G), with two rows of denticles; 11) abdomen (Fig. 65L) with color pattern of two dark brown bands, one at anterior of abdomen with segments I, I-III or II-III dark, and the other at posterior of abdomen with segments VII-VIII, VII-IX or VIII dark; 12) posterior margins of abdominal terga with spines (Fig. 65M). Distribution: Argentina, Bolivia, Brazil, Colombia, Costa Rica, Ecuador, Panamá and Peru. Biology: The nymphs have been collected from waters with sandy substrate. KEY

TO

SOUTH AMERICAN VARIPES

Nymphs 1 –

Femora I without transverse row of spine-like setae (Fig. 67F) ........ Varipes singuil Femora I with transverse row of spine-like setae (Figs. 65I, 66K) ......................... 2

IP P

C A

B IV

Plate 64. Varipes singuil, male imago. Figs. 64A-C. 64A, lateral view; 64B, fore wing; 64C, genitalia v.v. (IV = intercalary vein; IP = internal projection of the basal segment; P = projection of the hind margin of styliger plate). Figs. 64A-C modified from NIETO, 2004b.

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2(1)

Femora I with transverse row of spine-like setae in proximal third (Fig. 65I), antennae short, about 1.5 times of head capsule; right prostheca reduced to a simple long fine seta (Fig. 65E) ...................................................................................... 3 Femora I with transverse row of spine-like setae in distal half (Fig. 66K), antennae long, about three times of head capsule; right prostheca slender and bifid (Fig. 66H) ...................................................................................................................................... 4 Hypopharynx: lingua and superlinguae rounded apically (Fig. 65F); maxillae with palpi 1.4 to 1.5 times longer than galea-lacinia (Fig. 65G); tarsal claws with two rows of 9 or 10 denticles each (Fig. 65J) ................................................Varipes cajuato Hypopharynx: lingua and superlinguae pointed apically (Fig. 66D); maxillae with palpi reaching apex of galea-lacinia (Fig. 66E); tarsal claws with two rows of 5 to 7 denticles each (Fig. 66F) ................................................................ Varipes lasiobrachius Labrum: dorsal subapical setae as in Fig. 66G; hypopharynx: superlingua rounded (as in Fig. 65F), maxillae with palpi elongate, reaching beyond apex of galealacinia (Fig. 66I) .......................................................................................... Varipes minutus Labrum: dorsal subapical setae as in Fig. 66A; hypopharynx: superlinguae pointed (Fig. 66B); maxillae with palpi robust barely reaching apex of galea-lacinia (Fig. 66C) ................................................................................................................ Varipes helenae

– 3(2) – 4(2) –

CLAVE

PARA LOS

VARIPES

SUDAMERICANOS

Ninfas 1 – 2(1) – 3(2) – 4(2) –

Fémures I sin una hilera transversa de setas como espinas (Fig. 67F) ...... Varipes singuil Fémures I con una hilera transversa de setas como espinas (Figs. 65I, 66K) .......... 2 Fémures I con una hilera transversa de setas como espinas en el tercio basal (Fig. 65I); antenas cortas, 1,5 veces el ancho de la cabeza; prosteca derecha reducida a una seta larga y fina (Fig. 65E) ........................................................................................ 3 Fémures I con una hilera transversa de setas como espinas en el tercio distal (Fig. 66K); antenas largas, casi 3 veces el ancho de la cabeza, prosteca derecha alargada y bífida (Fig. 66H) .............................................................................................................. 4 Hipofaringe (Fig. 65F): lingua y superlingua redondeada apicalmente; maxilas (Fig. 65G) con palpos 1,4 a1,5 veces más largos que la galea-lacinia; uñas tarsales con dos hileras de 9 a 10 dentículos (Fig. 65J) ..............................................Varipes cajuato Hipofaringe: lingua y superlingua agudas apicalmente (Fig. 66D); maxilas con palpos subigual en longitud al ápice de la galea-lacinia (Fig. 66E); uñas tarsales con 2 hileras de 6 a 7 dentículos (Fig. 66F) ......................................... Varipes lasiobrachius Labro dorsalmente con setas subapicales como en la Fig. 66G; hipofaringe: superlinguas redondeadas apicalmente (como en la Fig. 65F), maxilas: palpos alargados, más largos que el ápice de la galea-lacinia (Fig. 66I) ......... Varipes minutus Labro dorsalmente con setas subapicales como en la Fig. 66A; hipofaringe: superlinguas agudas (Fig. 66B); maxilas: palpos robustos apenas subiguales al ápice de la galea-lacinia (Fig. 66C) .......................................................... Varipes helenae

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P

P

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D

S

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E L

SL

III II

F

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I TS

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L D

K

M

TS Plate 65. Varipes cajuato, nymph. Figs. 65A-N. 65A, antenna d.v. Figs. 65B-H, mouthparts: 65B, labrum, left d.v., right v.v.; 65C, left mandible v.v.; 65D, left mandible, molars detail; 65E, right mandible v.v.; 65F, hypopharynx v.v.; 65G, maxilla v.v.; 65H, labium, left d.v., right v.v. 65I, leg I; 65J, detail of tarsal claw I; 65K, leg III; 65L, abdominal terga I-X; 65M, posterior margin of tergum IV; 65N, gill I. (D = denticles; I = incisors; L = lingua; P = prostheca; S = scales; SL = superlingua; TS = transverse row of setae; Roman numbers = segment numbers). Figs. 65A-N modified from NIETO, 2004b.

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Varipes cajuato NIETO Varipes cajuato NIETO, 2004b: 164.

Taxonomy: Varipes cajuato is known only from nymphs, which can be distinguished from the other species of the genus by the following combination of characters: 1) Antennae short, about 1.5 times of head capsule; 2) labrum with dorsal surface with two short subapical setae near midline and three-four short setae laterally (Fig. 65B); 3) right mandible with incisors fused apically, prostheca (Fig. 65E) reduced to a simple long fine seta; 4) lingua and superlinguae rounded apically (Fig. 65F); 5) maxillae with palpi 1.5 times length of galea-lacinia (Fig. 65G); 6) segment II of labial palpi (Fig. 65H) with a thumb-like distomedial projection, segment III broad and rounded; 7) femora (Figs. 65I, K) with a transverse row of long spine-like setae on proximal third; 8) tarsal claws (Fig. 65J) widened with 2 rows of 9-10 denticles each; 9) abdominal gills I and II not racquet-shape (Fig. 65N). Distribution and Biology: Bolivia. The nymphs were found in mountain rain forests. Varipes helenae SALLES & BATISTA Varipes helenae SALLES & BATISTA, 2004: 2; NIETO, 2004b: 166.

Taxonomy: Varipes helenae is known only from nymphs, which can be distinguished from the other species of the genus by the following combination of characters: 1) Antennae long, about 3 times of head capsule; 2) labrum (Fig. 66A) with dorsal surface with two long subapical setae near midline and 2-3 setae laterally; 3) right mandible (as in Fig. 66H) with incisors cleft in 2 sets, prostheca slender and bifid; 4) hypopharynx (Fig. 66B): lingua rounded and superlinguae pointed apically; 5) maxillae with palpi barely reaching apex of galea-lacinia (Fig. 66C); 6) segment II of labial palpi (as in Fig. 66J) with a rounded distomedial projection, segment III rounded; 7) femora I with a transverse row of long spine-like setae on distal half (as in Fig. 66K); 8) tarsal claws widened with two rows of fourfive denticles each (Fig. 66L); 9) abdominal gills I and II racquet-shaped (Figs. 66N-O). Distribution: Brazil (Mato Grosso). Varipes lasiobrachius LUGO-ORTIZ & MCCAFFERTY Varipes lasiobrachius LUGO-ORTIZ & MCCAFFERTY, 1998: 69-70; SALLES & BATISTA, 2004: 3; NIETO, 2004b: 167.

Taxonomy: Varipes lasiobrachius is known only from nymphs, which can be distinguished from the other species of the genus by the following combination of characters: 1) Antennae long, about 3 times of head capsule; 2) labrum with dorsal surface with 2 short subapical setae near midline and three-four short setae laterally (as in Fig. 65B); 3) right mandible with incisors fused apically, prostheca reduced to a simple long fine seta (as in Fig. 65E); 4) hypopharynx with lingua and superlinguae pointed apically (Fig. 66D); 5) maxillae with palpi reaching apex of galea-lacinia (Fig. 66E); 6) segment II of labial palpi with moderately developed and somewhat narrowly pointed distomedial projection, segment III broadly rounded (as in Fig. 65H); 7) femur I with a transverse row of long spine-like setae on

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L

SL

E A C

D

B F D III

I

G

II

P

J

I

H

K TS

L D

M N O

TS

Plate 66. Varipes, nymphs. Figs. 66A-C, V. helenae: 66A, labrum d.v.; 66B, hypopharynx v.v.; 66C, right maxilla v.v. Figs. 66D-F, V. lasiobrachius: 66D, hypopharynx v.v.; 66E, maxilla v.v.; 66F, tarsal claw I. Figs. 66G-O, V. minutus: 66G, labrum, left d.v., right v.v.; 66H, right mandible v.v.; 66I, right maxilla v.v.; 66J, labium, left d.v., right v.v.; 66K, leg I; 66L, tarsal claw I; 66M, leg III; 66N, gill I; 66O, gill II. (D = denticles; L = lingua; P = prostheca; SL = superlingua; TS = transverse row of setae; Roman numbers = segment numbers). Figs. 66A-C modified from SALLES & BATISTA, 2004; 66D-F from LUGO-ORTIZ & MCCAFFERTY, 1998; 66G-O modified from NIETO, 2004b.

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proximal third (as in Fig. 65I); 8) tarsal claws widened with two rows of five-seven denticles each (Fig. 66F); 9) abdominal gills I and II not racquet-shaped (as in Fig. 65N). Distribution: Colombia and Ecuador. Varipes minutus NIETO Varipes minutus NIETO, 2004b: 167.

Taxonomy: Varipes minutus is known from nymphs and male and female subimagos. This species can be distinguished from the other species of the genus by the following combination of characters. In the nymph: 1) Antennae long, about 3 times of head capsule; 2) labrum: dorsal surface with one long subapical setae near midline and two-three long setae laterally (Fig. 66G); 3) right mandible (Fig. 66H) with incisors cleft in 2 sets, prostheca slender and bifid; 4) hypopharynx: lingua and superlingua rounded apically (as in Fig. 65F); 5) maxillae with palpi 1.5 times length of galea-lacinia (Fig. 66I); 6) segment II of labial palpi (Fig. 66J) with a rounded distomedial projection, segment III rounded; 7) femora I (Fig. 66K) with a transverse row of long spine-like setae on distal half; 8) tarsal claws widened with two rows of four-five denticles each (Fig. 66L); 9) abdominal gills I and II (Figs. 66N-O) racquet-shaped. Distribution and Biology: Argentina (Córdoba, La Rioja and Tucumán). Varipes singuil NIETO Varipes singuil NIETO, 2004b: 169.

Taxonomy: Varipes singuil is known from nymphs and male imagos. This species can be distinguished from the other species of the genus by the following combination of characters. In the imago: 1) Eyes with apical portion yellowish, stalk orange-yellow; 2) abdomen with segment I yellowish-brown, segments II-VI pale yellow, segments VII-IX pale brown. In the nymph: 1) Antennae short, 1.5 times of head capsule; 2) labrum: dorsal surface with two subapical setae near midline and two-three long setae laterally (Fig. 67A); 3) right mandible (Fig. 67B) with incisors cleft in 2 sets, prostheca slender and bifid; 4) hypopharynx: lingua with a rounded projection apically (Fig. 67C); 5) maxillae with palpi 1.5 times length of galea-lacinia (Fig. 67D); 6) segment II of labial palp (Fig. 67E) with a large rounded distomedial projection, segment III conical; 7) femora I (Fig. 67F) without a transverse row of long spine-like setae; 8) tarsal claws (Fig. 67G) elongate with two rows of five-six denticles each; 9) abdominal gills I and II not racquet-shaped (Fig. 67I). Distribution and Biology: Argentina (Tucumán and Salta). The nymphs were found in mountain rain forest. Genus Waltzoyphius MCCAFFERTY & LUGO-ORTIZ Waltzoyphius MCCAFFERTY & LUGO-ORTIZ (in LUGO-ORTIZ & MCCAFFERTY, 1995a: 239); ORTH et al., 2000: 36; THOMAS & PERU, 2002: 15. (Type-species: Waltzoyphius fasciatus MCCAFFERTY & LUGO-ORTIZ, original designation).

Taxonomy: This genus was established by MCCAFFERTY & LUGO-ORTIZ (in LUGOORTIZ & MCCAFFERTY, 1995a). They included one species Waltzoyphius fasciatus, known

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from nymphs. Later, THOMAS & PERU (2002) described another species, W. roberti, known also from nymphs. The adult is currently unknown. Adult Characteristics: Unknown. Nymphal Characteristics: 1) Antennae long, 3 times of head capsule; 2) labrum (Fig. 68A) dorsally with lateral tufts of long setae, apical margin with two kinds of setae: 18-20 SL

L

P

A

C

B III

II

I

D

E

I F

H TS

G D

Plate 67. Varipes singuil, nymph. Figs. 67A-I. Figs. 67A-E, mouthparts: 67A, labrum, left d.v., right v.v.; 67B, right mandible v.v.; 67C, hypopharynx v.v.; 67D, maxilla v.v.; 67E, labium, left d.v., right v.v. 67F, leg I; 67G, tarsal claw I; 67H, leg III; 67I, gill IV. (D = denticles; L = lingua; P = prostheca; SL = superlingua; TS = transverse row of setae; Roman numbers = segment numbers). Figs. 67A-I modified from NIETO, 2004b.

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blunt setae at middle and seven-eight long spine-like setae laterally (Fig. 68B); 3) mandibles (Figs. 68C-E) with incisors cleft in two sets, inner set of incisors positioned at right angle to outer set; 4) left mandible with prostheca with 1-2 denticles apically (Fig. 68D), right prostheca bifid (Fig. 68F); 5) hypopharynx (Fig. 68G): lingua subequal in length to superlinguae, subrectangular and without projection apically; 6) maxillae: galea apically with a row of long setae, longer than marginal setae (Fig. 68I), palpi two-segmented, long, two times length of galea-lacinia (Fig. 68H); 7) labium (Fig. 68J): glossae equal in length and two times width of paraglossae, with row of spines on outer margin, paraglossae with a row of long setae, palpi segment II with a strong distomedial projection (Figs. 68M, O-P), segment III conical; 8) thoracic mesosternun with a projection (Fig. 68Q); 9) legs very long (Fig. 68K), femora with a dorsal row of short spines; 10) tarsal claws with 2 rows of denticles (Fig. 68L); 11) hind wing pads absent; 12) abdominal gills on segments I-VII, elongate, 1.5 times length of each segment; 13) cerci with three-five long spines laterally every four segments, terminal filament with two-three long spines centrally every four segments. Distribution: Brazil, Colombia, French Guiana, Paraguay. Biology: Nymphs inhabits small streams and rivers with slow current. KEY

TO

SOUTH AMERICAN WALTZOYPHIUS

Nymphs 1

Tarsal claws with 12 to 14 denticles in each row; segment II of labial palpi as in Fig. 68M; gills with tracheae weakly tinged with pigment (Fig. 68N) ............. Waltzoyphius fasciatus Tarsal claws with 8 to 10 denticles in each row; segment II of labial palpi as in Figs. 68O-P; gills with tracheae darkly pigmented (Fig. 68R) ............... Waltzoyphius roberti

–

CLAVE

PARA LOS

WALTZOYPHIUS

SUDAMERICANOS

Ninfas 1 –

Uñas tarsales con 12 a 14 dentículos en cada hilera; segmento II del palpo labial como en la Fig. 68M; branquias con las tráqueas pobremente pigmentadas (Fig. 68N) .................................................................................................... Waltzoyphius fasciatus Uñas tarsales con 8 a 10 dentículos en cada hilera; segmento II del palpo labial como en la Figs. 68O-P; branquias con las tráqueas principal y secundarias fuertemente pigmentadas (Fig. 68R) ............................................... Waltzoyphius roberti

Waltzoyphius fasciatus MCCAFFERTY & LUGO-ORTIZ Waltzoyphius fasciatus MCCAFFERTY & LUGO-ORTIZ (in LUGO-ORTIZ & MCCAFFERTY, 1995a: 241); DOMÍNGUEZ et al., 2002: 462; SALLES et al., 2004: 7.

Taxonomy: This species, known from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Segment II of labial palpi as in Fig. 68M; 2) tarsal claws with 12-14 denticles in each row; 3) gills with tracheae weakly tinged with pigment (Fig. 68N). Distribution: Brazil, Colombia and Paraguay.

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B L

E I LS

G J L

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H

D

M

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II

N I III III II

A

O I

P

II

Q

R

Plate 68. Waltzoyphius, nymphs. Figs. 68A-L, Waltzoyphius sp. Figs. 68A-J, mouthparts: 68A, labrum, left d.v., right v.v.; 68B, labrum, setae of apical margin; 68C, left mandible v.v.; 68D, left mandible, incisors and prostheca, detail; 68E, right mandible v.v.; 68F, right mandible, incisors and prostheca, detail; 68G, hypopharynx v.v.; 68H, maxilla v.v.; 68I, maxilla, apical portion, detail; 68J, labium, glossae and paraglossae, left d.v., right v.v. 68K, leg I; 68L, tarsal claw I. Figs. 68M-N, W. fasciatus: 68M, labium, left d.v., right v.v.; 68N, gill VI. Figs. 68O-R, W. roberti: Fig. 68O, labial palp v.v.; 68P, labial palp d.v.; 68Q, thoracic sterna v.v.; 68R, gill IV. (A = mesosternal apophysis; D = denticles; I = incisors; L = lingua; LS = long setae; P = prostheca; SL = superlingua; Roman numbers = segment numbers). Figs. 68M-N modified from LUGO-ORTIZ & MCCAFFERTY, 1995a; 68O-Q from THOMAS & PERU, 2002.

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Waltzoyphius roberti THOMAS & PERU Waltzoyphius roberti THOMAS & PERU, 2002: 15.

Taxonomy: This species, known from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Segment II of labial palpi as in Figs. 68O-P; 2) tarsal claws with 8-10 denticles in each rows; 3) gills with tracheae darkly pigmented (Fig. 68R). Distribution and Biology: French Guiana. This species was collected in small rivers or streams with slow currents. Specimens were found on dead plant material. Genus Zelusia LUGO-ORTIZ & MCCAFFERTY Zelusia LUGO-ORTIZ & MCCAFFERTY, 1998: 70. (Type-species: Zelusia principalis LUGO-ORTIZ & MCCAFFERTY, original designation).

Taxonomy: This genus was established by LUGO-ORTIZ & MCCAFFERTY (1998) to include one species, Zelusia principalis, known from nymphs. This is the only species presently described in the genus. Adult Characteristics: Unknown. Nymphal Characteristics: 1) Antennae with scape longer than pedicel; 2) labrum (Fig. 69A) wider than long, medially raised; 3) mandibles (Figs. 69B-C) with incisors fused apically; 4) left mandible (Fig. 69B): prostheca robust with denticles apically; 5) right mandible (Fig. 69C): prostheca bifid; 6) hypopharynx (Fig. 69D): lingua slightly longer than superlinguae and rounded apically; 7) maxillae (Fig. 69E) with palpi two-segmented, longer than galea-lacinia; 8) labium (Fig. 69F): glossae subequal in length to paraglossae and with a row of setae on the external margin, paraglossae with 2 rows of spine-like setae, segment II of palpi without lateral projection; 9) legs (Fig. 69G), dorsal edge of femora with a row of spine-like setae; 10) tarsal claws (Fig. 69H) with one row of small denticles; 11) hind wing pads absent; 12) posterior margin of abdominal terga with irregular spines (Fig. 69I); 12) gills present on segments II-VII, elongate (Fig. 69J); 13) terminal filament subequal in length to cerci. Distribution: Brazil and Colombia. Zelusia principalis LUGO-ORTIZ & MCCAFFERTY Zelusia principalis LUGO-ORTIZ & MCCAFFERTY, 1998: 72; DOMÍNGUEZ et al., 2002: 462; SALLES et al., 2004: 7.

Taxonomy: This species, known from nymphs, can be distinguished by the following combination of characters: 1) Labrum (Fig. 69A), dorsally with a row of long robust subapical setae on either side of midline; 2) maxillae (Fig. 69E) with palpi twice length of galea-lacinia, segment II 1.5 times length of segment I; 3) posterior margin of paraprocts as in Fig. 69K. Distribution: Discussed in the generic description.

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Plate 69. Zelusia principalis, nymph. Figs. 69A-K. Figs. 69A-F, mouthparts: 69A, labrum d.v.; 69B, left mandible v.v.; 69C, right mandible v.v.; 69D, hypopharynx v.v.; 69E, maxilla v.v.; 69F, labium, left d.v., right v.v. 69G, leg I; 69H, tarsal claw; 69I, posterior margin of abdominal tergum IV; 69J, gill IV; 69K, paraproct. (D = denticles; L = lingua; P = prostheca; SL = superlingua; Roman numbers = segment numbers). Figs. 69A-K modified from LUGO-ORTIZ & MCCAFFERTY, 1998.

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FAMILY CAENIDAE Taxonomy: The family Caenidae is represented in South America by four genera with 26 recognized species. Other species are considered nomina dubia in this work, because of poor descriptions and the fact that their types have been lost. Nevertheless, the genus Brachycercus is included in the key because of the presence of undescribed species in collections. There also exist in collections numerous undescribed species, especially in the genus Caenis. Caenis was recently treated by MALZACHER (2001) and there are modern treatments of the genera Cercobrachys by SOLDÁN (1986) and Brasilocaenis by MALZACHER (1986, 1990). BANKS (1900) established the tribe Caenini and KLAPÁLEK (1909) recognized it as a family. This placement was followed by ULMER (1920a) and SPIETH (1933), but not by NEEDHAM et al. (1935) who maintained it as a subfamily of the Baetidae. After the work by EDMUNDS & TRAVER (1954) the family level was used quite consistently in the literature. Adult Characteristics: 1) Eyes of imagos small (but at least 2 African genera show large eyes in males) and undivided; 2) hind wings absent, anal area of fore wings expanded, veins MP2 and IMP almost as long as vein MP1 (Fig. 72A); 3) a median lighter mark (ommation) present on mesonotum (Fig. 72B); 4) species small, generally with short and robust bodies, especially in females; and 5) forceps 1-segmented (Figs. 71A, 72D). Nymphal Characteristics: 1) Operculate gills of abdominal segment II quadrangular, overlapping on median line of body (Fig. 75A); 2) gills on abdominal segments III-VI fringed with long filaments; 3) gills on abdominal segment I filamentous (Fig. 75A); and 4) hind wing pads absent. Representatives of this family can initially be confused with those of the family Leptohyphidae, as much by their appearance as by the places where they are found, but they can be distinguished easily by wing venation and the presence of an ommation in the adults, and by the form of gills in the nymphs. Biology: The nymphs of this family are found in practically every type of aquatic habitat, generally preferring lagoons, lakes, and backwaters of rivers. They also can be found aggregated in vegetation in current zones and in floating lentic plants. They tolerate a wide range of temperatures and varying levels of pollution. Distribution: Practically cosmopolitan, except for New Zealand and various oceanic islands. KEYS TO SOUTH AMERICAN GENERA Adults 1 – 2(1) –

Prosternum longer than wide, with longitudinal ridges forming a triangular or trapezoid shaped structure (prosternal triangle) (Figs. 72F, J, P) ............................. 2 Prosternum wider than long, not as above ................................................................... 3 Styliger plate elongated caudally; forceps close to lateral margins of styliger plate or fused with it (Figs. 71A, F) ....................................................................... Brasilocaenis Styliger plate shaped differently, with an incision between the lateral margin and the forceps (Fig. 72D) ...............................................................................................Caenis

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Pedicel of antennae 1.5 to 2.0 times as long as scape (Fig. 70B) ........... Brachycercus Pedicel of antennae subequal to 1.3 times length of scape (Fig. 77C) ...... Cercobrachys

Nymphs 1 – 2(1) – 3(1) –

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Head with 3 ocellar tubercles (Figs. 70A, 77A), labial and maxillary palpi 2segmented, and gill basket present ................................................................................. 2 Head without ocellar tubercles (Fig. 75A), labial and maxillary palpi 3-segmented, and gill basket absent ........................................................................................................ 3 Pedicel of antennae a maximum of 1.1 to 1.3 times as long as scape (Fig. 77A); legs with setae as long as tibia (Fig. 77D) ................................................... Cercobrachys Pedicel of antennae a minimum of 1.5 to 2.0 times as long as scape (Fig. 70A); legs with setae much shorter than tibia (Fig. 70C) ..................................... Brachycercus Coxae of middle and hind legs with a large dorsolateral projection (Figs. 71I, K, M); dorsal surface of operculate gill with short strong and blunt spines; margin of the segments VIII and IX with strong teeth .............................................. Brasilocaenis Coxae sicle-shaped or with rounded projections of medium length (Figs. 76D-H) (abdominal gill I may be confused with a projection of hind coxa); operculate gill generally without short strong spines, sometimes with long bristles; margin of the segments VIII and IX mostly without teeth ........................................................Caenis PARA LOS

GÉNEROS

SUDAMERICANOS

Adultos 1 – 2(1) – 3(1) –

Presterno más largo que ancho, con carinas longitudinales formando una estructura triangular o trapezoidal (triángulo presternal) (Figs. 72F, J, P) ............. 2 Presterno más ancho que largo, no como arriba ........................................................ 3 Placa estilígera elongada posteriormente; fórceps cercanos a los márgenes laterales de la placa estilígera o fusionados con ella (Figs. 71A, F) ....... Brasilocaenis Placa estilígera de diferente forma, con una incisión entre su márgen lateral y los fórceps (Fig. 72D) ......................................................................................................Caenis Pedicelo de las antenas 1,5 a 2,0 veces más largo que el escapo (Fig. 70B) ............ ............................................................................................................................... Brachycercus Pedicelo de las antenas subigual a 1,3 veces la longitud del al escapo (Fig. 77C) ..... .............................................................................................................................. Cercobrachys

Ninfas 1 – 2(1) –

Cabeza con 3 tubérculos ocelares (Figs. 70A, 77A), palpos labiales y maxilares bisegmentados, cámara branquial presente ...................................................................... 2 Cabeza sin tubérculos ocelares (Fig. 75A), palpos labiales y maxilares trisegmentados, cámara branquial ausente ........................................................................ 3 Pedicelo de las antenas como máximo 1,1 a 1,3 veces más largo que el escapo (Fig. 77A); patas con setas tan largas como la tibia (Fig. 77D) ....................... Cercobrachys Pedicelo de las antenas como mínimo 1,5 a 2,0 veces más largo que el escapo (Fig. 70A); patas con setas más cortas que la tibia (Fig. 70C) .......................... Brachycercus

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Coxas medias y posteriores con una larga proyección dorsal (Figs. 71I, K, M); superficie dorsal de la branquia opercular con espinas romas cortas y fuertes; margen de los segmentos abdominales VIII y IX con fuerte dientes ...................... Brasilocaenis Coxas en forma de hoz o con proyecciones redondeadas de longitud media (Figs. 76D-H) (branquia abdominal I puede ser confundida con una proyección de la coxa posterior); branquia opercular generalmente sin espinas cortas, a veces con largas setas; márgenes de los segmentos abdominales VIII y IX generalmente sin dientes ........................................................................................................................... Caenis

Genus Brachycercus CURTIS Brachycercus CURTIS, 1834: 122; SOLDÁN, 1986: 291. (Type-species: Brachycercus harrisella CURTIS, subsequent designation by LESTAGE, 1924b: 61). Eurycaenis BENGTSSON, 1917: 186. (Type-species: Brachycercus harrisella CURTIS, original designation).

Taxonomy: Brachycercus, widely distributed in the Holarctic Region, was revised by SOLDÁN (1986). The record of distribution from Argentina is based on 1 species (Brachycercus tenella) described by NAVÁS (1932a), and its placement in this genus is dubious. Nevertheless, ORTH et al. (2000) confirmed the presence of this genus in South America based on a nymph of an undescribed species from French Guiana. Adults Characteristics: 1) Pedicel of antennae at least 1.5 to 2 times as long as scape (Fig. 70B); and 2) prosternum broad with coxae widely separated. Nymphal Characteristics: 1) Pedicel of antennae at least 1.5 to 2 times as long as scape (Figs. 70A-B); 2) ocellar tubercles well developed (Fig. 70A); 3) maxillary and labial palpi 2-segmented; 4) prosternum broad, and anterior coxae widely separated; 5) legs long and thin, with short setae (Fig. 70C); 6) tibiae approximately equal in length to tarsi and claws without denticles (Fig. 70C); and 7) posterolateral projections of abdominal segment VI not curved dorsally over abdomen (Figs. 70D-E). Distribution and Biology: Holarctic and Neotropical (French Guiana and Argentina). The nymphs of this genus appear to prefer backwater areas of streams or

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Plate 70. Brachycercus sp., nymph. Figs. 70A-E.70A, head; 70B, scape and pedicel of antenna, detail; 70C, middle leg; 70D, posterolateral spines of abdominal segments, d.v.; 70E, same, l.v. Figs. 70A-E modified from SOLDAN, 1986.

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rivers, with sandy bottoms, in which they are found buried or covered with silt. They have also been collected from lake shores. Nevertheless, there are no data about the biology of the South American species. Brachycercus tenella (NAVÁS) Nomen Dubium Eurycaenis tenella NAVÁS, 1932a: 112 (male, female). Brachycercus tenella; THEW, 1960a: 202; HUBBARD, 1982a: 261. Brachycercus tenellus; SOLDÁN, 1986: 294.

Taxonomy: Brachycercus tenella was described from imagos of both sexes. Unfortunately, the types have been lost and the original description contains no morphological characters, which would permit us to identify this species. Serious doubts exist about its reference to the genus Brachycercus. We consider the species a nomen dubium. Distribution: Argentina (Buenos Aires prov.). Genus Brasilocaenis PUTHZ Brasilocaenis PUTHZ, 1975: 411; MALZACHER, 1986: 84; MALZACHER, 1998: 3. (Type-species: Brasilocaenis irmleri PUTHZ, original designation).

Taxonomy: Brasilocaenis was established for 1 species, B. irmleri, which is known from imagos of both sexes and nymphal exuviae. Later, MALZACHER (1986, 1990, 1998) revised the genus and described another five species. MALZACHER (1998) divided the genus in 2 species groups: the irmleri-group (with B. irmleri, B. puthzi, and B. renata) and the septentrionalis-group (with B. septentrionalis and B. mendesi). This author also stated that the identity of B. intermedia, known only from nymphs, is doubtful. MALZACHER indicated the possibility that Brasilocaenis was, in reality, part of Caenis, making the latter a paraphyletic group, because the imagos are differentiated from the other caenid genera only by characters of the male genitalia, but some of these species are similar to Caenis. The most important characters for their separation are found in the nymphs. Adult Characteristics: 1) Pedicel of antennae twice as long as scape; 2) prosternum longer than wide, with anterior coxae contiguous; and 3) styliger plate posteriorly elongated often covering penes (Fig. 71A, D-G). Nymphal Characteristics: 1) Pedicel of antennae twice as long as scape; 2) ocellar tubercles absent; 3) maxillary and labial palpi 3-segmented; 4) prosternum longer than wide, with anterior coxae contiguous; 5) dorsolateral spines present on middle and hind coxae (Figs. 71I, K, M); 6) dorsal surface of operculate gills on abdominal segment II covered with short strong and blunt spines; 7) abdominal segments with long sharp posterolateral projections; and 8) anterolateral margins of pronotum generally forming a characteristic angle (Figs. 71H, J, L); 9) lateral margins of abdominal segments VII-IX besides the bristles with strong teeth. Distribution: Brazil and Colombia.

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KEYS

TO

BRASILOCAENIS

Adult Males 1 – 2(1) – 3(1) – 4(3) –

Basolateral sclerites parallel, laying close to the lateral sclerites; penes with laterally protruding lobes, structured median part visible (Figs. 71F-G) ................................ ............................................................................................................ septentrionalis-group 2 Basolateral sclerites diverging, laying apart from the lateral sclerites; penes not as above (Figs. 71A, D-E) ............................................................................. irmleri-group 3 Styliger plate with tongue-shaped posterolateral processes (TP in Fig. 71F) ......... ................................................................................................................. Brasilocaenis mendesi Styliger plate not as above (Fig. 71G) ................................... Brasilocaenis septentrionalis Apex of styliger plate broadly truncated; apex of forceps straight or slightly curved (Fig. 71E) ................................................................................... Brasilocaenis renata Apex of styliger plate rounded, apex of forceps strongly curved (Figs. 71A, D) ...... 4 Apex of forceps strongly curved toward median line (Fig. 71A); flagellum of antennae not expanded basally (Fig. 71B) ...................................... Brasilocaenis irmleri Apex of forceps recurved and parallel to the median line (Fig. 71D); flagellum of antennae swollen basally (Fig. 71C) .................................................. Brasilocaenis puthzi

Nymphs11 1 – 2(1) –

CLAVES

Anterolateral margins of the pronotum markedly angular (Fig. 71J); coxal processes on mid and posterior legs long and apically acute (Fig. 71K) ................. .................................................................................................................. Brasilocaenis irmleri Anterolateral margins of pronotum not as above (Fig. 71H, L); coxal processes on mid and posterior legs short and apically rounded (Figs. 71I, M) .......................... 2 Coxal processes with parallel sides and rounded apically (Fig. 71I); anterior margin of pronotum angular and directed anteriorly (Fig. 71H) ......... Brasilocaenis intermedia Coxal processes short, tapered, and rounded apically (Fig. 71M); anterolateral margins of pronotum blunt and directed laterally (Fig. 71L) ....... Brasilocaenis puthzi PARA

BRASILOCAENIS

Machos Adultos 1 – 2(1) –

11

Escleritos basolaterales paralelos, cercanos a los escleritos laterales; penes con lóbulos laterales protruyentes y zona media rugosa visible (Figs. 71F-G) .............. ............................................................................................................ grupo-septentrionalis 2 Escleritos basolaterales divergentes, alejados de los escleritos laterales; penes no como arriba (Figs. 71A, D-E) .................................................................. grupo-irmleri 3 Placa estilígera con procesos posterolaterales en forma de lengua (TP en Fig. 71F) ................................................................................................................. Brasilocaenis mendesi Placa estilígera no proyectada como en el caso anterior (Fig. 71G) ......................... ....................................................................................................... Brasilocaenis septentrionalis

Brasilocaenis renata is not included in the key because it is known only from a single nymphal exuviae.

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Plate 71. Brasilocaenis, male imagos. Figs. 71A-M. B. irmleri: 71A, male genitalia, v.v.; 71B, antennae. B. puthzi: 71C, antenna; 71D, male genitalia. B. renata: 71E, male genitalia. B. mendesi i: 71F, male genitalia. B. septentrionalis: 71G, male genitalia. Nymphs: B. intermedia: 71H, anterolateral margins of pronotum; 71I, mesocoxal process. B. irmleri: 71J, anterolateral margins of pronotum; 71K, mesocoxal process. B. puthzi: 71L, anterolateral margins of pronotum; 71M, mesocoxal process (F = forceps; FL = flagellum; P = penes; SP = Styliger plate; TP = tongue processes). Figs. modified from MALZACHER, 1986, except 71F from MALZACHER, 1998 and 71G from MALZACHER, 1990.

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Ápice de la placa estilígera ampliamente truncada; ápice de los fórceps rectos o suavemente curvados (Fig. 71E) ........................................................ Brasilocaenis renata Ápice de la placa estilígera redondeada; ápice de los fórceps fuertemente curvados (Figs. 71A, D) ...................................................................................................................... 4 Ápice de los fórceps fuertemente curvados hacia la línea media (Fig. 71A); flagelo de las antenas no ensanchado (Fig. 71B) ........................................ Brasilocaenis irmleri Ápice de los fórceps curvados de manera que terminan paralelos a la línea media (Fig. 71D); flagelo de las antenas hinchado (Fig. 71C) ................ Brasilocaenis puthzi

Ninfas12 1 – 2(1) –

Margen anterolateral del pronoto terminando en punta (Fig. 71J); procesos coxales en patas medias y posteriores puntiagudos y largos (Fig. 71K) ................... .................................................................................................................. Brasilocaenis irmleri Margen anterolateral del pronoto no terminando en punta (Figs. 71H, L); procesos coxales en patas medias y posteriores más cortos y redondeados (Figs. 71I, M) .................................................................................................................................. 2 Espinas coxales con lados paralelos (Fig. 71I) y redondeada apicalmente; margen anterolateral del pronoto dirigidos hacia adelante (Fig. 71H) ....... Brasilocaenis intermedia Espinas coxales cortas, triangulares y redondeadas en el extremo (Fig. 71M); margen anterolateral del pronoto dirigido lateralmente (Fig. 71L) ............. Brasilocaenis puthzi

Brasilocaenis intermedia MALZACHER Brasilocaenis intermedia MALZACHER, 1986: 89 (nymph); MALZACHER, 1998: 1.

Taxonomy: Brasilocaenis intermedia is known only from nymphal exuviae. The assignment of this species to Brasilocaenis must be confirmed when the male imagos are known because the nymph has some characters that do not correspond to the generic description. It can be characterized by the following combination of characters: 1) Length of body of male 2.8-3.0 mm, of female 3.4-4.0 mm; 2) anterolateral margins of pronotum rounded and directed anteriorly (Fig. 71H); 3) coxal processes of middle and hind legs with parallel sides and rounded apically, with small evenly spaced denticles (Fig. 71I); and 4) posterolateral projections on abdominal segments V-VIII long and pointed. Distribution: Brazil (Tocantins river). Brasilocaenis irmleri PUTHZ Brasilocaenis irmleri PUTHZ, 1975: 412 (male, female, nymph); IRMLER, 1975: 348; MALZACHER, 1986: 85 (male, female, nymph, egg); MALZACHER, 1998: 3.

Taxonomy: Brasilocaenis irmleri, the type-species of the genus, was described from imagos of both sexes and nymphs. It can be characterized by the following combination of characters. In the imagos: 1) Body length of male 3.1-3.5 mm, of female 3.5-4.6 mm; 2) base of antennal flagellum slender (Fig. 71B); 3) male genitalia as in Fig. 71A, with 12

No se incluye B. renata por ser conocida de una única exuvia ninfal.

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forceps enlarged near base and narrow toward apex; apex of forceps curved inwardly; and 4) meso- and metanota light brown, abdomen yellowish with darker spots on terga II-III and VII. In the nymph: 1) Body length of male 2.9-3.2 mm, of female 3.6-4.2 mm; 2) anterolateral margins of pronotum forming a very marked angle (Fig. 71J); 3) coxal processes of middle and hind legs long, sharply tapered, and with dentate borders (Fig. 71K); and 4) posterolateral projections of abdominal segment V-VIII long and sharp, protruding more than in other species; 5) hind margin of the ninth sternite rounded. Distribution: Brazil (Manaus, Rivers Negro, Amazonas, Solimões, Madeira, TapajosCururu, Paru, Belem); Colombia (Leticia). Brasilocaenis mendesi MALZACHER Brasilocaenis mendesi MALZACHER, 1998: 2 (male).

Taxonomy: Brasilocaenis mendesi was described from male imagos collected in Brazil (Pantanal), recently we collected material from Colombia (Leticia). This species can be separated from the other species of the genus by the following combination of characters: 1) Body length 2.6-3.3 mm; 2) base of antennal flagellum hardly swollen (similar to Fig. 71B); 3) genitalia as in Fig. 71F, with forceps fused to lateral margins of styliger plate, except for apical part, and styliger plate with posterolateral tongue-like processes; and 4) meso- and metanota light-yellowish, with rest of body whitish, with some areas intensively shaded with blackish brown. Distribution: Brazil (Pantanal), Colombia (Leticia). Brasilocaenis puthzi MALZACHER Brasilocaenis puthzi MALZACHER, 1986: 86 (male, female, nymph, egg); MALZACHER, 1998: 3.

Taxonomy: Brasilocaenis puthzi was described from imagos of both sexes, nymphs, and eggs. This species can be characterized by the following combination of characters. In the imagos: 1) Body length of male 2.5-3.0 mm, of female 2.7-3.5 mm; 2) base of antennal flagellum swollen (Fig. 71C); 3) male genitalia with forceps markedly narrowed only in apical third and curved as in Fig. 71D; and 4) coloration similar to B. irmleri, although somewhat lighter. In the nymph: 1) Body length of male 2.6-3.0 mm, of female 3.1-3.8 mm; 2) anterolateral margins of pronotum rounded (Fig. 71L); 3) coxal processes of middle and hind legs short, apically tapered, and gently arched (Fig. 71M); and 4) posterolateral projections of abdominal segments scarcely evident; 5) hind margin of the ninth sternite truncated. Distribution: Brazil (regions of Rivers Tapajos and Tocantins, Negro, Itu, Madeira). Mostly in creeks and small streams. Brasilocaenis renata MALZACHER Brasilocaenis renata MALZACHER, 1986: 88 (male, female, nymph, egg); MALZACHER, 1998: 3.

Taxonomy: Brasilocaenis renata was described from male subimagos, a nymphal exuviae and females from the same locality tentatively assigned to this species, and can be characterized by the following combination of characters. In the adults: 1) Body length of

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male 2.8 mm, of female 3.7-4.0 mm; 2) flagellum apparently normal (not mentioned in original description); 3) genitalia as in Fig. 71E with forceps smoothly curved on external margin and pointed apically; and 4) head and pronotum yellowish, and meso- and metanota and pleura yellowish brown, strong dark maculae on abdominal terga I-II and VII-VIII, on these last segments maculae emarginated laterally, delimiting oval or semicircular light areas. The nymph is known only from 1 exuviae and is very similar to B. irmleri. It shows a rounded process apically on abdominal tergum II and another very large process on tergum I. Distribution: Brazil (Rio Tarumã). Brasilocaenis septentrionalis MALZACHER Brasilocaenis septentrionalis MALZACHER, 1990: 37 (male); MALZACHER, 1998: 3.

Taxonomy: Brasilocaenis septentrionalis was described from a single male which can be characterized by the following combination of characters: 1) Body length 3.0 mm; 2) base of antennal flagellum clearly enlarged, but not swollen; 3) genitalia as in Fig. 71G, with forceps fused to lateral margins of styliger plate, except for apical part, and apex curved toward median line; and 4) meso- and metanota yellowish brown, with rest of body whitish yellow. Distribution: Brazil (Pará: Rio Paru). Genus Caenis STEPHENS Caenis STEPHENS, 1835: 60; MALZACHER, 1986: 90. (Type-species: Caenis macrura STEPHENS, subsequent designation by WESTWOOD, 1840).

Taxonomy: Caenis was established in 1835 for a species from England. This genus is well represented in South America, although the species described up to now for the region surely represent only a small portion of those existing. At present, the genus is being revised by P. MALZACHER. He indicates the possibility that Brasilocaenis is in reality part of Caenis, which would make the genus paraphyletic. Twenty two species of Caenis have been described from South America, of which we consider 4 as nomina dubia. Of the remaining eighteen, 8 are known from nymphs. MALZACHER (1986, 1990) recognized 2 groups of species in South America based on the genitalia: the Caenis fittkaui group (forceps strong, sclerotized, and apically pointed) and the Caenis reissi group (forceps short, weaker, apically rounded and surface covered with setae). This last group was further subdivided (MALZACHER, 2001) into 4 species groups from Central and South America (and a fifth group from West Africa). Adults Characteristics: 1) Pedicel of antennae twice as long as scape (Fig. 72E); 2) with longitudinal ridges forming a triangular or trapezoid shaped structure (Figs. 72F, J, P); 3) styliger plate not very elongated caudally (see Figs. 72D, I), and forceps not fusing with it. In this genus, most characters are those of the male genitalia. Details of the styliger plate and its connection with sternum IX include the “styliger sclerite” referring to the sclerotized posterior region, the apophyses of this region (AS), the basal “central sclerite” (CS), and the chitinous and epidermal pigments (CP and EP) (Figs. 72D. 73A, 73N). Nymphal Characteristics: 1) Pedicel of antennae twice as long as scape; 2) ocellar tubercles absent (Fig. 75A); 3) maxillary and labial palpi 3-segmented; 4) prosternum longer

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Plate 72. Caenis, male imagos. Figs. 72A-T. Caenis sp.: 72A, fore wing; 72B, mesonotum. C. argentina: 72C, outline of the lateral part of the abdomen, d.v.; 72D, genitalia; 72E, antennae; 72F, prosternal triangle; 72G-72H, details of forceps. C. burmeisteri: 72I, genitalia; 72J, prosternal triangle; 72K, detail of forceps. C. candelata: 72L, genitalia; 72M, detail of forceps. C. chamie: 72N, genitalia; 72O, detail of forceps; 72P, prosternal triangle. C. cuniana: 72Q, genitalia; 72R, detail of forceps; 72S, prosternal triangle. C. cigana: 72T, genitalia. (AS = apophyses of styliger sclerite; CS = central sclerite; EP = epidermal pigments; F = forceps; OM = ommation; P = penes). Figs. 72C-H modified from MALZACHER, 2001; 72I-K from MALZACHER, 1990; 72L-M from MALZACHER, 1986, 72N-P from ALBA-TERCEDOR & MOSQUERA, 1999; 72Q-S from DA-SILVA, 1993a and 72T from PEREIRA & DA-SILVA, 1990.

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than wide, with anterior coxae contiguous; 5) fore legs almost as long as middle and hind legs, and tarsal claws short; 6) dorsal face of gills of abdominal segment II not covered with spines; and 7) posterolateral projections on median abdominal segments (Fig. 72C) not upcurved; 8) gill of abdominal segment II with a submarginal row of microtrichiae on the ventral side. Distribution and Biology: Cosmopolitan, except New Zealand and oceanic islands. In South America it is widely distributed as far as Patagonia and Central Chile. Biological data exist for some species: C. cigana and C. cuniana of Brazil, C. chamie from Colombia, and an unidentified Caenis species from southern Argentina. The nymphs can be found in ponds or rivers, and are not good swimmers; generally they are found in vegetation or among organic material deposited on the bottom in areas of low current, although at times they can be collected in zones of rapid current. In some cases they can be found on sandy bottoms. The operculate gills protect the respiratory surfaces of the other gills. The nymphs are collectorsgatherers, and the stomach contents are composed of vegetable fibers and organic detritus. They tolerate wide variations of water temperature and oxygen levels, and some degree of contamination. Nymphal development varies greatly, according to the area and time of year, the shortest developmental time recorded being about 1 month. The time of emergence varies from midday until a little after dusk. In South America, the nuptial flight takes place around midday or later in the afternoon, though at times males are collected late in the morning among very numerous swarms of Tricorythodes. The flight consists of rapid vertical movements, between 1.5 and 3 m above the water surface, although in general the swarms are not as numerous as those of Tricorythodes. The subimagos, which emerge toward the evening, are attracted to light in large numbers, molting to imagos in a few minutes to a few hours. KEYS

TO

SOUTH AMERICAN CAENIS

Adult Males 1 – 2(1) – 3(2) – 4(2) –

Apex of the forceps sharp and sclerotized (as in Figs. 72I, Q) ............................... 2 Apex of the forceps rounded (as in Figs. 72D, N) ..................................................... 6 Lobes of the penes forming blunt angles laterally, without tubercles on the ventral surface of the penes (Figs. 72I, 74L) ............................................................................. 3 Lobes of the penes rounded laterally, small tubercles on the ventral surface of the penes (Figs. 72L, Q, 73E) ................................................................................................. 4 Prosternal triangle with each side of equal length, apex closed and sharp (Fig. 72J); apex of the forceps sharpening abruptly, basal half with numerous setae (Figs. 72I, K) ............................................................................................ Caenis burmeisteri Prosternal triangle short and broadly rounded apically (Fig. 74M); apex of the forceps narrowing gradually toward the apex, without setae on the basal part (Fig. 74N) ....... .................................................................................................................... Caenis pseudamica Forceps narrowing gradually toward the apex, with 3 subapical denticles, surface of the forceps without setae (Fig. 72R); lobes of the penes fused, without vestige of a dividing groove (Fig. 72Q) ................................................................ Caenis cuniana Apex of the forceps sharpening abruptly, some setae present on the surface of the forceps (Figs. 72M, 73F); shallow groove on the ventral surface of the penes (Figs. 72L, 73E) .................................................................................................................. 5

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Forceps approximately 6 times longer than width of median part, about 6 setae present on their surface (Fig. 72M) ........................................................ Caenis candelata – Forceps approximately 9 to 11 times longer than width of median part, approximately 15 to 20 setae present on their surface (Fig. 73F)..................................... Caenis fittkaui 6(1) Finger-like median process present in abdominal tergum II (Figs. 73I, M); fore leg twice as long as hind leg or longer; forceps and sclerites of the genitalia strongly colored .................................................................................................................................. 7 – Abdominal tergum II without median projections (but lateral projections may be present); fore leg at most 1.8 times the length of hind leg; forceps and sclerites of the genitalia nearly uncolored ........................................................................................ 11 7(6) Lateral filaments of abdominal segments relatively short; lobes of penes short and laterally rounded (Fig. 74A); apophyses of the styliger sclerite long and thin, somewhat distanced from the base of the forceps (Fig. 74A) .... Caenis plaumanni – Lateral filaments of abdominal segments medium to long (Fig. 72C); lobes of penes longer and generally laterally acute (Figs. 72D, 73A, G, N); apophyses of the styliger sclerite short and close to base of forceps (Figs. 72D, 73A, G, N) .......... 8 8(7) Finger-like process of abdominal tergum II relatively short (Fig. 73I); apex of forceps with small projections (Fig. 73H) ................................................ Caenis gonseri – Finger-like process of abdominal tergum II long (Fig. 73M); apex of forceps smoothly rounded (Figs. 72G-H, 73B-C, 73O-P) ....................................................... 9 9(8) Lobes of penes laterally pointed; apophyses of styliger sclerite bent inwards (Fig. 73A); dilated base of the antennal bristle short and more or less triangular; basal half of the forceps with trichomae with arrowhead-like dilated tips ........ Caenis dominguezi – Lobes of penes ovoid to rounded; apophyses of styliger sclerite straight (Figs. 72D, 73N); base of the antennal bristle hardly dilated; trichomae of the forceps not with arrowhead-like dilated tips ............................................................................. 10 10(9) Central sclerite of styliger plate elongate (Fig. 72D); epidermal pigmentation reduced on basal part of styliger (Fig. 72D); a brownish species ... Caenis argentina – Central sclerite circular (Fig. 73N); styliger plate heavily pigmented (Fig. 73N); a very dark blackish-brown species ...............................................................Caenis ludicra 11(6) Lobes of the penes totally fused, without vestige of a groove dividing them (Fig. 72T) .............................................................................................................................Caenis cigana – Groove separating the lobes of the penes visible (Figs. 74E, H) .......................... 12 12(11) Penes with clearly laterally protruding lobes (Figs. 72N, 73J, Q) .......................... 13 – Lobes of penes very short and not or only a little bit protruding (Figs. 74E, H, O) .............................................................................................................................................. 15 13(12) Posterior margin of styliger plate straight, fore margin of the styliger sclerite convex (Fig. 73J). .............................................................................................. Caenis grimi – Posterior margin of styliger plate more or less triangular (Figs. 72N, 73Q); fore margin of the styliger sclerite concave or straight (Figs. 72N, 73Q). ................... 14 14(13) Apical margin of penes convex; apophyses of the styliger sclerite bent inwardly (Fig. 73Q) ........................................................................................................... Caenis pflugfelderi – Apical margin of penes straight; apophyses of the styliger sclerite pointed and straight (Fig. 72N) .......................................................................................... Caenis chamie

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15(12) Lobes of the penes very short and swollen at base (Fig. 74O); styliger plate usually covering the penes; prosternal triangle narrow and triangular, usually closed toward apex (Fig. 74R) ................................................................................ Caenis sigillata – Lobes of the penes relatively long (Figs. 72T, 74E, H); styliger plate not covering penes; prosternal triangle broad and usually open toward apex (Figs. 74F, K) ......... 16 16(15) Prosternal triangle very broad (Fig. 74F); apex of genital forceps rounded with margins ornamented with small wart-like bumps (Fig. 74G); basal dilation of the antennal bristle about 1.5 times the length of the pedicel .......... Caenis quatipuruica – Prosternal triangle not as above (Fig. 74K); apex of forceps without ornamented margins (Figs. 74I-J); basal dilation of the antennal bristle shorter than the length of the pedicel .................................................................................................... Caenis reissi Nymphs13 1 – 2(1) 3(2) – 4(3) – 5(4) – 6(2) –

CLAVES

Denticles on fore and middle tarsal claws absent or small (as in Figs. 75B-C) .... 2 Denticles on fore and middle tarsal claws large (Figs. 75E-F) .......... Caenis cuniana Denticles on fore and middle tarsal claws present, small (as in Figs. 75B-C) ...... 3 Denticles on fore and middle tarsal claws absent ....................................................... 6 Coxal processes on middle and hind legs short, semicircular (as in Fig. 76F) ..... 4 Coxal processes on middle and hind legs long and tapered apically (Fig. 76G) .... ............................................................................................................................Caenis ludicra Dorsal surface of body and operculate gills with semicircular scale-shaped microtrichia; lateral margins of abdominal segments VIII-IX with small teeth .... ..................................................................................................................... Caenis plaumanni Dorsal surface of body and operculate gills without semicircular scale-shaped microtrichia; lateral margins of abdominal segments VIII-IX smooth ................. 5 Pronotum short, not projected anteriorly (Fig. 76A), tarsal claws as in Figs. 75BD; posteromedian process on the hind margin of abdominal tergum II long and apically rounded ......................................................................................... Caenis argentina Pronotum longer, projected anteriorly (Fig. 76B), tarsal claws as in Figs. 75H-J; posteromedian process on the hind margin of abdominal tergum II short and broad triangular, apically pointed ............................................................. Caenis fittkaui Coxal processes on middle and hind legs wide and rounded (Fig. 76D); apex of ninth sternum with median emargination (Fig. 76J) ............................... Caenis chamie Coxal processes on middle and hind legs narrow (Fig. 76H); apex of ninth sternum rounded (Fig. 76L) .................................................................. Caenis pflugfelderi PARA LOS

CAENIS

SUDAMERICANOS

Adultos Machos 1 –

13

Ápice de los fórceps agudos y esclerotizados (como en Figs. 72I, Q) ................... 2 Ápice de los fórceps redondeado (como en Figs. 72D, N) ...................................... 6

Caenis cigana PEREIRA & DA-SILVA (1990b) was not included in the key due to lack of data.

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–

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Lóbulos de los penes formando ángulos romos lateralmente, sin tubérculos sobre la superficie ventral de los penes (Figs. 72I, 74L) ....................................................... 3 Lóbulos de los penes redondeados lateralmente, pequeños tubérculos sobre la superficie ventral de los penes (Figs. 72L, Q, 73E) .................................................... 4 Triángulo prosternal con cada lado de igual longitud, ápice cerrado y agudo (Fig. 72J); ápice de los fórceps aguzados abruptamente, 1/2 basal con numerosas setas (Figs. 72I, K) ............................................................................................ Caenis burmeisteri Triángulo prosternal corto y ampliamente redondeado apicalmente (Fig. 74M); ápice de los fórceps afinándose gradualmente hacia el ápice, sin setas en la parte basal (Fig. 74N) ...................................................................................... Caenis pseudamica Fórceps afinándose gradualmente hacia el ápice, con 3 dentículos subapicales, superficie de los fórceps sin setas (Fig. 72R); lóbulos de los penes fusionados, sin vestigio de surco que los divida (Fig. 72Q) ............................................ Caenis cuniana Apice de los fórceps aguzados abruptamente, algunas setas presentes en la superficie de los fórceps (Figs. 72M, 73F); surco poco profundo en la cara ventral de los penes (Figs. 72L, 73E) .......................................................................................... 5 Fórceps aproximadamente 6 veces más largos que el ancho en la parte media, alrededor de 6 setas presentes en su superficie (Fig. 72M) ............... Caenis candelata Fórceps aproximadamente 9 a 11 veces más largos que el ancho en la parte media, aproximadamente 15 a 20 setas presentes en su superficie (Fig. 73F) .... .......................................................................................................................... Caenis fittkaui Tergo abdominal II con una proyección media dorsal en forma de dedo (Figs. 73I, M); pata anterior 2 veces la longitud de la pata posterior o más larga; fórceps y escleritos de la genitalia fuertemente coloreados ........................................................ 7 Tergo abdominal II sin proyección media dorsal (puede haber proyecciones laterales); pata anterior como máximo 1,8 veces más larga que la pata posterior; fórceps y escleritos de la genitalia casi descoloridos ................................................ 11 Filamentos laterales de los segmentos abdominales relativamente cortos; lóbulos de los penes cortos y redondeados lateralmente (Fig. 74A); apófisis del esclerito estilígero largas y delgadas, algo distanciadas de la base de los fórceps (Fig. 74A) ..................................................................................................................... Caenis plaumanni Filamentos laterales de los segmentos abdominales medianos a largos (Fig. 72C); lóbulos de los penes más largos y generalmente lateralmente aguzados (Figs. 72D, 73A, G, N); apófisis del esclerito estilígero cortas y cercanas a la base de los fórceps (Figs. 72D, 73A, G, N) ....................................................................................... 8 Proyección media dorsal en forma de dedo del tergo abdominal II relativamente corta (Fig. 73I); ápice de los fórceps con pequeñas poyecciones (Fig. 73H) .......... ............................................................................................................................ Caenis gonseri Proyección media dorsal en forma de dedo del tergo abdominal II larga (Fig. 73M); ápice de los fórceps redondeados y lisos (Figs. 72G-H, 73B-C, 73O-P) .................. 9 Lóbulos de los penes lateralmente aguzados; apófisis del esclerito estilígero curvadas hacia la línea media (Fig. 73A); base dilatada del flagelo antenal corta y más o menos triangular; mitad basal de los forceps con tricomas con extremo en forma de flecha ....................................................................................... Caenis dominguezi

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– 10(9) – 11(6) – 12(11) – 13(12) – 14(13) – 15(12) – 16(15) –

Lóbulos de los penes ovoides a redondeados; apófisis del esclerito estilígero rectas (Figs. 72D, 73N); base del flagelo antenal no dilatada; tricomas de los fórceps no como arriba ....................................................................................................................... 10 Esclerito central de la placa estilígera elongado (Fig. 72D); pigmentación epidérmica reducida a la parte basal de la placa estilígera (Fig. 72D); especie de coloración parda ......................................................................................... Caenis argentina Esclerito central circular (Fig. 73N); placa estilígera fuertemente pigmentada (Fig. 73N); especie muy oscura, pardo negruzca .............................................. Caenis ludicra Lóbulos de los penes totalmente fusionados, sin vestigios de un surco dividiéndolos (Fig. 72T) .................................................................................Caenis cigana Surco visible separando los lóbulos peneanos (Figs. 74E, H) ................................ 12 Penes con lóbulos laterales claramente protruyentes (Figs. 72N, 73J, Q) ........... 13 Lóbulos de los penes muy cortos, apenas o no proyectados lateralmente (Figs. 74E, H, O) ......................................................................................................................... 15 Márgen posterior de la placa estilígera recto, márgen anterior del esclerito estilígero convexo (Fig. 73J) ........................................................................... Caenis grimi Márgen posterior de la placa estilígera más o menos triangular (Figs. 72N, 73Q); márgen anterior del esclerito estilígero cóncavo o recto (Figs. 72N, 73Q) ......... 14 Márgen apical de los penes convexo; apófisis del esclerito estilígero curvadas hacia la línea media (Fig. 73Q) ....................................................................... Caenis pflugfelderi Márgen apical de los penes recto; apófisis del esclerito estilígero aguzadas y rectas (Fig. 72N) ......................................................................................................... Caenis chamie Lóbulos de los penes muy cortos e hinchados en la base (Fig. 74O); placa estilígera usualmente cubriendo los penes; triángulo prosternal angosto y triangular, usualmente cerrado hacia el ápice (Fig. 74R) ......................................... Caenis sigillata Lóbulos de los penes relativamente largos (Figs. 72T, 74E, H); la placa estilígera no cubre los penes; triángulo prosternal ancho y usualmente abierto hacia el ápice (Figs. 74F, K) ..................................................................................................................... 16 Triángulo prosternal muy ancho (Fig. 74F); ápice de los fórceps redondeados con pequeñas protuberancias en forma de verrugas (Fig. 74G); ensanchamiento basal del flagelo antenal alrededor de 1,5 la longitud del pedicelo ................. Caenis quatipuruica Triángulo prosternal no como arriba (Fig. 74K); ápice de los fórceps sin ornamentación en los márgenes (Figs. 74I-J); ensanchamiento basal del flagelo antenal más corto que la longitud del pedicelo ......................................... Caenis reissi

Ninfas

14

1

Dentículos de las uñas tarsales anteriores y medias ausentes o pequeños (como en Figs. 75B-C) ......................................................................................................................... 2 Dentículos de las uñas tarsales anteriores y medias grandes (Figs. 75E-F) ............. .......................................................................................................................... Caenis cuniana Dentículos de las uñas tarsales anteriores y medias presentes, pequeños (como en Figs. 75B-C) ......................................................................................................................... 3

– 2(1)

14

Caenis cigana PEREIRA & DA-SILVA (1990b) no fue incluida en la clave debido a falta de datos.

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B

E

209

F

C

CP

AS

D J

G

K

H

L I Q M O

R

N

P

T EP

U

S

CS

Plate 73. Caenis, male imagos. Figs. 73A-U. C. dominguezi: 73A, genitalia; 73B-C, details of forceps; 73D, prosternal triangle. C. fittkaui: 73E, genitalia; 73F, detail of forceps. C. gonseri: 73G, genitalia; 73H, detail of forceps; 73I, finger-like process of abdominal tergum II, l.v. C. grimi: 73J, genitalia; 73K-L, detail of forceps. C. ludicra: 73M, finger-like process of abdominal tergum II, l.v.; 73N, genitalia; 73O-P, detail of forceps. C. pflugfelderi: 73Q, genitalia; 73R-S, details of forceps; 73T-U, variations in prosternal triangle. (AS =apophyses of styliger sclerite; CP = chitinous pigments; CS = central sclerite; EP = epidermal pigments). Figs. 73A-D, G-P modified from MALZACHER, 2001; 73E-F from MALZACHER, 1986 and 73Q-U from MALZACHER, 1990.

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Dentículos de las uñas tarsales anteriores y medias ausentes ................................... 6 Procesos coxales de las patas medias y posteriores cortos, semicirculares (as in Fig. 76F) ............................................................................................................................... 4 Procesos coxales de las patas medias y posteriores largos y aguzados apicalmente (Fig. 76G) .........................................................................................................Caenis ludicra Superficie dorsal del cuerpo y branquias operculares con microtriquios semicirculares en forma de escama; márgenes laterales de los segmentos abdominales VIII-IX con pequeños dentículos ............................... Caenis plaumanni Superficie dorsal del cuerpo y branquias operculares sin microtriquios semicirculares en forma de escama; márgenes laterales de los segmentos abdominales VIII-IX lisos ................................................................................................ 5 Pronoto corto, no proyectado anteriormente (Fig. 76A), uñas tarsales como en las Figs. 75B-D; proceso posteromediano en el margen posterior del tergo abdominal II largo y redondeado apicalmente ........................................................ Caenis argentina Pronoto más largo, proyectado anteriormente (Fig. 76B), uñas tarsales como en las Figs. 75H-J; proceso posteromediano en el margen posterior del tergo abdominal II corto y ancho, triangular, aguzado apicalmente ............ Caenis fittkaui Procesos coxales de las patas medias y posteriores, anchos y redondeados (Fig. 76D); ápice del noveno esternito con emarginación mediana (Fig. 76J) .............. ............................................................................................................................ Caenis chamie Procesos coxales de las patas medias y posteriores más delgados (Fig. 76H); ápice del noveno esternito redondeado (Fig. 76L) ..................................... Caenis pflugfelderi

Caenis argentina NAVÁS Caenis argentina NAVÁS, 1915b: 12 (male); HUBBARD, 1982a: 261; MALZACHER 2001: 2 (male, female, nymph, egg).

Taxonomy: This species was inadequately described by NAVÁS (1915b) based on a series of specimens (apparently males) caught in a spider web, and in very bad shape. Recently, MALZACHER (2001) redescribed the male imagos and described the females, nymphs and eggs for the first time. The species can be separated from the other species of the genus by the following combination of characters. In the adult: 1) Body length of male 4.0-4.7 mm, of female 5.0-5.5 mm; 2) prosternal triangle equal-sided, pointed, and with slightly concave sides (Fig. 72F); 3) finger-like process on abdominal tergum II long (as in Fig. 73M); 4) lateral filaments of abdominal segments long (Fig. 72C); 5) posterior margin of styliger plate straight (Fig. 72D); 6) central sclerite elongate and elliptical, styliger sclerite with long and straight apophyses, epidermal pigments absent or limited to basal part (Fig. 72D); 7) lobes of penes rounded laterally, with vestiges of a dividing groove (Fig. 72D); 8) forceps rounded apically, surface with 4-5 setae (Figs. 72G-H), inner margin of forceps basally protruding; 9) abdominal terga pale yellowish brown, with clear “paratergal” marks; in general aspect a species with a medium brown color. In the nymph: 1) Body length of male 4.5-5.0 mm, of female 6.0-6.5 mm; 2) lateral margins of pronotum straight (Fig. 76A); 3) coxal processes of middle and hind legs short, semicircular or sickle-shaped; 4) tarsal claws with several small denticles (Figs. 75B-D); 5) sternum IX

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with a blunt apical margin (Fig. 76I); and 6) hind margin of the head with laterally turned long bristles. Distribution: Argentina (Buenos Aires, Tucumán, Salta). Caenis burmeisteri MALZACHER Caenis burmeisteri MALZACHER, 1990: 35 (male).

Taxonomy: This species, known only from male imagos, can be characterized in the following manner: 1) Length of body 3.3-3.5 mm; 2) prosternal triangle pointed (Fig. 72J); 3) finger-like process of abdominal tergum II absent; 4) lateral filaments of abdominal segments short; 5) posterior margin of styliger plate slightly rounded (Fig. 72I); 6) central sclerite elongated, apophyses of styliger sclerite long and bent inward, epidermal pigments present near anterolateral corners of sternum IX (Fig. 72I); 7) apical margin of penes convex (Fig. 72I) and lobes with blunt lateral angles; 8) apex of forceps sharp and sclerotized, base swollen, outer margin slightly concave, numerous setae on basal half (Fig. 72K), and a few setae near apex; 9) abdominal terga diffusely pigmented, denser toward posterior angles and middle of terga I-III; in general aspect a yellowish brown species. Distribution: Paraguay (Río Ypané), Argentina (Formosa). Caenis candelata MALZACHER Caenis candelata MALZACHER, 1986: 93 (male).

Taxonomy: This species, known only from males, can be characterized in the following manner: 1) Length of body 2.6-2.8 mm; 2) finger-like process on abdominal tergum II absent; 3) lateral filaments of abdominal segments apparently short or absent; 4) posterior margin of styliger plate almost straight (Fig. 72L); 5) central sclerite elongated, apophyses of styliger sclerite straight, without marked pigments (Fig. 72L); 6) lobes of penes rounded, with a constriction in the basal 1/4, and small tubercles distributed over ventral surface (Fig. 72L); 7) apex of forceps sharp and sclerotized, positioned almost parallel to median line, surface of forceps with a few setae (approximately 6; Fig. 72M), forceps approximately 6 times longer than median width; 8) coloration of body almost lost because of poor fixation, although apparently paler than C. fittkaui; in general aspect a light yellowish brown species. Distribution: Brazil (Rio Solimões). Caenis chamie ALBA-TERCEDOR & MOSQUERA Caenis chamie ALBA-TERCEDOR & MOSQUERA 1999: 61 (male, female, nymph).

Taxonomy: This species is known from all stages and appears to belong to the pflugfelderi-group of species as defined by MALZACHER (2001) for C. pflugfelderi and C. panamensis. Adults can be characterized as follows: 1) Body length of male 2.9-3.7 mm, of female 3.5-3.7 mm; 2) prosternal triangle with concave sides, and anteriorly broad (Fig. 72P); 3) finger-like process on abdominal tergum II absent; 4) lateral filaments of abdominal segments apparently short or lacking; 5) posterior margin of styliger plate

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I

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R Plate 74. Caenis, male imagos. Figs. 74A-R. C. plaumanni: 74A, genitalia; 74B-C, details of forceps; 74D, prosternal triangle. C. quatipuruica: 74E, genitalia; 74F, prosternal triangle; 74G, detail of forceps. C. reissi: 74H, genitalia; 74I-J, details of forceps; 74K, prosternal triangle. C. pseudamica: 74L, genitalia; 74M, prosternal triangle; 74N, details of forceps. C. sigillata: 74O, genitalia; 74P-Q, details of forceps; 74R, prosternal triangle. (AS = apophyses of styliger sclerite; WB = wart-like bumps). Figs. 74A-D modified from MALZACHER, 2001; 74E-K, O-R from MALZACHER, 1986 and 74L-N from MALZACHER, 1990.

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more or less triangular (Fig. 72N); 6) central sclerite apparently semicircular, apophyses of styliger sclerite pointed and straight, sternum IX apparently without marked pigments; 7) lobes of penes well developed and rounded, a median groove with a pair of small indentations present (Fig. 72N); 8) forceps relatively long with basal and median projections (Fig. 72O); 9) abdominal terga with gray-brown pigments; in general aspect a yellowish brown species. In the nymph: 1) Length of male 3.0-3.5 mm, of female 4.3-5.3; 2) anterior margin of pronotum with an acute sublateral projection; 3) coxal processes of middle and hind legs well developed, semicircular (Fig. 76D); 4) fore tarsal claw with several small denticles, middle and hind claws without denticles; 5) apical margin of sternum IX with a median emargination (Fig. 76J). Distribution and Biology: Colombia (Valle del Cauca). MOSQUERA et al. (2001) reported that nymphs inhabit in the muddy bottom of a stream with abundant detritus. The mean values of some parameters in this stream are: temperature (19.5ºC), pH (7.52), oxygen saturation (64%). In the field subimagos were observed to emerge at early morning (6-7 a.m.), mainly during rainy or cloudy days. The imaginal molt took place after a few minutes (5-6 min) but no swarming flight was observed. Caenis cigana PEREIRA & DA-SILVA Caenis cigana PEREIRA & DA-SILVA, 1990b: 1 (male, female, nymph, egg).

Taxonomy: Caenis cigana was described from imagos of both sexes, nymphs, and eggs. Description and figures are somewhat incomplete and determination of this species should be confirmed with the study of type material. It can be characterized in the following manner. In the imagos: 1) Body length of male 2.6-2.7 mm, of female 3.2-4.8 mm; 2) finger-like process on abdominal tergum II apparently absent; 3) lateral filaments of abdominal segments probably short or absent; 4) posterior margin of styliger plate more or less straight (Fig. 72T); 5) original drawings not showing the characteristics of central sclerite and apophyses of styliger sclerite; 7) lobes of penes totally fused, without vestiges of a dividing groove (Fig. 72T); 8) apex of forceps rounded and somewhat curved toward median line; and 9) abdomen light brown; in general aspect a orangebrown species. In the nymph: 1) Body length 4.8-5.2 mm; 2) tarsal claws with relatively small denticles. The coxal processes and sternum IX were not described. Distribution and Biology: Brazil (Rio de Janeiro). The nymphs were collected in a dammed area of a river with a temperature between 16ºC and 23ºC at an altitude of 200 m. The nymphs were found among organic material deposited on a rocky bottom. The adults emerged throughout the day (PEREIRA & DA-SILVA, 1990b). Caenis cuniana FROEHLICH Caenis sp. DEMOULIN, 1955a: 4 (nymph). Caenis cuniana FROEHLICH, 1969: 104 (female, nymph); HUBBARD, 1982a: 261; DA-SILVA, 1993a: 413 (male). Caenis cuniana?; MALZACHER, 1986: 94 (nymph, egg).

Taxonomy: Caenis cuniana was originally described from nymphs and parthenogenetic females. Later the male imago was described by DA-SILVA (1993a). Caenis cuniana can be

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characterized in the following manner. In the imagos: 1) Body length of male 1.9 mm, of female 2.5-3.8 mm; 2) prosternal triangle narrowly tapered and open anteriorly (Fig. 72S); 3) finger-like process on abdominal tergum II absent; 4) lateral filaments of abdominal segments absent on segments I-VII, short on VIII-IX; 5) posterior margin of styliger plate slightly concave (Fig. 72Q); 6) central sclerite not illustrated, apophyses of styliger sclerite pointed and bent outward (Fig. 72Q), sternum IX apparently without marked pigments; 7) lobes of penes fused, club shaped, without vestiges of a dividing groove, and tubercles present on ventral surface (Fig. 72Q); 8) forceps sharp and sclerotized apically, somewhat curved toward median line and with 3 subapical denticles, surface without setae (Fig. 72R); and 9) abdominal terga yellowish brown, with dark brown stigmatic maculae; in general aspect an orange-brown species. In the nymph: 1) Body length 3.8-4.8 mm; 2) lateral margins of pronotum almost parallel, a little broader anteriorly (similar to C. fittkaui, Fig. 76B); 3) coxal processes of middle and hind legs narrow and sickle-shaped with margins smooth (Fig. 76E); and 4) tarsal claws of fore and middle legs (Figs. 75E-F)

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Plate 75. Caenis, nymphs. Figs. 75A-P. 75A, Caenis sp. habitus. Figs. 75B-P, Fore, middle and hind tarsal claws: 75B-D, C. argentina; 75E-G, C. cuniana; 75H-J, C. fittkaui; 75K-M, C. ludicra; 75N-P, C. plaumanni. Fig. 75A, drawing by A. Dupuy; Figs. 75B-D, K-P modified from MALZACHER, 2001; 75EJ, from MALZACHER, 1986.

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with denticles which increase in size toward apex, those of hind legs (Fig. 75G) with 3-6 small denticles followed by a row of bifid or multifid setules; and 5) sternum IX rounded. Distribution and Biology: Brazil (Pará, Rio de Janeiro and São Paulo). The nymphs of this species have been collected on the bottom or in vegetation in shallow parts of pools and impoundments, especially where there is rooted vegetation. The nymphs tolerate a wide range of variations in water temperature. This is a species which is principally parthenogenetic (FROEHLICH, 1969), numerous females having been collected, but only 1 male which was recently described (DA-SILVA, 1993a). The biology of this species was studied in detail by FROEHLICH (1969) and later by DA-SILVA (1993a) from whom we have taken the following data. First instar nymphs measure approximately 0.4 mm and the antennae and cerci have five segments, whose number grows incrementally at successive molts. Nymphs of the first three instars do not have gills. The period required for development from oviposition to emergence varies between 50 and 84 days. The nymphs are scrapers and the stomach contents are composed principally of vegetable fibers and organic detritus. The mature nymphs climb or float to the surface. Emergence is rapid, with the wings expanding instantaneously and the subimagos flying immediately. The subimaginal molt occurs between 4-7 minutes after emergence. Emergence takes place in the afternoon or soon after nightfall and can occur any time throughout the year. Soon after the molt, the females are ready for oviposition. The eggs are extruded in a mass which disintegrates on contact with the water, and the eggs fall separately to the bottom. During oviposition, a rupture normally occurs in the intersegmental membrane between sterna VII and VIII where the eggs are extruded followed by the air-filled sac. The normal number of eggs varies between 100 and 300. The time required for eclosion varies in the laboratory between 7 and 29 days. FRANCISCHETTI et al. (2001) reported that nymphs of this species from Rio de Janeiro (Brazil) are scrapers, and that analyses of gut contents revealed the presence of algae, plant fibers and detritus. Relative importance of each item varied seasonally, in relation to water level. Caenis dominguezi MALZACHER Caenis dominguezi MALZACHER, 2001: 9 (male).

Taxonomy: This species was described from male imagos. It can be characterized in the following manner. In the imago: 1) Body length 3.5-3.6 mm; 2) prosternal triangle with concave sides, apically acute or open (Fig. 73D); 3) finger-like process on abdominal tergum II long (as in Fig. 73M); 4) lateral filaments of abdominal segments long (except short on VII -VIII); 5) posterior margin of styliger plate slightly rounded (Fig. 73A); 6) central sclerite diffuse, apophyses of styliger sclerite bent inwards, chitinous lateral structures of sternum IX strongly colored (Fig. 73A); 7) lobes of penes long, rounded or pointed (Fig. 73A); 8) apex of forceps rounded (Figs. 73B-C), and basal half with modified setae with arrowhead-shaped tips; 9) abdominal terga yellowish brown, evenly and sparsely pigmented; in general aspect a species with a medium brown color and a dark brown thorax. Distribution: Argentina (Córdoba).

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Caenis fittkaui MALZACHER Caenis fittkaui MALZACHER, 1986: 91 (male, female, nymph, egg).

Taxonomy: This species was described from imagos of both sexes, nymphs, and eggs. It can be characterized in the following manner. In the imago: 1) Body length of male 2.83.7 mm, of female 3.2-5.3 mm; 2) prosternal triangle not mentioned in original description; 3) finger-like process on abdominal tergum II absent; 4) lateral filaments of abdominal segments apparently short or absent; 5) posterior margin of styliger plate almost straight (Fig. 73E); 6) central sclerite elongate and constricted medially, apophyses of styliger sclerite long, thin and straight, sternum IX apparently without strong pigmentation (Fig. 73E); 7) lobes of penes fused, forming a rounded head, with a constriction in basal 1/4, and small tubercles distributed on ventral surface (Fig. 73E); 8) apex of forceps sharp, sclerotized, and somewhat directed toward median line, surface with approximately 15-20 setae (Fig. 73F), forceps 9-11 times longer than median width; 9) abdominal terga diffusely tinted, segment II with a medial stripe and segment VII with black spots on margins; in general aspect a yellowish brown species. In the nymph: 1) Body length of male 3.2-3.8 mm, of female 3.85.0 mm; 2) lateral margins of pronotum parallel or gently diverging anteriorly, sometimes gently convex (Fig. 76B); 3) coxal processes on middle and hind legs rounded, and margins with small denticles (Fig. 76F); 4) tarsal claws with small denticles (Figs. 75H-J); and 5) sternum IX (Fig. 76N) truncated at posterior apex (sometimes somewhat sinuous). Distribution: Brazil (Belém Rio Tapajós-Cururu-São Manuel, Santarém, Rio Madeira near junction with Rio Amazonas). Caenis gonseri MALZACHER Caenis gonseri MALZACHER, 2001: 11 (male).

Taxonomy: This species was described from male imagos. It can be characterized in the following manner. In the imago: 1) Body length 2.8-3.8 mm; 2) prosternal triangle equilateral with straight or concave sides, rounded anteriorly; 3) finger-like process on abdominal tergum II short (Fig. 73I); 4) lateral filaments of abdominal segments long (except short on VII-VIII); 5) posterior margin of styliger plate slightly rounded (Fig. 73G); 6) central sclerite circular (size variable), styliger sclerite with long apophyses (sometimes bent inwards), sternum IX pigmented basally (Fig. 73G); 7) lobes of penes long, rounded to pointed (Fig. 73G); 8) apex of forceps rounded, forceps curved and broader basally, tip with small projections (Fig. 73H); 9) abdominal terga yellowish brown, evenly pigmented; in general aspect a species with a medium yellowish brown color; and 10) caudolateral processes of the sternum IX apically more or less rounded. Distribution: Argentina (Río Negro); Chile. Caenis grimi MALZACHER Caenis grimi MALZACHER, 2001: 15 (male, female, eggs).

Taxonomy: This species was described from imagos of both sexes and eggs. It can be characterized in the following manner. In the imago: 1) Body length of male 1.9-2.6 mm, of

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female 3.0-3.7 mm; 2) prosternal triangle broad with anteriorly rounded apex and straight or slightly concave sides; 3) finger-like process on abdominal tergum II absent; 4) lateral filaments of abdominal segments very short; 5) posterior margin of styliger plate straight (Fig. 73J); 6) central sclerite diffuse, dome-shaped, styliger sclerite with concave fore margin, apophyses bent inward (Fig. 73J); 7) lobes of penes long and triangular (Fig. 73J); 8) forceps short and tapering with apex rounded (Figs. 73K-L); 9) abdominal terga I-II with narrow transverse bands, others with sublateral blotches, becoming weaker posteriorly; in general aspect a pale species. Distribution: Argentina (Formosa, Misiones, Buenos Aires); Uruguay (Co. Largo, Tacuarembó, Salto). Caenis ludicra NAVÁS Caenis ludiera [sic] NAVÁS, 1920e: 54 (male); ALBA-TERCEDOR & PETERS, 1985: 219. Caenis ludicra; NAVÁS, 1922b: 199; NAVÁS, 1924a: 359; NAVÁS, 1930e: 130; NAVÁS, 1930d: 360; NAVÁS, 1931b: 322; HUBBARD, 1982a: 261; DOMÍNGUEZ, 1989b: 272; MALZACHER, 2001: 6 (male, nymph). Ordella ludicra; LESTAGE, 1931b: 59.

Taxonomy: This species was described from males (NAVÁS, 1920e) and, in subsequent works, NAVÁS corrected the typographical error in the original name (ludiera for ludicra). According to NAVÁS, this species is similar to C. argentina, although shorter (body length 2.6-3.8 mm). Recently MALZACHER (2001) redescribed the male imagos and described the probable nymphs from Brazil, but these nymphs belong to an undescribed species (Molineri, in prep.). The species can be characterized in the following manner. In the imago: 1) Body length of male 2.6-3.8 mm; 2) prosternal triangle anteriorly truncate; 3) finger-like process on abdominal tergum II very long (Fig. 73M); 4) lateral filaments of abdominal segments long (as in Fig. 72C); 5) posterior margin of styliger plate almost straight or slightly rounded (Fig. 73N); 6) central sclerite circular, styliger sclerite with short and rounded apophyses, epidermal pigments distributed over styliger and forceps (Fig. 73N); 7) lobes of penes rounded to very slightly pointed laterally, with vestiges of a dividing groove (Fig. 73N); 8) forceps rounded apically and strongly sclerotized basally, surface with 6-8 setae (Figs. 73O-P); 9) abdominal terga yellowish brown, strongly pigmented with blackish brown; in general aspect a dark colored species. In the nymph: 1) Body length of male 3.8-4.2 mm, of female 4.7-5.4 mm; 2) lateral margins of pronotum straight (Fig. 76A); 3) coxal processes of middle and hind legs long, more or less tapering (Fig. 76G); 4) tarsal claws (Figs. 75B-D) with many small denticles, apex relatively short and less curved than C. argentina; and 5) sternum IX with a median emargination (Fig. 76I). Distribution: Argentina (Córdoba, Santiago del Estero, Tucumán). Caenis nemoralis NAVÁS Caenis nemoralis NAVÁS, 1922c: 386 (female); HUBBARD, 1982a: 261; DOMÍNGUEZ, 1989b: 272. Ordella nemoralis; LESTAGE, 1931b: 60.

Taxonomy: This species was inadequately described based on females. Therefore, until the type material is studied and males are known it is not possible to characterize this species. Distribution: Argentina (Buenos Aires, La Plata).

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Caenis pflugfelderi MALZACHER Caenis pflugfelderi MALZACHER, 1990: 31 (male, female, nymph, egg); ALBA-TERCEDOR & MOSQUERA, 1999: 65; MALZACHER, 2001: 18.

Taxonomy: Caenis pflugflederi was described from imagos of both sexes, nymphs, and eggs. It can be characterized in the following manner. In the imagos: 1) Body length of male 2.4-2.8 mm, of female 3.5 mm; 2) prosternal triangle broadly rounded anteriorly (Figs. 73T-73U); 3) finger-like process on abdominal tergum II absent; 4) lateral filaments of abdominal segments very short; 5) posterior margin of styliger plate more or less an expanded triangle (Fig. 73Q); 6) central sclerite elongate, anterior margin of styliger sclerite concave, apophyses bent inward (Fig. 73Q); 7) penes with oval lobes and a strong ventral transverse ridge (Fig. 73Q); 8) apex of forceps rounded and somewhat curved toward median line, surface densely covered with small setae (Figs. 73R-S); and 9) abdominal terga I-II with prominent transverse bands, terga III-VI with weaker pattern, and VIII-IX with dark spots on lateral areas; in general aspect a yellowish brown species. In the nymph: 1) Body length of male 2.8 mm, of female 4.0 mm; 2) lateral margins of pronotum rounded and expanded anteriorly (Fig. 76C); 3) coxal process on middle and hind legs of medium size, margins strongly dentate (Fig. 76H); 4) tarsal claws of fore and middle legs without denticles, claw of hind legs with a dense row of micro-denticles; and 5) sternum IX apically rounded (Fig. 76L). Distribution: Brazil (Manaus). Caenis plaumanni MALZACHER Caenis plaumanni MALZACHER, 2001: 13 (male, nymph).

Taxonomy: Caenis plaumanni was described from male imagos and nymphs. It can be characterized in the following manner. In the imagos: 1) Body length 2.7-2.8 mm; 2) prosternal triangle equilateral or a little elongated, anteriorly pointed or rounded (Fig. 74D); 3) finger-like process on abdominal tergum II long (as in Fig. 73M); 4) lateral filaments of abdominal segments very short; 5) posterior margin of styliger plate straight (Fig. 74A); 6) central sclerite circular; apophyses of styliger sclerite long and very thin (Fig. 74A); 7) penes with short and rounded lobes (Fig. 74A); 8) apex of forceps rounded and somewhat curved toward the median line (Figs. 74B-C); 9) abdominal terga yellowish, weakly pigmented, and in general aspect a species with medium yellowish brown color. In the nymph: 1) Body length 3.5-5.0; 2) lateral margins of pronotum straight (as in Fig. 76A); 3) coxal process of middle and hind legs semicircular, margin with blunt teeth and very small scale-shaped setae; 5) tarsal claws (Figs. 75N-P) with several small denticles, those on hind claws more numerous and basal teeth stronger than in C. argentina; 6) sternum IX with truncate apex (Figs. 76M-N); 7) body surface with characteristic semicircular scaleshaped and ramified microtrichia; and 8) lateral margin of the abdominal segments VIII-IX with small teeth. Distribution: Brazil (Rio Grande do Sul); Argentina (Misiones); Uruguay (Piraraja, Cebollati).

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Plate 76. Caenis, nymphs. Figs. 76A-N. Pronotum (left side only): 76A, C. argentina; 76B, C. fittkauiC. cuniana; 76C, C. pflugfelderi. Mesocoxal processes: 76D, C. chamie; 76E, C. cuniana; 76F, C. fittkaui; 76G, C. ludicra; 76H, C. pflugfelderi. Sternum IX: 76I, C. argentina; 76J, C. chamie; 76K, C. fittkaui; 76L, C. pflugfelderi; 76M, C. plaumanni, male; 76N, C. plaumanni, female. Figs. 76A, G, I, M-N modified from MALZACHER, 2001; 76B, E-F, H, K-L from MALZACHER, 1986; 76C from MALZACHER, 1990 and 76D, J from ALBA-TERCEDOR & MOSQUERA, 1999.

Caenis pseudamica MALZACHER Caenis pseudamica MALZACHER, 1990: 34 (male).

Taxonomy: This species, known only from male imagos, is very similar to several species from North America, especially C. amica (MALZACHER 1990). It can be characterized in the following manner: 1) Body length 3.4-4.0 mm; 2) prosternal triangle very short and broadly rounded anteriorly, or somewhat emarginated laterally (dotted line, Fig. 74M); 3) finger-like process on abdominal tergum II absent; 4) lateral filaments of abdominal segments of medium length; 5) styliger plate membranous, often difficult to recognize, with more or less rounded posterior margin (Fig. 74L); 6) central sclerite elongate, apophyses of styliger sclerite broad basally and very long, the fore margin of the styliger sclerite is therefore strongly

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concave, sternum IX with black pigment at base (Fig. 74L); 7) penes with long, oval laterally directed, somewhat pointed lobes (Fig. 74L); 8) apex of forceps sharp and sclerotized, anterior third curved toward median line, and with few apical setae (Fig. 74N); and 9) abdominal terga I-V strongly pigmented with black lines on posterior margins, and terga VII-VIII with diffuse triangular spots laterally; in general aspect a yellowish brown species. Distribution: Paraguay (San Bernardino). Caenis quatipuruica MALZACHER Caenis quatipuruica MALZACHER, 1986: 96 (male).

Taxonomy: This species, known only from males, can be characterized in the following manner: 1) Body length 2.3 mm; 2) dilated part of the antennal bristle 1.5 the length of the pedicel; 3) prosternal triangle very broad, opening anteriorly (Fig. 74F); 4) finger-like process on abdominal tergum II absent; 5) lateral filaments of abdominal segments apparently short or absent; 6) posterior margin of styliger plate rounded Fig. 74E); 6) central sclerite more or less circular, outline blurred by surrounding pigments, apophyses of styliger sclerite short and rounded (Fig. 74E); 7) lobes of penes ellipsoidal and clearly separated from basal shaft (Fig. 74E); 8) apex of forceps rounded with margins ornamented with small wart-like bumps (WB in Fig. 74G), surface densely covered with strong setae which project gently outwards (Fig. 74G); and 9) abdomen yellowish brown, abdominal tergum IX brown basally and laterally, styliger plate and central sclerite brown; in general aspect a brownish species. Distribution: Brazil (Belém). Caenis reissi MALZACHER Caenis reissi MALZACHER, 1986: 95 (male, female, egg).

Taxonomy: This species, described from imagos of both sexes, can be characterized in the following manner: 1) Body length of male 2.0-2.4 mm, of female 3.2 mm; 2) antennal bristle with slightly dilated base, dilation short and asymmetric; 3) prosternal triangle usually open anteriorly (Fig. 74K); 4) finger-like process on abdominal tergum II absent; 5) lateral filaments of abdominal segments apparently short or lacking; 6) posterior margin of styliger plate gently convex (Fig. 74H); 7) central sclerite dome-shaped, apophyses of styliger sclerite straight (Fig. 74H); 8) apical margin of penes elliptically curved; lobes of penes flat located on wide shaft, only weakly distinct (Fig. 74H); 9) apex of forceps rounded, although somewhat tapered toward apex and gently curved toward median line in basal third to half; surface with numerous recumbent setae (Figs. 74I-J); and 10) abdominal terga I-II with apical margins dark and 2 oblong lateral spots; in general aspect a light brownish yellow species. Distribution: Brazil (Belterra, below Rio Tapajós). Caenis sigillata MALZACHER Caenis sigillata MALZACHER, 1986: 96 (male).

Taxonomy: This species, described from male imagos, can be characterized in the following manner: 1) Body length 1.7-2.3 mm; 2) antennal bristle with slightly dilated base,

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dilation short and symmetrical; 3) prosternal triangle narrow and rounded anteriorly (Fig. 74R); 4) finger-like process on abdominal tergum II absent; 5) lateral filaments of abdominal segments apparently short or lacking; 6) styliger plate usually covering penes completely, with posterior margin somewhat convex (Fig. 74O); 7) central sclerite elongate and domeshaped, apophyses of styliger sclerite short and straight (Fig. 74O); 8) penes very short and swollen toward base with lobes weakly or not differentiated (Fig. 74O); 9) forceps short and broad, apex rounded with sharp spines apically, surface with very fine setae (Figs. 74P-Q); and 10) body strongly pigmented; in general aspect a reddish brown species. Distribution: Brazil (Rio Paru). Caenis albata NAVÁS Nomen Dubium Caenis albata NAVÁS, 1920c: 12; HUBBARD, 1982a: 261. Ordella albata; LESTAGE, 1931b: 59.

Taxonomy: This species was described very inadequately based on a single female. Because the type is apparently lost and it is not possible to recognize it based on the original description, we consider this species nomen dubium. Distribution: Argentina (Buenos Aires). Caenis axillata NAVÁS Nomen Dubium Caenis axillata NAVÁS, 1930d: 359; ULMER, 1938: 107; HUBBARD, 1982a: 261; ALBA-TERCEDOR & PETERS, 1985: 219; ALBA-TERCEDOR & MOSQUERA, 1999: 65.

Taxonomy: This species which was described very inadequately, were outstanding by “a greenish plate at base of main veins” to which the name alludes. Given that the description is not sufficient to identify this species, and that the type is very damaged, we consider this species nomen dubium. Distribution: Chile. Caenis chilensis NAVÁS Nomen Nudum Caenis chilensis [nomen nudum]; BERTHÉLEMY, 1965: 2; ALBA-TERCEDOR & MOSQUERA, 1999: 62.

Taxonomy: Material labeled by NAVÁS as “Typus” of this species is deposited in the Museum d´Histoire Naturelle de Paris, although apparently it was never described. ALBATERCEDOR & MOSQUERA (1999) considered this species nomen nudum. Distribution: Chile. Caenis nigella NAVÁS Nomen Dubium Caenis nigella NAVÁS, 1933b: 82; HUBBARD, 1982a: 261.

Taxonomy: This species was described very inadequately and the type is apparently lost. Therefore we consider this species nomen dubium. Distribution: Chile.

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Genus Cercobrachys SOLDÁN Cercobrachys SOLDÁN, 1986: 336. (Type-species: Cercobrachys etowah SOLDÁN, original designation).

Taxonomy: Cercobrachys, known principally on the basis of nymphs, was established by SOLDÁN in 1986, including at that time 2 species from South America. Adult Characteristics: 1) Antennal pedicel at most 1.1 to 1.5 times as long as scape; 2) prosternum broad and anterior coxae widely separated; 3) spines on abdominal segment VI considerably curved toward median line; and 4) grooves of the forceps tortuous. Nymphal Characteristics: 1) Antennal pedicel at most 1.1 to 1.5 times as long as scape (Figs. 77A-B); 2) ocellar tubercles short (Fig. 77A); 3) maxillary and labial palpi 2segmented (Fig. 77B); 4) prosternum broad and anterior coxae widely separated; 5) legs long and thin, with long setae (Fig. 77D); 6) tibiae usually equal in length to or shorter than tarsi, and claws generally without denticles (Fig. 77D); and 7) posterolateral projections on abdominal segment VI curved toward median line (Figs. 77E-H) to form a “gill basket.” Distribution and Biology: Holarctic, Neotropical and Oriental. Very little is known about the biology of the nymphs of this genus, except for 1 species from North America which has been collected from streams with sandy bottoms covered with silt, and from logs in deep water. KEY

TO

SOUTH AMERICAN CERCOBRACHYS

Nymphs 1 Pronotum with a submarginal border; base of lateral ocellar tubercles narrower than ocelli; segment II of maxillary palpi 1/2 width and twice length of segment I (Fig. 77B); operculate gills on abdominal segment II at least 1.6 times as long as wide; posterolateral spine of abdominal segment VI 3 times longer than wide in dorsal view, not continuing to median line (Figs. 77E-F) ..... Cercobrachys peruanicus – Pronotum smooth, without grooves; base of lateral ocellar tubercles as wide as ocelli; segment II of maxillary palpi almost as wide and 1.7 to 1.9 times as long as segment I; operculate gills on abdominal segment II at most 1.3-1.4 times as long as wide; posterolateral spine of abdominal segment VI 5-6 times longer than wide in dorsal view, continuing to the median line (Figs. 77G-H) ...................................... .......................................................................................................... Cercobrachys colombianus CLAVE

PARA LOS

CERCOBRACHYS

SUDAMERICANOS

Ninfas 1

Pronoto con un reborde submarginal; base de los tubérculos ocelares laterales más angosto que los ocelos; segmento II de los palpos maxilares 1/2 del ancho y 2 veces el largo del segmento I (Fig. 77B); branquia opercular del segmento abdominal II por lo menos 1,6 veces más larga que ancha; espina posterolateral del segmento abdominal VI 3 veces más larga que ancha en vista dorsal, no contiguas en el medio (Figs. 77E-F) .......................................... Cercobrachys peruanicus

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Pronoto liso, sin surcos; bases de los tubérculos ocelares laterales tan anchas como el ocelo; segmento II de los palpos maxilares casi tan ancho y 1,7 a 1,9 veces más largo que el segmento I; branquia opercular del segmento abdominal II como máximo 1,3 a 1,4 veces más larga que ancha; espinas del segmento VI 5-6 veces más larga que ancha en vista dorsal, tocándose en el medio (Figs. 77G-H) ........... .......................................................................................................... Cercobrachys colombianus

Cercobrachys colombianus SOLDÁN Cercobrachys colombianus SOLDÁN, 1986: 340 (nymph); MALZACHER, 1986: 98.

Taxonomy: Cercobrachys colombianus, known only from nymphs, can be characterized by the following combination of characters: 1) Ocellar tubercles large, triangular, and wide at base; 2) pronotal spines absent; 3) operculate gill symmetrical, 1.3 times longer than wide; and 4) lateral spines of abdominal segment VI very long, bent medially and touching in the middle (Figs. 77G-77H). Distribution: Colombia (Prov. de Tolima); Brazil (Rio Marauia, Tapuruquara). Cercobrachys peruanicus SOLDÁN Brachycercus nymph EDMUNDS, JENSEN & BERNER, 1976: 266. Cercobrachys peruanicus SOLDÁN, 1986: 343 (nymph).

Taxonomy: Cercobrachys peruanicus is known only from nymphs, and can be characterized by the following combination of characters: 1) Ocellar tubercles rounded and as long as width of an ocellus; 2) pronotal spines very obtuse, forming a submarginal border; 3) operculate gill elongated, 1.6 times longer than wide; and 4) lateral spines of abdominal segment VI shorter, not touching in the middle (Figs. 77E-77F). Distribution: Peru (San Martín prov. [Río Tulumayo] and Loreto prov. [Río Yurac]) and Bolivia (Tarija).

A

B

C

D

E

F

G

H

Plate 77. Cercobrachys, nymphs. Figs. 77A-H. Cercobrachys sp.: 77A, head; 77C, scape and pedicel of antenna. C. peruanicus: 77B, maxillary palp; 77D, middle leg; 77E, posterolateral spines of abdominal segments, d.v.; 77F, same, l.v. C. colombianus: 77G, posterolateral spines of abdominal segments, d.v.; 77H, same, l.v. Figs 77A-H modified from SOLDÁN, 1986.

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FAMILY COLOBURISCIDAE Taxonomy: This family is composed by three genera: Coloburiscoides (Australia), Coloburiscus (New Zealand), and Murphyella (South America). This group is related to the family Oligoneuriidae, showing nymphs of similar aspect, with specialized structures used for filtering food. Adult Characteristics: 1) Fore wings with numerous longitudinal and cross veins (Fig. 78A); 2) hind wings large (Fig. 78B); 3) first tarsal segment of fore legs very short; 4) terminal filament (adult and nymph) present, reduced or vestigial. Nymphal Characteristics: 1) Pro- and mesonotum fused forming a notal shield (Fig. 79A); 2) fore and middle legs with rows of long filtering setae, dorsum of all femora with transverse rows of stout spines (Fig. 79A); 4) abdominal gills present on segments I-VII (in Australian and New Zealand genera) or completely lost (in the South American Murphyella). Distribution: Australia, New Zealand, Southern Argentina and Chile. Biology: Nymphs inhabit cold water rivers and streams. Genus Murphyella LESTAGE Metamonius [partim]; NEEDHAM & MURPHY, 1924: 29; NAVÁS, 1935: 140. Murphyella LESTAGE, 1930b: 439; LESTAGE, 1931b: 49; ULMER, 1932: 217; EDMUNDS & TRAVER, 1954: 237; DEMOULIN, 1955b: 5; DEMOULIN, 1955d: 9; EDMUNDS et al., 1963: 11; HUBBARD, 1982a: 12; HUBBARD, 1985: 11; DOMÍNGUEZ et al., 1994: 34; KLUGE, 2004: 128. (Type-species: Murphyella needhami LESTAGE, original designation). Heptagenia [partim]; NAVÁS, 1930a: 331. Dictyosiphlon LESTAGE, 1931b: 47. (Type-species: Heptagenia molinai NAVÁS, monotypy).

A

B

C

Plate 78. Murphyella needhami, male imago. Figs. 78A-C. 78A, fore wing; 78B, hind wing; 78C, genitalia v.v. Figs. 78A-C modified from DEMOULIN, 1955d.

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Taxonomy: This genus, known only based on a single species, is closely related to Coloburiscus from New Zealand. Adult Characteristics: 1) The cross veins of the fore wings are anastomosed (Fig. 78A); 2) the costal projection of the hind wings is sharp (Fig. 78B); 3) remnants of the thoracic gills are present; 4) the male genitalia are as in Fig. 78C; 5) terminal filament vestigial. Nymphal Characteristics: 1) Abdominal gills are absent (Fig. 79A); 2) with respiratory evaginations on the thoracic sterna and at the base of maxillae; 3) terminal filament vestigial. Distribution: Southern Argentina and Chile. Biology: The nymphs inhabit rivers with well-oxygenated cold water. Sometimes they were collected from sand and woody debris on the bottom of swift current streams, but are also found under rocks. The nymphs present tufts of setae on the fore legs that are used to filter particles from the water current. Murphyella needhami LESTAGE Metamonius? sp.? NEEDHAM & MURPHY, 1924: 29. Murphyella needhami LESTAGE, 1930b: 439; LESTAGE, 1931b: 49 (nymph); LESTAGE, 1935a: 97; ULMER, 1938: 96; DEMOULIN, 1955b: 5 (male); DEMOULIN, 1955c: 2; DEMOULIN, 1955d: 9 (male, nymph); EDMUNDS et al., 1963: 29 (nymph); KOSS & EDMUNDS, 1974: 304 (egg); HUBBARD, 1985: 12; DOMÍNGUEZ et al., 1994: 34 (male, nymph). Heptagenia? molinai NAVÁS, 1930a: 331 (female); ALBA-TERCEDOR & PETERS, 1985: 222. Dictyosiphlon molinai; LESTAGE, 1931b: 50; LESTAGE, 1935a: 97; ULMER, 1938: 90 (male, female, nymph). Metamonius needhami NAVÁS [nec LESTAGE], 1935: 140; DEMOULIN, 1955c: 3. Murphyella molinai; HUBBARD, 1982a: 272.

Taxonomy: This is the single species of the genus and can be characterized, in addition to the characters mentioned in the generic description, by the presence on abdominal terga I-X of a medio-longitudinal brown band and a trapezoidal macula on the latero-posterior angles of the abdominal terga. Distribution and Biology: See generic section.

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A

Plate 79. Murphyella needhami. Fig. 79A, nymphal habitus. Drawing by C. Molineri.

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FAMILY CORYPHORIDAE Taxonomy: This monotypic family was erected recently by MOLINERI et al. (2001) for Coryphorus aquilus PETERS. This relictual taxon apparently represents the sister group of the family Leptohyphidae (MOLINERI & DOMÍNGUEZ, 2003), showing many unique characters. As the nymphs have some similarities with the Afrotropical genus Machadorythus, they were once classified in the same subfamily (Machadorythinae, Tricorythidae). Adult Characteristics: 1) Absence of cubital intercalaries in fore wings (Figs. 80AB); and 2) eyes of male greatly enlarged (Fig. 80C). C D A

B A CuP

G

CuA

E

I

F J M

H

K

L

Plate 80. Coryphorus aquilus. Figs. 80A-E, imago: 80A, male fore wing; 80B, female fore wing; 80C, male head, d.v.; 80D, detail of fore tarsal claw of male imago; 80E, male genitalia, v.v. Figs. 80F-M, nymph: 80F, head, frontal view; 80G, labrum, d.v.; 80H, left mandible, d.v.; 80I, labial palp, detail; 80J, labium, v.v. at left, d.v. at right; 80K, right maxilla, v.v.; 80L, hypopharynx, v.v.; 80M, right mandible, d.v. Figs. 80C-E modified from MOLINERI et al., 2001; 80F-M from PETERS, 1981.

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Nymphal Characteristics: 1) Quadrangular operculate gills on segment II (Fig. 81I); forming a unique gill chamber (Figs. 81G-H). Distribution: Northern Brazil, Colombia, French Guyana. Biology: The biology of this taxon is almost unknown. Nymphs were collected in streams of the Amazonic and Orinoquian basins. Adults were attracted to light traps. Genus Coryphorus PETERS Coryphorus PETERS, 1981: 209 (nymph); LANDA & SOLDÁN, 1985: 104; PETERS & PETERS, 1993: 45; MCCAFFERTY & WANG, 2000: 58; MOLINERI et al., 2001: 118 (adults, egg). (Type-species: Coryphorus aquilus PETERS, original designation).

Taxonomy: Coryphorus was described from nymphs by PETERS (1981), WIERSEMA & MCCAFFERTY (2000) proposed it to be a member of their subfamily Tricorythodinae, but MOLINERI et al. (2001), who described adults of both sexes and eggs, showed that Coryphorus is not closely related to leptohyphid genera (also see MOLINERI & DOMÍNGUEZ 2003). Only one species is known, but ORTH et al. (2000) reported a probably undescribed species of the genus from French Guiana. Adult Characteristics: 1) Absence of intercalaries in cubital field of fore wings (Figs. 80A-B); 2) hind wings absent in both sexes; 3) eyes of male very large (Fig. 80C); 4) male with large, fused, distally broadened penes and short unsclerotized forceps (Fig. 80E); 5) fore tarsal claws of a pair dissimilar in male imagos (Fig. 80D). Nymphal Characteristics: In the nymphs (Fig. 81A): 1) Vertex of head with a pair of long slender submedian tubercles (Fig. 80F); 2) labrum without anteromedian deep emargination (Fig. 80G); 3) mandibles as in Figs. 80H, M; 4) labium broad (Fig. 80J), labial palpi 3-segmented (Figs. 80I-J); 5) maxillary palpi absent (Fig. 80K); 6) hypopharynx as in Fig. 80L; 7) legs long and slender (Figs. 81B-D), with serrated hind margin (detail in Fig. 81E); 8) basal 1/2 of tarsal claws (Fig. 81F) with six small blunt denticles; 9) gills on abdominal segments II-V (Fig. 81G), dorsally rectangular (Fig. 81I) and ventrally bifurcated (Figs. 81J-K), partially confined to a chamber formed by projections of terga III-VI curved dorsally (Figs. 81G-H). Distribution and Biology: Forested streams in northern Brazil, Colombia, and French Guiana. Nymphs of the only known species were found in a permanent stream, approximately 2 m wide and with a current velocity of 30 cm s-1; the water temperature was 28ºC. The stream was strongly shaded by the surrounding Amazon forest, the stream bed was sandy with many leaves and submerged branches, under which the nymphs were collected. The water was tea-colored, because of the presence of humic substances. The adults were captured with a light trap, imagos at dusk and subimagos at dawn. Coryphorus aquilus PETERS Coryphorus aquilus PETERS, 1981: 211 (nymph); MOLINERI et al., 2001: 118 (adults and egg).

Taxonomy: Coryphorus aquilus is the only known species in the genus, and can be recognized by the characters listed in the generic diagnosis.

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B C D E

F

A

G DE

GB

H

DE

I

J

K

Plate 81. Coryphorus aquilus, nymph. Figs. 81A-K. 81A, habitus; 81B, hind leg; 81C, middle leg; 81D, fore leg; 81E, detail of hind margin of fore leg; 81F, detail of fore tarsal claw; 81G, abdomen, l.v. (gill II-IV extracted); 81H, abdomen, d.v.; 81I, gill II, d.v.; 81J, gill II, v.v.; 81K, gill III, v.v. (DE = dorsal extensions; GB = gill base). Figs. 81A-F, I-K, modified from PETERS, 1981.

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FAMILY EPHEMERIDAE Taxonomy: The family Ephemeridae contains the following genera: Eatonica, Eatonigenia, Ephemera, Hexagenia, Ichthybotus, and Litobrancha. Only Hexagenia is represented in South America. Adult Characteristics: 1) Very large mayflies, male with moderate to large eyes; 2) all legs of both sexes functional and well developed; 3) fore wings (Fig. 82A) with veins MP2 and CuA strongly divergent from MP1 at base; vein A1 attached to hind margin by 2 to many veinlets; 4) forceps of male three or four-segmented (Figs. 82C, E); 5) terminal filament normal or vestigial in both sexes. Nymphal Characteristics: 1) Mandibular tusks upcurved distally (Fig. 3F); 2) gills arising dorsally, gills I reduced, gills II-VII with fringed margin (Fig. 83A); 3) hind tibiae with an apical projection (Fig. 3B); 4) three caudal filaments present. Distribution: Cosmopolitan (except Australia). Biology: The nymphs are burrowers in the bottom of rivers and lakes, making Ushaped tunnels in relatively soft mud or clay substrates. They feed by filtering suspended particles, or crawl out of the tunnels to collect food directly from the bottom. Adults emerge synchronously in huge numbers. Genus Hexagenia WALSH Hexagenia WALSH, 1863: 197. (Type-species: Ephemera limbata SERVILLE, subsequent designation by EATON, 1868a: 85).

Taxonomy: The genus Hexagenia consists of two subgenera: Hexagenia s.s. (distributed from Mexico north to Canada) and Pseudeatonica (from Mexico south to Argentina). Adult Characteristics: 1) Cross veins in fore wings not bunched up at bulla and from three to 14 veins connecting vein A1 to posterior margin (Fig. 82A); 2) genital forceps three- or four-segmented (Figs. 82C, E); and 3) abdomen generally with a distinctive dark color pattern (Fig. 82D). Nymphal Characteristics: 1) Mandibular tusks well developed, projecting in front of head, curved upwards distally, with long setae on medial and lateral margins (Fig. 3F); and 2) dorsal abdominal gills present, gills on segment I reduced, and dorsal portion of gills on segment II with basal portion expanded. Distribution: Americas and possibly Asia. Subgenus Hexagenia (Pseudeatonica) SPIETH Hexagenia (Pseudeatonica) SPIETH, 1941: 269. (Type-species: Hexagenia mexicana EATON, original designation).

Taxonomy: Hexagenia (Pseudeatonica) is the only subgenus of this family present in South America. The nymphs are indistinguishable from those of the other subgenus, Hexagenia (Hexagenia), but the geographic disjunction (Hexagenia s.s. distributed from Mexico northward) should permit separation of the subgenera. The adults are distinct and can be distinguished by the following combination of characters: 1) Vein A1 of fore wings connected to wing margin by three to six veins (Fig. 82A); and 2) genital forceps three-segmented, first segment straight

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and approximately 1/2 length of second which is long, robust, and strongly curved in basal 1/ 3, and third segment short and thin (Figs. 82C, E). Some authors have doubts about whether Hexagenia (Pseudeatonica) really belongs to Hexagenia or should be considered a separate genus. Distribution: South America to southern Mexico. KEY

HEXAGENIA (PSEUDEATONICA)

TO

Imagos 1 – 2(1) – CLAVE

Wing membranes uniformly tinted brown .......... Hexagenia (Pseudeatonica) mexicana Wing membranes hyaline or tinted yellow, but not uniformly ................................. 2 Abdominal sterna tinted reddish-brown laterally ..... Hexagenia (Pseudeatonica) albivitta Abdominal sterna pale, not stained laterally ....... Hexagenia (Pseudeatonica) callineura PARA

HEXAGENIA (PSEUDEATONICA)

Imagos 1 –

Membrana de las alas teñidas uniformemente de castaño .......................................... ....................................................................................... Hexagenia (Pseudeatonica) mexicana Membrana de las alas hialina o teñidas de amarillento pero no de forma uniforme ..... 2

A2 A1

A

B

C

D

E

Plate 82. Hexagenia (Pseudeatonica), adults. Figs. 82A-E: H. (P.) albivitta: 82A, male fore wing; 82B, male hind wing; 82C, male genitalia, v.v.; 82D, abdominal color pattern, d.v. H. (P.) mexicana: 82E, male genitalia, v.v. Fig. 82C modified from NEEDHAM & MURPHY, 1924; 82D from SPIETH, 1941 and 82E from KIMMINS, 1960.

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Esternitos abdominales teñidos de ferruginoso lateralmente .................................... ......................................................................................... Hexagenia (Pseudeatonica) albivitta Esternitos abdominales pálidos, no teñidos lateralmente ........................................... ....................................................................................... Hexagenia (Pseudeatonica) callineura

Hexagenia (Pseudeatonica) albivitta (WALKER) Baetis albivitta WALKER, 1853: 566; HAGEN, 1861: 304. Hexagenia albivitta; EATON, 1871: 64 (male); EATON, 1873: 392; EATON, 1883-1888: 49 (male, female); HAGEN, 1890: 11 (male, female); ULMER, 1920c: 108; NEEDHAM & MURPHY, 1924: 27 (male, female); NAVÁS, 1930e: 130; LESTAGE, 1931b: 59; NAVÁS, 1933c: 63; TRAVER, 1947b: 370; KIMMINS, 1960: 309. Hexagenia (Pseudeatonica) albivitta; SPIETH, 1941: 270 (male, female); MCCAFFERTY, 1970: 225 (nymph); HUBBARD, 1982a: 262. Palingenia continua WALKER, 1860: 199 (male). Hexagenia continua KIMMINS, 1960: 309. Palingenia dorsigera [nomen nudum] HAGEN, 1861: 304. Hexagenia benedicta NAVÁS, 1922a: 55 (female); LESTAGE, 1931b: 59; ALBA-TERCEDOR & PETERS, 1985: 222. Hexagenia dominans NAVÁS, 1936b: 732.

Taxonomy: Hexagenia (Pseudeatonica) albivitta has a wide range, both geographically and ecologically, since it has been collected from cold mountain streams in Patagonia to much warmer and slower waters of Brazil and Costa Rica. The imagos can be differentiated by the following combination of characters: 1) Membrane of fore wings tinted yellowish gray, much darker in costal and subcostal areas; 2) cross veins of hind wings shaded brown (Fig. 82B); 3) abdominal terga with a black saw-tooth line running length of abdomen dorsal to whitish spiracular line (Fig. 82D); 4) abdominal sterna reddish brown laterally, with median line lighter; and 5) genitalia as in Fig. 82C. The nymph of this species was illustrated by MCCAFFERTY (1970), although he gave no specific characters. The association was based on the similarity of color patterns between the nymphs and adults of this species, both collected at the same locality. Distribution and Biology: Central America, Colombia, Brazil, Guyana, Paraguay, Uruguay, and Argentina. The nymphs of this species have been collected burrowing in deposits of mud, in places in the river with little current or between rocks. Because of the large latitudinal range of this species, there may be seasonal emergence at some localities and not at others. According to people in areas where the species has been collected, massive emergences of adults occur after the rainy season. Hexagenia (Pseudeatonica) callineura BANKS Hexagenia callineura BANKS, 1914: 613 (female). Hexagenia (Pseudeatonica) callineura; SPIETH, 1941: 271 (female); MCCAFFERTY, 1970: 225; HUBBARD, 1982a: 262.

Taxonomy: Hexagenia (Pseudeatonica) callineura is known only from female imagos, which can be differentiated by the following combination of characters: 1) Wing membrane hyaline; 2) cross veins of hind wings shaded with blackish; 3) color pattern of abdominal terga similar to H. (P.) albivitta; and 4) abdominal sterna without spots. Distribution: Colombia and Ecuador.

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Hexagenia (Pseudeatonica) mexicana EATON Hexagenia mexicana EATON, 1883-1888: 50, (male); ULMER, 1921: 235; KIMMINS, 1934: 339 (male). Hexagenia (Pseudeatonica) mexicana; SPIETH, 1941: 272 (male); TRAVER, 1947b: 370; MCCAFFERTY, 1970: 225; HUBBARD, 1982a: 262. Pseudeatonica mexicana; KIMMINS, 1960: 312.

Taxonomy: Hexagenia (Pseudeatonica) mexicana is known only from male imagos and can be distinguished by the following combination of characters: 1) Wing membranes transparent brownish; 2) hind wings darker than fore wings with cross veins brownish black; 3) abdominal color pattern formed by three yellowish areas, one median area separated from two lateral ones by brownish black areas; 4) abdominal sterna pale; and 5) genitalia as in Fig. 82E. Distribution: Mexico, Costa Rica, Colombia, Ecuador, and Peru.

A

Plate 83. Hexagenia (Pseudeatonica) albivitta. Fig. 83A, nymphal habitus. Drawing by A. Dupuy.

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FAMILY EUTHYPLOCIIDAE The family Euthyplociidae, following the classification of GILLIES (1980) and LANDA & SOLDÁN (1985), is divided into two subfamilies: Euthyplociinae and Exeuthyplociinae. The first of the subfamilies (Euthyplociinae) is distributed in South America (Campylocia, Euthyplocia, and Mesoplocia), Madagascar (Proboscidoplocia), and Asia (Polyplocia). Some records cite Euthyplocia from Asia but this is improbable. The second subfamily (Exeuthyplociinae) is restricted to Africa (Afroplocia and Exeuthyplocia). SUBFAMILY EUTHYPLOCIINAE Adults Characteristics: 1) Eyes of both sexes of approximately same small size; 2) vein R1 of fore wings bifurcated at least 1/6 of distance from base to margin, vein MA bifurcated near or further than bifurcation of vein R1, and cubital region with 6 or more sigmoidal veins uniting vein CuA with posterior margin (Figs. 84A, D, G, 85A, 87A); and 3) legs well developed. Nymphal Characteristics: 1) Mandibular tusks longer than head, with numerous long setae (Fig. 86A); 2) legs not modified for burrowing (Fig. 86A); 3) abdominal gills small and ovoid on segment I and those on segments II-VII extended laterally from abdomen and with margins fringed (Fig. 86A). Distribution: South and Central America, Madagascar, Asia. Biology: The nymphs are large crawlers but can burrow horizontal trenches in the fine sediment below rocks, using their mandibular tusks. Apparently they are collectors gatherers of fine particulate organic matter. Subimagos generally emerge at dusk, molting to imago in a few minutes and then performing the mating swarm. KEYS TO SOUTH AMERICAN GENERA Adults 1 – 2(1) –

Cubital area of fore wings with one to three intercalaries parallel to CuA (Figs. 84A, D, G); vein CuA not connected to the posterior margin by more than 10 sigmoidal intercalaries; forceps one-segmented .......................................... Campylocia Cubital area of fore wings without intercalaries parallel to CuA (Figs. 85A, 87A); vein CuA connected to the posterior margin by more than 10 sigmoidal intercalaries (Figs. 85A, 87A); forceps two-segmented .............................................. 2 Hind wing triangular, almost half the length of the fore wing (Fig. 85B) .............. ............................................................................................................................... Euthyplocia Hind wing elongate, approximately 1/4 the length of fore wing (Figs. 87B-C) .... ................................................................................................................................. Mesoplocia

Nymphs 1 –

Antennae shorter than mandibular tusks; fore tibial spine approximately 1/4 length of tarsus (Fig. 84J) ................................................................................. Campylocia Antennae 3 times as long as mandibular tusks (Fig. 86A) or longer ...................... 2

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2(1) –

Fore tibial spine approximately 1/3 length of tarsus .................................. Mesoplocia Fore tibial spine more than 1/2 as long as tarsus (Fig. 85I) .................... Euthyplocia

CLAVES

PARA LOS

GÉNEROS

SUDAMERICANOS

Adultos 1 – 2(1) –

Area cubital de las alas anteriores con una a tres intercalares paralelas a CuA (Figs. 84A, D, G); vena CuA no conectada al margen posterior por mas de 10 sigmoidales intercalares; fórceps unisegmentados ...................................... Campylocia Area cubital de las alas anteriores sin intercalares paralelas a CuA (Figs. 85A, 87A); vena CuA conectada al margen posterior por más de 10 intercalares sigmoidales; fórceps bisegmentados ...................................................................................................... 2 Ala posterior triangular, casi la mitad de la longitud del ala anterior (Fig. 85B) .... ............................................................................................................................... Euthyplocia Ala posterior alargada, de aproximadamente 1/4 de la longitud del ala anterior (Figs. 87B-C) .................................................................................................................... Mesoplocia

Ninfas 1 – 2(1) –

Antenas más cortas que los colmillos mandibulares; espina de la tibia anterior aproximadamente 1/4 de la longitud del tarso (Fig. 84J) ............................ Campylocia Antenas tres veces más largas que los colmillos mandibulares (Fig. 86A), o más ...... 2 Espina de la tibia anterior aproximadamente 1/3 de la longitud del tarso ...... Mesoplocia Espina de la tibia anterior más de 1/2 de la longitud del tarso (Fig. 85I) ...... Euthyplocia

Genus Campylocia NEEDHAM & MURPHY Campylocia NEEDHAM & MURPHY, 1924: 25; BERNER & THEW, 1961: 329; DEMOULIN, 1966b: 140. (Type-species: Euthyplocia anceps EATON, original designation). Longinella GROS & LESTAGE, 1926: 158. (Type-species: Euthyplocia guntheri NAVÁS, original designation).

Taxonomy: The genus Campylocia was established by NEEDHAM & MURPHY (1924) who included three species in addition to the type-species Euthyplocia anceps EATON: Euthyplocia burmeisteri HAGEN, Campylocia ampla NEEDHAM & MURPHY, and Euthyplocia guntheri NAVÁS. GROS & LESTAGE (1926) created the new genus Longinella and designated Euthyplocia anceps as the type-species. ULMER (1932, 1942) eventually synonymized Longinella with Campylocia and synonymized all the species belonging to Campylocia at that time (C. ampla, C. burmeisteri, C. guntheri, and C. intercalata BANKS) with C. anceps, leaving this the only species in the genus. DEMOULIN (1952b) revised the subfamily Euthyplociinae and separated Campylocia and Polyplocia by wing characters. BERNER & THEW (1961) revised Campylocia once again and described the new species C. dochmia. In addition, based on a large series of specimens, they proposed as a new generic character the presence of five segments in the mesothoracic legs of the adults. PEREIRA & DA-SILVA (1990a) described a new species from Brazil, based on nymphs and adults of both sexes. Presently, the genus consists of these three species, two of which are known from both nymphs and adults.

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Adult Characteristics: 1) Cubital area of fore wings with 1 to 3 intercalary veins parallel to vein CuA (arrows in Figs. 84A, D, G), these intercalaries connected to hind margin by 6 or more sigmoidal intercalary veins (Figs. 84A, D, G); 2) hind wings subtriangular and almost 1/ 2 length of fore wings (Figs. 84B, E, H); 3) genital forceps without terminal segment (Figs. 84C, F, I); and 4) terminal filament of male imago not reduced. Nymphal Characteristics: 1) Antennae shorter than mandibular tusks; 2) tibial spine of fore legs approximately 1/4 length of tarsus (Fig. 84J); and 3) fore tarsus without a distal projection. Distribution: Lowlands of Central and South America. Biology: The nymphs can be found below rocks in warm rivers and streams. The imagos fly at dusk below the canopy. KEYS

TO

SOUTH AMERICAN CAMPYLOCIA

Male Imagos 1 – 2(1) –

Penes lobes widely divergent along the medial line, penes without median lobes (Fig. 84C); fore wings with one cubital intercalary vein (parallel to vein CuA) (in large individuals two intercalaries may be present) (Fig. 84A) ................... Campylocia anceps Inner margin of the penes lobes adjacent to the median line, median lobes present on penes (Figs. 84F, I); fore wings with two or three cubital intercalary veins (parallel to vein CuA) (Figs. 84D, G) ............................................................................. 2 Two intercalary cubital veins present in fore wings (Fig. 84D) ....... Campylocia dochmia Three intercalary cubital veins present in fore wings (Fig. 84G) ............................... ..............................................................................................................Campylocia bocainensis

Nymphs 1 –

CLAVES

Abdominal gills light-colored with darker fringe; antennae 3/4 to subequal in length to the mandibular tusks ............................................................. Campylocia anceps Abdominal gills dark brown with lighter fringe; antennae approximately 2/3 the length of the mandibular tusks .....................................................Campylocia bocainensis PARA LOS

CAMPYLOCIA

SUDAMERICANOS

Imagos Machos 1 – 2(1) –

Lóbulos de los penes ampliamente divergentes sobre la línea medial, penes sin lóbulos mediales (Fig. 84C); alas anteriores con una vena intercalar cubital (paralela a CuA) (en individuos grandes puede haber dos ICu) (Fig. 84A) .......................... Campylocia anceps Margen interno de los lóbulos de los penes adyacentes a la línea medial, lóbulos mediales presentes en los penes (Figs. 84F, I); alas anteriores con dos ó tres venas intercalares cubitales (paralelas a CuA) (Figs. 84D, G) .............................................. 2 Dos venas intercalares cubitales presentes en las alas anteriores (Fig. 84D) .......... .................................................................................................................. Campylocia dochmia Tres venas intercalares cubitales presentes en las alas anteriores (Fig. 84G) .......... ..............................................................................................................Campylocia bocainensis

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C A

B

F D LP

E

G

LP

H

J

I

Plate 84. Campylocia. Figs. 84A-J. Figs. 84A-I, male imago. C. anceps: 84A, fore wing; 84B, hind wing; 84C, genitalia, v.v. C. dochmia: 84D, fore wing; 84E, hind wing; 84F, genitalia, v.v. C. bocainensis: 84G, fore wing; 84H, hind wing; 84I, genitalia, v.v. 84J, nymph: Campylocia sp.: 84J, fore leg. (LP = medial lobes of penes; Arrows = intercalary veins). Figs. 84G-I modified from PEREIRA & DA-SILVA, 1990a.

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Ninfas 1 –

Branquias abdominales claras orladas de filamentos oscuros; antenas 3/4 o subiguales a la longitud de los colmillos mandibulares ................... Campylocia anceps Branquias abdominales oscuras orladas de filamentos claros; antenas 2/3 la longitud de los colmillos mandibulares ....................................... Campylocia bocainensis

Campylocia anceps (EATON) Euthyplocia anceps EATON, 1883-1888: 38 (male); HAGEN, 1888: 226 (male, female); ULMER 1920c: 104; ULMER, 1920b: 3 (nymph); LESTAGE, 1924c: 38; GROS & LESTAGE, 1926: 121 (nymph). Campylocia anceps; NEEDHAM & MURPHY, 1924: 27 (male, female); GROS & LESTAGE, 1926: 162; ULMER, 1932: 207; ULMER, 1942: 101 (male, female); SPIETH, 1943: 1 (male, female); TRAVER, 1944: 10 (nymph); DEMOULIN, 1952b: 13; KIMMINS, 1960: 307 (male); BERNER & THEW, 1961: 329 (male); HUBBARD, 1982a: 262; PEREIRA & DA-SILVA, 1990a: 2. Euthyplocia burmeisteri; LESTAGE, 1924c: 39; NEEDHAM & MURPHY, 1924: 25. Campylocia burmeisteri; NEEDHAM & MURPHY, 1924: 25. Euthyplocia intercalata BANKS, 1918: 10 (female); GROS & LESTAGE, 1926: 122. Euthyplocia guntheri; NAVÁS, 1920b: 414 (male); NEEDHAM & MURPHY, 1924: 25; GROS & LESTAGE, 1926: 121. Longinella guntheri; LESTAGE, 1931b: 59; GROS & LESTAGE, 1926: 161. Campylocia ampla NEEDHAM & MURPHY, 1924: 25 (female, egg); GROS & LESTAGE, 1926: 162.

Taxonomy: Campylocia anceps, originally described in Euthyplocia, is the type-species of the genus and is known from adults of both sexes and nymphs. The adults of this species can be separated from the other species of the genus by the following combination of characters: 1) Penes lobes diverge along median line (Fig. 84C); 2) subgenital plate with a convex to almost truncated posterior border (Fig. 84C); and 3) one ICu (intercalary cubital) vein in fore wings (Fig. 84A), but some large individuals (mainly females) present two ICu veins. In the nymphs: 1) gills clear-gray, with black fringe and general coloration of body brownish orange, darker on pronotum and dorsum of head; and 2) antennae 3/4 to subequal in length to mandibular tusks. Distribution: Brazil, Peru, Colombia, Venezuela, Ecuador, French Guiana, Guyana, Surinam, Central America. Campylocia bocainensis PEREIRA & DA-SILVA Campylocia bocainensis PEREIRA & DA-SILVA, 1990a: 2 (male, female, nymph).

Taxonomy: Campylocia bocainensis, known from adults and nymphs, can be distinguished by the following combination of characters. In the adult: 1) Inner margin of penes lobes joining at median line and medial lobes present (Fig. 84I); 2) subgenital plate with a slightly emarginated border (Fig. 84I); and 3) three ICu veins in fore wings (Fig. 84G). In the nymph: 1) abdominal gills dark brown, with lighter fringe; and 2) antennae approximately 2/3 length of mandibular tusks. Distribution and Biology: Brazil (São Paulo). The nymphs were collected in backwater areas, in a river some 70 cm deep. The subimagos emerged during dusk, molting to imagos after some 15 minutes. The imagos fly during dusk in large numbers near the surface of the water.

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Some specimens recently collected in Misiones (NE Argentina) apparently pertain to this species. Nevertheless, some variations occur, and for example, wing venation characteristics are intermediate with C. dochmia. For the moment, we do not know if the Argentinean material represents a new species or it suggests, instead, that C. bocainensis and C. dochmia are synonyms. Campylocia dochmia BERNER & THEW Campylocia dochmia BERNER & THEW, 1961: 332 (male, female); HUBBARD, 1982a: 262; PEREIRA & DASILVA, 1990a: 2.

Taxonomy: Campylocia dochmia, known only from adults of both sexes, can be separated from the other known species of the genus based on the following combination of characters: 1) Inner margins of penes lobes parallel to the median line (Fig. 84F); 2) medial lobes present on penes (Fig. 84F); 3) subgenital plate with a slightly emarginated posterior border (Fig. 84F); and 4) two ICu veins in fore wings (Fig. 84D). Distribution: Brazil (Minas Gerais). Genus Euthyplocia EATON Euthyplocia EATON, 1871: 67; DEMOULIN, 1966b: 140. (Type-species: Palingenia hecuba HAGEN, monotypy).

Taxonomy: The genus Euthyplocia was established by EATON (1871) based on a female described by HAGEN as Palingenia hecuba. Presently, the genus contains two described species, one known only from males and the other from all stages. Adult Characteristics: 1) Vein CuA of fore wings connected with posterior margins by more than 10 sigmoidal intercalary veins (Fig. 85A); 2) hind wings triangular, almost 1/ 2 length of fore wings (Fig. 85B); 3) genital forceps with a small terminal segment (Figs. 85E, H); and 4) terminal filament of male imago not reduced. Nymphal Characteristics: 1) Antennae 3 times as long as mandibular tusks (Fig. 86A); 2) tibial spine on anterior legs approximately 1/2 length of tarsus (Fig. 85I); and 3) fore tarsus with a distal projection. Distribution: South and Central America. Biology: Nymphs have been collected from small streams to medium-sized rivers, below rocks. Adults generally emerge in late afternoon but SWEENEY et al. (1995) reported adult activity during early morning; subimaginal stage lasts around 15 minutes. SWEENEY et al. (1995) also found that the egg stage lasted 2 months and nymphal stage around 2 years. KEY

SOUTH AMERICAN EUTHYPLOCIA

TO

Imagos 1 –

Light-colored species; last segment of the forceps 2 times as long as wide; penes lobes (Figs. 85F-H) scarcely or not at all narrowed apically, rounded at the apex .... .................................................................................................................... Euthyplocia hecuba Dark-colored species; last segment of the forceps 4 times as long as wide; penes lobes (Figs. 85C-E) elongated and narrowed toward the apex ................... Euthyplocia haenschi

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CLAVE

PARA LOS

EUTHYPLOCIA

SUDAMERICANOS

Imagos 1

Coloración general clara; último segmento de los fórceps 2 veces más largo que ancho; lóbulos del pene (Figs. 85F-H) apenas o nada estrechados apicalmente, redondeados en el ápice ....................................................................... Euthyplocia hecuba Coloración general oscura; último segmento de los fórceps 4 veces más largo que ancho; lóbulos del pene (Figs. 85C-E) alargados y adelgazados hacia la parte apical ................................................................................................................. Euthyplocia haenschi

–

A

B

C

F D

G I

E

H

Plate 85. Euthyplocia. Figs. 85A-I. Figs. 85A-H, male imago. Euthyplocia sp.: 85A, fore wing; 85B, hind wing. E. haenschi: 85C, detail of penes, l.v.; 85D, detail of penes, d.v.; 85E, male genitalia, v.v. E. hecuba: 85F, detail of penes, l.v.; 85G, detail of penes, d.v.; 85H, male genitalia, v.v.; 85I, nymphal leg I. Figs. 85C-H modified from ULMER, 1942.

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Euthyplocia haenschi ULMER Euthyplocia haenschi ULMER, 1942: 100 (male); DEMOULIN, 1952b: 11; HUBBARD, 1982a: 263.

Taxonomy: Euthyplocia haenschi, known only from male imagos, can be separated from the other known species of the genus by the following combination of characters: 1) General coloration blackish with a dark gray abdomen; 2) last segment of forceps four times as long as wide (Fig. 85E); and 3) penes lobes (Figs. 85C-E) elongated and narrowed toward apical part and apex rolled up in ventral view. Distribution: Ecuador (Santa Inés). Euthyplocia hecuba (HAGEN) Palingenia hecuba HAGEN, 1861: 40. Euthyplocia hecuba; EATON, 1871: 67 (female); EATON, 1873: 392; EATON, 1883-1888: 37 (female); HAGEN, 1888: 226 (male); EATON, 1892: 3 (male, female); ULMER, 1920c: 104; ULMER, 1920b: 10 (nymph); ULMER, 1921: 251 (female); LESTAGE, 1924c: 38; GROS & LESTAGE, 1926: 121 (nymph); ULMER, 1942: 99 (male, female); DEMOULIN, 1952b: 11; HUBBARD, 1982a: 263; SWEENEY et al., 1995: 141.

Taxonomy: Euthyplocia hecuba can be characterized by the following combination of characters. In the imagos: 1) General coloration grayish violet; 2) last segment of genital forceps twice as long as wide (Fig. 85H); and 3) penes lobes (Figs. 85F-H) not narrowed apically, but rounded at apex with a weak groove. According to ULMER (1942) this species can at times be much smaller than E. haenschi. In the nymph: 1) General coloration grayish violet; and 2) gills blackish violet. The nymphs of E. hecuba were described by ULMER (1920b) from material collected from San Lorenzo, Jujuy, Argentina, but this must be regarded with caution since one of us (ED) reared nymphs from localities near that cited by ULMER and the adults apparently corresponded to E. haenschi. SWEENEY et al. (1995) working in Costa Rica suggested that E. hecuba may be in fact represented by more than one identity. Analyzing the genomic structure of a Costa Rican population of E. hecuba and that of the type locality in Mexico, they found a high degree of genetic differentiation. Distribution and Biology: Argentina, Brazil, Colombia, Ecuador, Peru, Venezuela, Central America, Mexico. According to EDMUNDS et al. (1976) the nymphs are found on the surface of relatively large rocks. In the northwest part of Argentina, nymphs have been collected from small streams to medium-sized rivers. Mature nymphs occur generally in zones from 3 to 10 cm depth in the stream margins, under small rocks where they remain immobile or try slowly to burrow. Smaller nymphs are found in deeper zones with more current, adhering to rocks. In NW Argentina, adults emerge in the late afternoon, a little before sundown. Male subimagos cling to tree trunks and other surrounding objects and molt to imago in around 15 minutes. The nuptial flight takes place above the underbrush but below the tree canopy, generally in areas 10 to 20 m from the river. The males perform an undulating flight, forming groups of between 5 and 20 individuals. The females approach the swarm and after copulation withdraw to perch on surrounding vegetation. Generally males are attracted to light in the early hours of night and the females around midnight. SWEENEY et al. (1995) studied with detail the life cycle of this species in two rivers from Costa Rica. They found that all the activity of the adult stage (molt to imago, swarming,

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mate, and oviposition) took place in the early morning (not lasting more than 3 hours). Egg development time was two months at 20ºC; nymph development took 22 months and involved a developmental diapause during the first year. Two different cohorts of nymphs were present during the study (4 years), one that emerges this year, and other, smaller, that emerges the next year. Thus, they found that this mayfly show a semivoltine life cycle. Genus Mesoplocia DEMOULIN Mesoplocia DEMOULIN, 1952b: 11; DEMOULIN, 1966b: 139; KLUGE & NARANJO, 1994: 781; KLUGE, 2004: 229. (Type-species: Mesoplocia intermedia DEMOULIN, original designation).

Taxonomy: Mesoplocia was described by DEMOULIN (1952b) based on a single female, which lacked the anterior legs. Male adult and nymph remained unknown, until KLUGE &

A

Plate 86. Euthyplocia sp. Fig. 86A, nymphal habitus. Drawing by A. Dupuy.

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NARANJO (1994) described a new species from Cuba (sub. nom. Euthyplocia inaccessibile). In the same paper, these authors proposed Mesoplocia as a junior synonym of Euthyplocia. We do not agree with this synonymy, because the shape of hind wings in both sexes and the reduced terminal filament of the male imago are conspicuous generic differences. The only described South American species, M. intermedia, is known from female imagos. Adult Characteristics: 1) Cubital area of fore wings without intercalary veins parallel to vein CuA, and vein CuA united to posterior margin of wing by more than 10 sigmoidal intercalary veins (Fig. 87A); 2) hind wings elongated, approximately 1/4 length of fore wings (Figs. 87B-C); 3) genital forceps with a small terminal segment (Fig. 87D); 4) terminal filament of male imago reduced. Nymphal Characteristics: 1) Antennae long, 3 times length of tusks; 2) tibial spine on anterior legs approximately 1/3 length of tarsus; and 3) fore tarsus with a distal projection. Distribution: Ecuador, Cuba and Dominican Republic. Biology: Unknown. Mesoplocia intermedia DEMOULIN Mesoplocia intermedia DEMOULIN, 1952b: 12 (female); HUBBARD, 1982a: 263; KLUGE & NARANJO, 1994: 781; KLUGE, 2004: 229.

Taxonomy: The female of Mesoplocia intermedia can be characterized in the following manner: 1) General coloration brownish gray; 2) pronotum about twice as long as wide; and 3) costal and subcostal areas of fore wings and base of hind wing shaded with violet. Distribution: Ecuador (Paramba).

A

B

C

D

Plate 87. Mesoplocia, imagos. Figs. 87A-D. M. intermedia: 87A, fore wing; 87B, hind wing; 87C, detail of hind wing. Mesoplocia sp.: 87D, male genitalia. Fig. 87D drawn by J. Peters, from material collected by R. E. Woodruff in Dominican Republic.

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FAMILY LEPTOHYPHIDAE Taxonomy: This group was first recognized as a subfamily (Leptohyphinae) within Tricorythidae (EDMUNDS & TRAVER, 1954) and elevated to family by LANDA (1973). Recently, WIERSEMA & MCCAFFERTY (2000) divided Leptohyphidae in 2 subfamilies: Tricorythodinae and Leptohyphinae, but MOLINERI & DOMÍNGUEZ (2003) presented evidence that these are not natural groups. The number of genera in this Pan-American family has doubled in the last five years. All genera have been reported from South America (MOLINERI, 2004; EMMERICH, 2004), except Ableptemetes WIERSEMA & MCCAFFERTY (2003) recorded from Mexico and Central America. Allenhyphes, Tricorythodes, Leptohyphes and Vacupernius extend into North America. Adult Characteristics: 1) Small to medium sized mayflies (body length, 2-10 mm); 2) head with undivided, non-sexually dimorphic eyes (except Leptohyphodes and 1 species of Leptohyphes); 3) robust thorax and wings with fringed posterior margin in imago; 4) hind wings reduced in size and with a long costal projection (Fig. 88C), but in many genera hind wings absent in females, and absent in both sexes in others; 5) imagos with normal legs, with tarsal claws of a pair dissimilar (one acutely pointed, other paddle like), except those in male fore leg (both paddle-like); 6) male genitalia variable: forceps 2 or 3-segmented, penes divided in distal half or completely fused, and penes with spines of various forms; 7) caudal filaments of imagos long and thin, almost glabrous; generally shorter in female but sometimes greatly reduced (females of Tricorythopsis). Nymphal Characteristics: 1) Stout bodies with relatively short legs (except some Tricorythodes species), and generally body and legs covered by thin setae; 2) abdomen short and with tracheal gills on segments II-VI (except Leptohyphodes, on II-V); 3) first pair of gills enlarged and forming an operculum that covers remaining gills; 4) number of lobes in each gill is important in generic diagnosis and is here expressed as a “gill formula”, for example “3/3/3/3/2” meaning that first 4 pairs of gills (gills II-V) have 3 lobes each, and that gills VI have only 2 lobes; 5) caudal filaments, generally as long as body, with crowns of setae at each articulation; and often can be very thick in mature male nymphs. Distribution: South to North America. Biology: The nymphs live in various lotic habitats, and they are found mainly in fastwater zones, among boulders, gravel or sand. The presence of operculate gills facilitates the toleration of suspended solids, and in some very turbid rivers the family forms a major component of the benthic community. Apparently, the nymphs feed on fine particulate matter deposited in the substratum. Subimagos generally emerge during the night, molting to imago after 15 minutes to few hours. Imagos usually perform the nuptial flight at early morning, but sometimes swarming extends later in the day. Fertilized female imagos extrude the eggs in a single mass that is retained in the apex of abdomen and reeleased upon contact with the water surface.

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KEYS TO SOUTH AMERICAN GENERA Adults 1 – 2(1) – 3(1) – 4(3) – 5(4) – 6(5) – 7(6) – 8(7)

–

Male fore wings with enlarged cubito-anal lobe (widest part of wing at basal 1/3) (Fig. 108A); hind wings absent in both sexes; female caudal filaments reduced in length, shorter than body ................................................................................................. 2 Male fore wings with normal cubito-anal lobe (widest part of wing at 1/2) (Fig. 90A); hind wings absent or present; female caudal filaments not reduced, longer than fore wing ..................................................................................................................... 3 Forceps 3-segmented, long and slender, with swelling at base of segment II (Figs. 108I-S) (except T. australis, Fig.108H); female cerci as long as abdomen (the terminal filament may be longer) ................................................................ Tricorythodes Forceps 2-segmented and short, segment II without basal swelling and directed laterally (Figs. 113D-K); female caudal filaments very short, less than half length of abdomen ..................................................................................................... Tricorythopsis Eyes of male large and dorsal, divided into a whitish upper portion and a blackish smaller lower portion (Figs. 97C-D); hind wings absent in both sexes; forceps 2-segmented (Fig. 97F) ................................................................................ Leptohyphodes Eyes of male small, black, and lateral, not divided into upper and lower portions; hind wings present at least in males; forceps 2-segmented (Fig. 90G) or 3-segmented (Figs. 88D, 93G, 102E) ...................................................................................................... 4 Mesoscutellum without membranous filaments or with filaments not extending beyond the tip of mesoscutellum; forceps 2-segmented (Fig. 90G); hind wings present in both sexes ....................................................................................... Haplohyphes Mesoscutellum with relatively long membranous filaments on the tip; forceps 3-segmented (Figs. 88D, 93G, 102E); hind wings present or absent in female ....... 5 Penes divided on distal half, Y or T shaped, apex of each lobe formed by a rounded lobe with an acute extension (Figs. 93G, M); female hind wings present or absent .............................................................................................................. Leptohyphes Penes completely fused, or, if divided apically not strongly divergent (Figs. 88D, 100E, 102E, 103A, 117E); female hind wings absent ................................................ 6 Male terminal filament with well developed ventral spine at base (Fig. 88E); penes with a deep median cleft and without conspicuous spines (Fig. 88D) ................. Allenhyphes Male terminal filament without ventral spine, although some setae may be present; penes with or without spines ........................................................................................... 7 Penes with a pair of long sclerotized lateral filaments inserted at base (Figs. 117G, J, M) ............................................................................................................... Yaurina Penes not as above (Figs. 100E, 102E, 103A) ............................................................. 8 Forceps segment I almost as long as segment II (Figs. 100E, H, K); penes with a pair of ventrally visible (not completely hidden by penes) and long lateral spines, slightly curved medially (Fig. 100G); female gonopore (median zone between abdominal sterna VII and VIII) generally blackish .................................. Lumahyphes Forceps segment I less than 1/2 length of segment II (Fig. 102E); penes with or without sclerotized spines (may be hyaline), may be curved ventrally, generally

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hidden by the penes in ventral view; female gonopore generally whitish or slightly shaded with gray ................................................................................................................. 9 Penes broad (distal half markedly wider than basal half) and without spines; without posterolateral projections on styliger plate .................................. Vacupernius Penes slender, slightly wider at apex or similar in width for entire length; with spines (may be hyaline) of varied form, with (Figs. 102E, I, L) or without (Figs. 103A, E, H) well developed posterolateral projections on styliger plate ................. .............................................................................................................................. Traverhyphes

Nymphs 1

–

2(1)

– 3(2) – 4(3) – 5(3)

–

Operculate gills semi-triangular, meeting or almost meeting medially (Fig. 98A); gills present on abdominal segments II-V, margins of ventral lamellae fringed with small imbricated lobes (Figs. 99N-Q); eyes of male divided in upper whitish portion and blackish lower portion ........................................................... Leptohyphodes Operculate gills triangular, semi-triangular or oval, never meeting medially (Fig. 91A); gills present on abdominal segments II-VI, ventral lamellae without imbricated lobes at margin (Figs. 92R-V); eyes of male not divided, blackish and small ...................................................................................................................................... 2 Small body size (length of mature nymph = 1.0 to 3.0 mm), body stout and robust (Figs. 114A-G); hind wingpads absent in both sexes; operculate gills oval with a median or subapical weak transverse line (Figs. 114A, 115H-I); gill formula 5/4/ 4/2/1 (Figs. 115I-M) ..................................................................................... Tricorythopsis Larger body size (mature nymph > 3 mm); hind wingpads present or absent; operculate gills without transverse line; gill formula never as above ...................... 3 Operculate gills triangular or semitriangular (Figs. 92Q, 111A-C, E), if oval or subquadrate (Figs. 111D, F) then with broad femora (nearly as wide as long, Figs. 111K-N); fore femora with transverse row of long setae (Figs. 92F, 111I, K) ....... 4 Operculate gills oval or semi-oval (Figs. 88M, 94A, 106A), fore femora with transverse row of spines of variable length but never with long setae (Figs. 94A, 104A) .................................................................................................................................... 5 Body stout, base of abdomen much wider than apex (Fig. 109A); fore femora with long setae only (Figs. 111G, I); hind wingpads absent; gill formula 3/3/3/3/2 (Figs. 110R-V) or less ..................................................................................... Tricorythodes Body elongated, base of abdomen slightly wider than apex (Fig. 91A); fore femora with long setae and stout spines on leading edge (Figs. 92F-G); hind wingpads present; gill formula 4/5-4/4/4/2 (Figs. 92R-V) ..................................... Haplohyphes Operculate gills with basal spine (Fig. 95I); caudal filaments with darker segments at base (female) or medially (male) (Fig. 94A); dorsum of middle and hind femora with a longitudinal ridge and with a transverse row of spines near base (Fig. 95Q) ............................................................................................................................... Leptohyphes Operculate gills without basal spine (Figs. 101R-S); caudal filaments not as above; dorsum of middle and hind femora without longitudinal ridge or transverse row of spines near base (Figs. 101G-H) ............................................................................... 6

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6(5)

Transverse dorsal row of spatulate setae at midlength of fore femora ................... .............................................................................................................................. Vacupernius – Transverse dorsal row of spatulate setae in distal 1/3 of fore femora (Figs. 101I, O, 107B, F, K, O) ............................................................................................................... 7 7(6) Body relatively stout (fig.118A); maxillary palpi 2-segmented (Figs. 88K, 119C); operculate gills ovoid and relatively small, not reaching posterior half of abdominal segment VII (Fig. 118A); gill formula: 3/3/3/3/1-2 ............................ 8 – Body relatively elongated (fig. 104A); maxillary palpi 2 or 3-segmented (Figs. 101D, 105F-I); operculate gills ovoid to subquadrate and relatively large, at least reaching posterior half of abdominal segment VII (Fig. 104A); gill formula: 3/34/3-4/3/1-2 ........................................................................................................................ 9 8(7) Abdominal gill VI with 2 lamellae (Fig. 88R); maxillary palpi generally with apical seta, palpi inserted in straight or slightly concave margin (Figs. 88K-L); fore tarsal claw with a subapical single row of 4 denticles .......................... Allenhyphes (in part) – Abdominal gill VI with 1 lamella (Fig. 119Q); maxillary palpi without apical seta, palpi inserted in a concave margin (Fig. 119C); fore tarsal claw with a subapical single row of 6 to 7 denticles (Fig.119K) .......................................................... Yaurina 9(7) Maxillary palpi 3-segmented and relatively large (Figs. 89J, R-S, 105I) ................ 10 – Maxillary palpi 2-segmented and relatively small (Figs. 105F-H) .............................. ..............................................................................................................Traverhyphes (in part) 10(9) Spatulate setae on dorsum of fore femora bifid or serrate (Figs. 101J, P); female mature nymph generally with a median black macula between abdominal sterna VII and VIII ...................................................................................................... Lumahyphes – Spatulate setae on dorsum of fore femora not bifid and with smooth apex (Figs. 89I, O, 107G); sterna of female mature nymph not as above ............................... 11 11(10) Maxillary palpi with small apical seta (Fig. 105I) ......................Traverhyphes (in part) – Maxillary palpi without apical seta (Figs. 89J, R-S) .................... Allenhyphes (in part) CLAVES

PARA LOS

LEPTOHYPHIDAE

SUDAMERICANOS

Adultos 1

2(1) –

Ala anterior del macho con lóbulo cubitoanal muy desarrollado (ala más ancha en el 1/3 basal) (Fig. 108A); alas posteriores ausentes en ambos sexos; hembras con filamentos caudales reducidos en longitud, más cortos que el cuerpo .................. 2 Lóbulo cubitoanal normal (ala anterior más ancha en la zona media) (Fig. 90A); alas posteriores presentes o ausentes; hembras con filamentos caudales más largos que las alas anteriores ........................................................................................................ 3 Fórceps trisegmentados, largos y delgados, segmento II con base globosa (Figs. 108IS) (excepto T. australis, Fig. 108H); cercos de las hembras de una longitud similar a la del abdomen (el filamento terminal puede ser más largo) .......................... Tricorythodes Fórceps bisegmentados y cortos, el segmento II no presenta un ensanchamiento globoso basal, y está dirigido hacia afuera (Figs. 113D-K); los tres filamentos caudales de la hembra muy cortos, menos de la mitad de la longitud del abdomen ............................................................................................................................ Tricorythopsis

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Ojos del macho grandes y dorsales, divididos en una porción dorsal blanquecina y una ventral más pequeña, negruzca (Figs. 97C-D); alas posteriores ausentes en ambos sexos; fórceps 2-segmentados (Fig. 97F) .................................... Leptohyphodes Ojos del macho pequeños y negruzcos, no divididos, laterales; alas posteriores presentes al menos en los machos; fórceps 2-segmentados (Fig. 90G) o 3segmentados (Figs. 88D, 93G, 102E) ............................................................................. 4 Filamentos membranosos del mesoscutellum ausentes o no sobresalen de este esclerito; fórceps 2-segmentados (Fig. 90G); alas posteriores presentes en ambos sexos .................................................................................................................... Haplohyphes Filamentos membranosos del mesoscutellum presentes y relativamente largos; fórceps 3-segmentados (Figs. 88D, 93G, 102E); alas posteriores de la hembra presentes o ausentes .......................................................................................................... 5 Penes divididos en su mitad apical, en forma de Y o T, el ápice de cada pene con un lóbulo y una extensión aguda (Figs. 93G, M); ala posterior de la hembra ausente o presente ............................................................................................. Leptohyphes Penes completamente fusionados, si presentan ranura apical entonces con los extremos no fuertemente divergentes (Figs. 100E, 102E, 103A, 117E); ala posterior de la hembra ausente ....................................................................................... 6 Filamento terminal del macho con una espina ventral fuerte en la base (Fig. 88E); penes con una profunda escotadura media y sin grandes espinas (Fig. 88D) ........ ............................................................................................................................... Allenhyphes Filamento terminal del macho sin espina ventral, puede tener setas delgadas; penes con o sin espinas ..................................................................................................... 7 Penes con un par de largos filamentos esclerosados laterales insertados en su base (Figs. 117G, J, M) .................................................................................................... Yaurina Penes con o sin espinas esclerosadas (pueden ser hialinas), de longitud variada, pero nunca insertadas en la base de los penes (Figs. 100E, 102E, 103A) ............. 8 Segmento I de los fórceps genitales masculinos casi tan largo como el segmento II (Figs. 100E, H, K); penes con un par de espinas laterales largas, ventralmente visibles (no cubiertas por los penes) y ligeramente curvadas medialmente (Fig. 100G); gonoporo de la hembra (zona media entre esternos abdominales VII y VIII) generalmente con mácula negruzca .................................................. Lumahyphes Segmento I de los fórceps de una longitud menor a 1/2 del segmento II (Fig. 102E); penes con o sin espinas esclerosadas (pueden ser hialinas), pueden estar curvadas ventralmente, generalmente cubiertas por los penes en vista ventral ........................ 9 Penes anchos (mitad distal marcadamente más ancha que la mitad basal) y sin espinas fuertes; sin fuertes proyecciones posterolaterales en la placa estilígera ..... .............................................................................................................................. Vacupernius Penes delgados, apenas más anchos distalmente o con un ancho similar en toda su longitud; con espinas esclerosadas (a veces hialinas) de formas variadas, con (Figs. 102E, I, L) o sin (Figs. 103A, E, H) fuertes proyecciones posterolaterales en la placa estilígera ................................................................................................... Traverhyphes

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Ninfas 1

–

2(1)

– 3(2)

– 4(3)

–

5(3)

– 6(5) –

Branquias operculares subtriangulares, tocándose (o casi) en la línea media del cuerpo (Fig. 98A); branquias presentes en los segmentos abdominales II-V, lamelas ventrales con una hilera marginal de pequeños lóbulos imbricados (Figs. 99N-Q); ojos del macho divididos en una porción blanquecina y una negruzca .. ........................................................................................................................... Leptohyphodes Branquias operculares triangulares, subtriangulares u ovales, pero nunca tocándose en la línea media (Fig. 91A); branquias presentes en los segmentos abdominales II-VI, sin lóbulos pequeños imbricados (Figs. 92R-V); ojos del macho no divididos, negruzcos y pequeños ................................................................. 2 Tamaño del cuerpo pequeño (longitud total ninfa madura = 1 a 3 mm), cuerpo compacto y robusto (Figs. 114A-G); pterotecas posteriores ausentes en ambos sexos; branquia opercular ovalada con una línea transversal más clara en la zona media o subapical (Figs. 114A, 115H-I); fórmula branquial 5/4/4/2/1 (Figs. 115I-M) ............................................................................................................. Tricorythopsis Cuerpo relativamente más grande (ninfa madura > 3 mm); pterotecas posteriores presentes o ausentes; branquias operculares sin línea transversa; fórmula branquial nunca como en el caso anterior. ..................................................................................... 3 Branquias operculares triangulares o subtriangulares (Figs. 92Q, 111A-C, E), si ovales o subcuadradas (Figs. 111D, F) entonces fémures anchos (casi tan anchos como largos, Figs. 111K-N); fémures anteriores con hilera transversa de setas largas (Figs. 92F, 111I, K) ................................................................................................ 4 Branquias operculares ovales o subovales (Figs. 88M, 94A, 106A), fémures anteriores con hilera transversa de espinas de longitud variada pero nunca con setas largas (Figs. 94A, 104A) .......................................................................................... 5 Cuerpo robusto, base del abdomen claramente más ancha que el ápice (Fig. 109A); fémures anteriores solo con largas setas (Figs. 111G, I); pterotecas posteriores ausentes; fórmula branquial 3/3/3/3/2 (Figs. 110R-V) o reducciones de esta ..... ............................................................................................................................. Tricorythodes Cuerpo elongado, base del abdomen apenas más ancha que el ápice (Fig. 91A); fémures anteriores con largas setas y con cortas espinas en el margen anterior (Figs. 92F-G); pterotecas posteriores presentes; fórmula branquial 4/5-4/4/4/2 (Figs. 92R-V) ..................................................................................................... Haplohyphes Branquia opercular con espina basal (Fig. 95I); filamentos caudales con segmentos más oscuros en la base (hembras) o en la zona media (machos) (Fig. 94A); dorso de los fémures medios y posteriores con una costilla longitudinal y con una hilera transversal de espinas en la base (Fig. 95Q) ................................................ Leptohyphes Branquia opercular sin espina basal (Figs. 101R-S); filamentos caudales no como en el dilema anterior; fémures medios y posteriores sin costilla longitudinal ni hilera transversa de espinas en el dorso (Figs. 101G-H) ........................................... 6 Hilera transversal dorsal de setas espatuladas en la mitad del fémur ... Vacupernius Hilera transversal dorsal de setas espatuladas en el 1/3 distal del fémur (Figs. 101I, O, 107B, F, K, O) ............................................................................................................... 7

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Cuerpo relativamente robusto (Fig. 118A); palpo maxilar 2-segmentado (Figs. 88K, 119C); branquias operculares ovoides y relativamente pequeñas, no alcanzando la mitad posterior del segmento abdominal VII (Fig. 118A); formula branquial: 3/3/3/ 3/1-2 ..................................................................................................................................... 8 – Cuerpo relativamente elongado (Fig. 104A); palpo maxilar 2 o 3-segmentado (Figs. 101D, 105F-I); branquias operculares ovoides a subcuadradas y relativamente grandes, al menos llegando a la mitad posterior del segmento abdominal VII (Fig. 104A); formula branquial: 3/3-4/3-4/3/1-2 ................................................................. 9 8(7) Branquia abdominal VI con 2 lamelas (Fig. 88R); palpo maxilar generalmente con seta apical, márgen donde se inserta el palpo, recto o apenas cóncavo (Figs. 88KL); uña tarsal anterior con una sola hilera subapical de 4 dentículos ....................... ............................................................................................................... Allenhyphes (in part) – Branquia abdominal VI con 1 lamela (Fig. 119Q); palpo maxilar sin seta apical, palpo insertado en un márgen cóncavo (Fig. 119C); uña tarsal anterior con una sola hilera subapical de 6 a 7 dentículos (Fig.119K) ....................................... Yaurina 9(7) Palpo maxilar 3-segmentado y relativamente grande (Figs. 89J, R-S, 105I) ........ 10 – Palpo maxilar 2-segmentado y relativamente pequeño (Figs. 105F-H) .................... ..............................................................................................................Traverhyphes (in part) 10(9) Setas espatuladas en el dorso de los fémures bífidas o serradas (Figs. 101J, P); ninfa hembra madura generalmente con una mácula media negruzca entre los esternos abdominales VII y VIII .................................................................. Lumahyphes – Setas espatuladas en el dorso de los fémures con ápice liso (Figs. 89I, O, 107G); esterno de la ninfa hembra madura no como arriba ................................................ 11 11(10) Palpo maxilar con una pequeña seta apical (Fig. 105I) ............Traverhyphes (in part) – Palpo maxilar sin seta apical (Figs. 89J, R-S) ............................... Allenhyphes (in part) Genus Allenhyphes HOFMANN & SARTORI Allenhyphes HOFMANN & SARTORI, in HOFMANN et al., 1999: 67; WIERSEMA & MCCAFFERTY, 2000: 342; MOLINERI & FLOWERS, 2001: 61; MOLINERI & DOMÍNGUEZ, 2003: 272; MOLINERI, 2004: 214. (Type-species: Leptohyphes flinti ALLEN, original designation).

Taxonomy: Allenhyphes HOFMANN & SARTORI (HOFMANN et al., 1999) was established for Leptohyphes flinti ALLEN (1973b), previously known from nymphs. WIERSEMA & MCCAFFERTY (2000) transferred 7 additional South American species previously described in Leptohyphes to Allenhyphes, but a recent cladistic analysis by MOLINERI (2004) does not support these combinations. Only 1 species undoubtedly pertaining to Allenhyphes is present in South America: A. flinti was reported from adults and nymphs from Venezuela (MOLINERI & FLOWERS, 2001). Allenhyphes asperulus (ALLEN) and A. spinosus (ALLEN), both known only from nymphs, will probably prove to be members of another genus. Their generic position will be clear only when adults are known. The other species treated as Allenhyphes by WIERSEMA & MCCAFFERTY (2000) are currently included in other genera: Traverhyphes (Mocoihyphes) edmundsi (ALLEN), Traverhyphes (Byrsahyphes) nanus (ALLEN), Tricorythopsis minimus (ALLEN) (=Tricorythopsis viriosus (ALLEN)), and Yaurina ralla (ALLEN).

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A LS

MP

B

TF

C

C

D

E

VS P

F

M

G H

N I

O AS

P K

J

Q L R

Plate 88. Allenhyphes flinti, male imago. Figs. 88A-R. 88A, fore wing; 88B, hind wing; 88C, same, enlarged; 88D, male genitalia, v.v.; 88E, male terminalia, l.v. Nymph: 88F, hypopharynx, v.v.; 88G, labrum, d.v.; 88H, labium, v.v.; 88I, left mandible, d.v.; 88J, right mandible, d.v.; 88K, detail of maxillary palp; 88L, maxilla, d.v.; 88M, gill II, d.v.; 88N-88R, gills II-VI, v.v. (AS = Apical seta; C = cercus; LS = Lateral Sclerotization; MP = Median projection of styliger plate; P = Penis; TF = Terminal filament; VS = Ventral spine). Figs. 88D-E modified from MOLINERI & FLOWERS, 2001.

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Adult Characteristics: 1) Male fore wings without enlarged cubito-anal lobe (Fig. 88A); 2) hind wings present in males, with 1 longitudinal vein (Figs. 88B-C), hind wings absent in females; 3) female caudal filaments longer than wings; 4) membranous filaments of mesoscutellum long and slender; 5) male forceps 3-segmented (Fig. 88D); 6) penes fused except for deep apical furrow (Fig. 88D); 7) penes without dorsal projections near apex; 8) base of male terminal filament with well developed ventral spine (Fig. 88E). Nymphal Characteristics: 1) Maxillary palpi small, 2-segmented, with apical seta in the type species (Figs. 88K-L); 2) labrum with broad anteromedian emargination and some subapical pectinate setae on dorsum (Fig. 88G); 3) remaining mouthparts as in Figs. 88F, H-J; 4) hind wingpads present in males, absent in females; 5) operculate gill oval, without ridges (Fig. 88M); 6) gill formula: 3/3/3/3/2 (Figs. 88N-R); 7) fore femora with transverse row of long spines (Fig. 89C); 8) femora II and III without basal transverse row of spines on dorsum (Figs. 89A-B); 9) tarsal claw with numerous marginal denticles and a single submarginal row near apex (Fig. 89D). Allenhyphes and Yaurina nymphs are hard to distinguish from each other, but they can be separated because species of Yaurina have a larger maxillary palpi without an apical seta (Fig. 119C); and the type species of Allenhyphes has an apical seta (Fig. 88K, L), although this seta is not present in other species dubiously included in the genus. Additional differences are: gill VI with 1 lobe in Yaurina, but bilobated in Allenhyphes (Fig. 88R); and Allenhyphes with pectinate setae on dorsum of labrum, and Yaurina with simple setae. Distribution: Southern North America (Texas) to Northern South America (Venezuela), French Antilles. Biology: Nymphs inhabit a wide variety of rivers and streams, from sea level to 600 m. Imagos perform the nuptial flight in the morning (HOFMANN et al., 1999). KEY

TO

SOUTH AMERICAN ALLENHYPHES

Nymphs 1 – 2(1)

Maxillary palpi 2-segmented, much reduced in size (Figs. 88K-L) ..... Allenhyphes flinti Maxillary palpi 3-segmented, relatively larger (Figs. 89J, R-S) .................................. 2 Femora with large blackish marks (Fig. 89F), and with relatively long dorsal spines (Fig. 89I) ................................................................................................ Allenhyphes spinosus Femora without marks (Figs. 89L-N), and with shorter dorsal spines (Fig. 89O) . ............................................................................................................... Allenhyphes asperulus

–

CLAVE

PARA LOS

ALLENHYPHES

SUDAMERICANOS

Ninfas 1 – 2(1) –

Palpos maxilares 2-segmentados, muy reducidos en tamaño (Figs. 88K-L) ........... ...................................................................................................................... Allenhyphes flinti Palpos maxilares 3-segmentados, relativamente grandes (Figs. 89J, R-S) .............. 2 Fémures con marcas negruzcas grandes (Fig. 89F), y con espinas relativamente largas en el dorso (Fig. 89I) .............................................................. Allenhyphes spinosus Fémures sin marcas (Figs. 89L-N), y con espinas más cortas (Fig. 89O) ................ ............................................................................................................... Allenhyphes asperulus

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Allenhyphes? asperulus (ALLEN) Leptohyphes asperulus ALLEN, 1967: 366; HUBBARD, 1982a: 273. Allenhyphes asperlus [sic]; WIERSEMA & MCCAFFERTY, 2000: 343. Allenhyphes? asperulus; MOLINERI, 2004: 214.

Taxonomy: Allenhyphes? asperulus (ALLEN, 1967) was described from a single nymph. WIERSEMA & MCCAFFERTY (2000) transferred it to Allenhyphes, but this was not supported in a cladistic analysis (MOLINERI, 2004). Nevertheless, its generic position will remain uncertain until the adult stage is known. For now it is treated as Allenhyphes. Allenhyphes asperulus can be separated from the other species of this group of genera by the following combination of characters: 1) Maxillary palpi 3-segmented, without apical seta (Figs. 89RS); 2) femoral spines relatively short and with smooth apex (Fig. 89O); 3) female gonopore, in mature nymphs, without black macula; 4) operculate gills broadly shaded with gray (Fig. 89Q); 5) legs and tarsal claw as in Figs. 89L-N, P, respectively. Distribution: Peru (near Tingo María). Allenhyphes flinti (ALLEN) Leptohyphes flinti ALLEN, 1973b: 365 (nymph). Allenhyphes flinti; HOFMANN & SARTORI, in HOFMANN et al., 1999: 67 (adults, nymph, egg); WIERSEMA & MCCAFFERTY, 2000: 342; MOLINERI & FLOWERS, 2001: 61.

Taxonomy: This species was first described from nymphs in Leptohyphes by ALLEN (1973b), HOFMANN et al. (1999) reared nymphs and demonstrated that the adults do not show the characters of Leptohyphes, they then established Allenhyphes to include this species. Allenhyphes flinti (ALLEN) can be separated from the other species of Leptohyphidae by the following combinations of characters. In the imago: see generic section. In the nymph: 1) Maxillary palpi 2-segmented, generally with apical seta (Figs. 88K-L); 2) femoral spines relatively short and with smooth apex (Fig. 89E); 3) female gonopore, in mature nymphs, without black macula; 4) operculate gills broadly shaded with gray (Fig. 88M); 5) legs as in Figs. 89A-C, fore tarsal claw with 2 sets of denticles (Fig. 89D). Distribution and Biology: French Antilles (Martinique, Dominica, Guadeloupe), Panama, Venezuela. HOFMANN et al. (1999), reported that this multivoltine species is commonly encountered in a wide variety of rivers and streams in Guadeloupe Island at altitudes ranging from 5 m to 600 m. The imagos were active at early morning. Allenhyphes? spinosus (ALLEN & ROBACK) Leptohyphes sp. 1 ROBACK, 1966: 150. Leptohyphes spinosus ALLEN & ROBACK, 1969: 372; HUBBARD, 1982a: 274. Allenhyphes spinosus; WIERSEMA & MCCAFFERTY, 2000: 366. Allenhyphes? spinosus; MOLINERI (2004): 214.

Taxonomy: Allenhyphes spinosus was described first in Leptohyphes (ALLEN & ROBACK, 1969) and later transferred to Allenhyphes (WIERSEMA & MCCAFFERTY, 2000). It is known from 2 nymphs collected in a locality near the type locality of A. asperulus. Both species are similar and, as

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explained before, the generic placement of A. spinosus is uncertain pending discovery of male adults. The nymphs of A. spinosus can be distinguished from the others of the group by the following combination of characters: 1) Maxillary palpi 3-segmented, without apical seta (Fig. 89J); 2) femoral spines relatively long and with smooth apex (Fig. 89I); 3) female gonopore, in A B C D E F G

J H

I K L

R

O

S

Q M

N

P

Plate 89. Allenhyphes, nymphs. Figs. 89A-S. Allenhyphes flinti: 89A, hind leg; 89B, midle leg; 89C, fore leg; 89D, detail of tarsal claw; 89E, detail of fore femoral spines. Allenhyphes spinosus: 89F, hind leg; 89G, fore leg; 89H, detail of tarsal claw; 89I, detail of fore femoral spines; 89J, maxilla, d.v.; 89K, gill II, d.v. Allenhyphes asperulus: 89L, hind leg; 89M, middle leg; 89N, fore leg; 89O, detail of fore femoral spines; 89P, detail of tarsal claw; 89Q, gill II, d.v; 89R, detail of maxillary palp; 89S, maxilla, d.v. Figs. 89A-S modified from MOLINERI, 2004.

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mature nymphs, without black macula; 4) operculate gills (Fig. 89K) and hind femora (Fig. 89F) with blackish marks; 5) tarsal claw with a double palisade of submarginal denticles (Fig. 89H). Distribution: Peru (near Tingo María). Genus Haplohyphes ALLEN Haplohyphes ALLEN, 1966: 566; DOMÍNGUEZ, 1984: 104; WIERSEMA & MCCAFFERTY, 2000: 343. (Type-species: Haplohyphes huallaga ALLEN, original designation).

Taxonomy: Haplohyphes is known from all the stages and currently includes 6 species: Haplohyphes aquilonius LUGO-ORTIZ and MCCAFFERTY (1995d), H. huallaga ALLEN (1966), H. mithras (TRAVER, 1958a), H. furtiva DOMÍNGUEZ (1984), H. dominguezi MOLINERI (1999a), and H. yanahuicsa MOLINERI (2003b). Adult Characteristics: 1) Male fore wings without well developed cubito-anal lobe (Fig. 90A); 2) hind wings present in both sexes, with 1-3 longitudinal veins; 3) female caudal filaments longer than fore wings; 4) membranous filaments of mesoscutellum absent; 5) male forceps 2-segmented (Fig. 90L); 6) penes divided in apical 1/2 or less, apical lobes of penes not strongly divergent (Fig. 90G); 7) male gonopore associated with a distal spine; 8) penes without dorsal projection. Nymphal Characteristics: 1) Hind wing pads present in both sexes; 2) operculate gill subtriangular, with 2 elevated ridges (Fig. 92Q); 3) gill formula: 4/5/4-5/4/2, lobes subtriangular; 4) fore femora with transverse row of long setae (Fig. 92F) and with stout spines on leading edge (Fig. 92G); 5) femora II and III with transverse row of dorsal setae at base (Figs. 92D-E); 6) maxillary palpi small, 2-segmented with apical seta (Fig. 92O); 7) labrum with deep anteromedian emargination and with numerous thick spines dorsally (Fig. 92L); 8) tarsal claw with numerous marginal denticles and a double row of submarginal denticles near apex (Fig. 92H); 9) body covered with long setae, abdomen cylindrical and long (Fig. 91A); 10) frontal and genal projections present. Distribution: South and Central America (from Costa Rica to NW Argentina). Biology: Nymphs of this genus are generally found in small to medium-sized pristine mountain streams (2-10 m wide), inhabiting sandy and stony substrates. Nymphs are mainly crawlers and rarely swim. The nuptial flight occurs early in the morning, with some males flying until noon. Male imagos perform the typical up and down movements except when windy; then, they hold position or alight on vegetation. The oblong or spherical swarms are composed of 20-80 males, and are located 2-4 m above riverbanks or water. KEYS

TO

HAPLOHYPHES

Adults 1 – 2(1) –

Posterior segments of abdomen blackish, much darker than rest of body (similar to Fig. 91A); genitalia and hind wings as in Figs. 90Q-S and P ............... Haplohyphes yanahuicsa Coloration of abdomen often darker posteriorly but never as above .................... 2 Apical spines of penes as long as or longer than apex of membranous lobes (Fig. 90I); hind wings as in Fig. 90H .................................................. Haplohyphes dominguezi Apical spines of penes shorter than apex of membranous lobes (Figs. 90G, M) ...... 3

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Hind wings with 1 longitudinal vein (Fig. 90J) ........................... Haplohyphes huallaga Hind wings with 2 or 3 longitudinal veins (Figs. 90C, F, K, N-O) ......................... 4 Membranous lobes of penes rounded (similar to Fig. 90I); hind wings with 2 longitudinal veins (Fig. 90K) ............................................................. Haplohyphes mithras Membranous lobes of penes obliquely truncated (Figs. 90G, M); hind wings with 3 longitudinal veins (Figs. 90C, F, N-O) ....................................................................... 5

A

D E B

F

C

H

G I L

J

K M N

O P

Q

S R

Plate 90. Haplohyphes, adults. Figs. 90A-S. H. aquilonius: 90A, male fore wing; 90B, male hind wing; 90C, idem, detail; 90D, female fore wing; 90E, female hind wing; 90F, idem, detail; 90G, male genitalia, v.v. H. dominguezi: 90H, male hind wing, detail; 90I, male genitalia, v.v. H. huallaga: 90J, male hind wing, detail. H. mithras: 90K, male hind wing, detail. H. furtiva: 90L, male forceps, detail; 90M, male genitalia, v.v.; 90N, male hind wing, detail; 90O, female hind wing, detail. H. yanahuicsa: 90P, male hind wing, detail; 90Q, genitalia, v.v.; 90R, male forceps, detail; 90S, genitalia, l.v. Figs. 90A-G, J-S modified from MOLINERI, 2003b; 90H-I from MOLINERI, 1999a.

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Segment I of forceps as long as or longer than segment II (Fig. 90L); penes divided on apical 0.23-0.26 (Fig. 90M) ............................................. Haplohyphes baritu Segment I of forceps slightly shorter than segment II (Fig. 90G); penes divided on apical 0.34-0.40 (Fig. 90G) ............................................................ Haplohyphes aquilonius

Nymphs 1

–

Apex of abdomen blackish, much darker than the rest of body (Fig. 91A) .......... ............................................................................................................ Haplohyphes yanahuicsa Coloration not as above .................................................................................................... 2 Fore wingbuds shaded with black only at basal half of costal margin .................... ................................................................................................................... Haplohyphes baritu Fore wingbuds shaded with black along entire costal margin ....... Haplohyphes aquilonius

CLAVES

PARA

– 2(1)

HAPLOHYPHES

Adultos 1 – 2(1) – 3(2) – 4(3) – 5(4) –

Últimos segmentos abdominales negruzcos, mucho más oscuros que el resto del cuerpo (Fig. 91A); genitalia y alas posteriores como en las Figs. 90Q-S y P, respectivamente .............................................................................. Haplohyphes yanahuicsa Coloración del abdomen a veces oscureciéndose posteriormente pero nunca como arriba ......................................................................................................................... 2 Espinas apicales de los penes tan o más largas que los lóbulos membranosos (Fig. 90I); alas como en las Fig. 90H .................................................. Haplohyphes dominguezi Espinas apicales de los penes más cortas que los lóbulos membranosos (Figs. 90G-M) ................................................................................................................................. 3 Alas posteriores con 1 vena longitudinal (Fig. 90J) ................... Haplohyphes huallaga Alas posteriores con 2 ó 3 venas longitudinales (Figs. 90C, F, K, N-O) ............... 4 Lóbulos membranosos de los penes redondeados (similar a Fig. 90I); alas posteriores con 2 venas longitudinales (Fig. 90K) ....................... Haplohyphes mithras Lóbulos membranosos de los penes oblicuamente truncados (Figs. 90G, M); alas posteriores con 3 venas longitudinales (Figs. 90C, F, N-O) ..................................... 5 Segmento I de los fórceps tan o más largo que el segmento II (Fig. 90L); penes divididos en los 0,23-0,26 apicales (Fig. 90M) ................................ Haplohyphes baritu Segmento I de los fórceps apenas más corto que el segmento II (Fig. 90G); penes divididos en los 0,34-0,40 apicales (Fig. 90G) .......................... Haplohyphes aquilonius

Ninfas 1 – 2(1) –

Extremo del abdomen negruzco, mucho más oscuro que el resto del cuerpo (Fig. 91A) ................................................................................................... Haplohyphes yanahuicsa Coloración no como arriba .............................................................................................. 2 Pterotecas anteriores sombreadas con negruzco solo en la mitad basal del margen costal ........................................................................................................ Haplohyphes baritu Pterotecas anteriores sombreadas con negruzco en todo el margen costal ............ .............................................................................................................Haplohyphes aquilonius

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Plate 91. Haplohyphes yanahuicsa. Fig. 91A, nymphal habitus. Modified from MOLINERI, 2003b.

Haplohyphes aquilonius LUGO-ORTIZ & MCCAFFERTY Haplohyphes aquilonius LUGO-ORTIZ & MCCAFFERTY, 1995d: 166; MOLINERI, 1999a: 29; MOLINERI, 2003b: 2.

Taxonomy: Haplohyphes aquilonius was described from Costa Rican nymphs by LUGOORTIZ & MCCAFFERTY (1995d). These authors stated that these nymphs could represent the unknown nymphal stage of H. mithras (TRAVER), described from imagos, also from Costa Rica. MOLINERI (2003b) tentatively associated both stages of H. aquilonius, the adults of this species showed marked differences with H. mithras. Haplohyphes aquilonius can be distinguished from the others species of the genus by the following combination of characters. In the imago: 1) hind wings with 3 longitudinal veins (Figs. 90C, F); 2) segment

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I of forceps shorter than segment II (Fig. 90G); 3) penes divided on apical 0.34-0.40 (Fig. 90G); 4) membranous lobes of penes obliquely truncated at apex (Fig. 90G); 5) apical spines of penes shorter than membranous lobes; 6) male fore wing length: 6.2-6.5 mm; 7) abdomen uniformly shaded with gray. In the nymph: 1) Fore wing buds shaded with gray at least along entire costal and subcostal areas; 2) head shaded with gray as in Fig. 92A. Distribution and Biology: Costa Rica and Colombia. Colombian material was collected in stony mountain streams, with forested margins. Adults were attracted to light traps. Haplohyphes baritu DOMÍNGUEZ Haplohyphes baritu DOMÍNGUEZ, 1984: 106; DOMÍNGUEZ et al., 1994: 97; LUGO-ORTIZ & MCCAFFERTY, 1995d: 169; MOLINERI, 1999a: 29; MOLINERI, 2003b: 4. Haplohyphes furtiva DOMÍNGUEZ, 1984: 108; DOMÍNGUEZ et al., 1994: 98; MOLINERI, 1999a: 29; MOLINERI, 2003b: 4. Haplohyphes furtivus [sic]; LUGO-ORTIZ & MCCAFFERTY, 1995d: 169.3

Taxonomy: Haplohyphes baritu can be distinguished from the others species of the genus by the following combination of characters. In the imago: 1) hind wings with 3 longitudinal veins (Figs. 90N-O); 2) segment I of forceps as long as or longer than segment II (Fig. 90L); 3) penes divided on apical 0.23-0.26 (Fig. 90M); 4) membranous lobes of penes obliquely truncated at apex (Fig. 90M); 5) apical spines of penes shorter than membranous lobes; 6) male fore wing length: 5.0-7.2 mm; 7) abdomen uniformly shaded with gray. In the nymph: 1) Fore wingbuds shaded with gray on basal half of costal margin; 2) head shaded with gray as in Fig. 92B. Distribution and Biology: Northwestern Argentina, Southwestern Bolivia. Nymphs inhabit clean and oxygenated stony streams at altitudes ranging from 500 to 1500 m. They seem to prefer patches of coarse sand in relatively swift current. Stomach contents consist of particulate organic matter suggesting that they feed as collectors or scrapers. Subimagos emerge at dusk, molting to imagos during the night. Swarming starts at early morning, and generally ends at midmorning or noon. Swarms are composed of 20-100 males that occupy a subspheric space 2-4 m above the river margins. A 2-years study in a small stream from NW Argentina showed a low density of nymphs (21 ind/m²); mature nymphs appeared in the warm season (spring-early summer) and nymphs were completely absent from February to September. Haplohyphes dominguezi MOLINERI Haplohyphes dominguezi MOLINERI, 1999a: 29; MOLINERI, 2003b: 9.

Taxonomy: Haplohyphes dominguezi is only known from male imago and can be differentiated from the other species of the genus by the following combination of characters: 1) hind wings with 2 long and 1 short longitudinal veins (Fig. 90H); 2) segment I of forceps as long as segment II (Fig. 90I); 3) penes divided on apical 0.30 (Fig. 90I); 4) membranous lobes of penes rounded at apex (Fig. 90I); 5) apical spines of penes longer than membranous lobes; 6) male fore wing length: 5.0 mm; 7) abdomen uniformly shaded with gray. Distribution: Ecuador (Zamora, Chimborazo).

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Haplohyphes huallaga ALLEN Haplohyphes huallaga ALLEN, 1966: 567 (male, female); PUTHZ, 1973: 93; HUBBARD, 1982a: 273; MOLINERI, 2003b: 10.

Taxonomy: Haplohyphes huallaga is the type-species of the genus, known from adults of both sexes. They can be differentiated from the other species of the genus by the following combination of characters: 1) hind wings with 1 longitudinal vein (Fig. 90J); 2) segment I of forceps as long as segment II; 3) penes divided on apical 0.20-0.30; 4) membranous lobes of penes obliquely truncated at apex; 5) apical spines of penes shorter than apex of membranous lobes; 6) male fore wing length: 4.0-5.0 mm; 7) abdomen uniformly shaded with gray. Besides these characters, ALLEN (1966) also reported that in the fore wings, the veins CuP and A are fused or nearly fused at wing margin. Distribution: Peru (Tingo María). Haplohyphes mithras (TRAVER) Leptohyphes mithras TRAVER, 1958a: 497. Haplohyphes mithras; ALLEN, 1966; DOMÍNGUEZ, 1984: 103; LUGO-ORTIZ & MCCAFFERTY, 1995d: 169; MOLINERI, 2003b: 10.

Taxonomy: Haplohyphes mithras was described by TRAVER (1958a) in Leptohyphes, and ALLEN (1966) transferred it to Haplohyphes. H. mithras was described from Costa Rica, and recently was reported from Colombia. It is known from adults of both sexes and can be distinguished from the other species of the genus by the following combination of characters: 1) hind wings with 2 longitudinal veins (Fig. 90K); 2) segment I of forceps as long as or longer than segment II; 3) penes divided on apical 0.40; 4) membranous lobes of penes rounded; 5) lateral spines of penes shorter than membranous lobes; 6) male fore wing length: 3.8 mm; 7) abdomen more or less uniformly shaded with gray. Distribution: Costa Rica and Colombia. Haplohyphes yanahuicsa MOLINERI Haplohyphes yanahuicsa MOLINERI, 2003b: 5.

Taxonomy: Haplohyphes yanahuicsa MOLINERI is known from all stages and can be distinguished from the other species of the genus by the following combination of characters. In the imago: 1) Hind wings with 2 longitudinal veins (Fig. 90P); 2) segment I and II of forceps subequal in length (Fig. 90R); 3) penes divided on apical 0.22 (Fig. 90Q); 4) membranous lobes of penes rounded (Fig. 90Q); 5) apical spines of penes shorter than membranous lobes; 6) male fore wing length: 4.0 mm; 7) general coloration pale except apical segments of abdomen blackish (Fig. 91A). In the nymph: 1) Markings on head not reaching base of antennae (Fig. 92C); 2) abdomen shaded with blackish on rear segments (as in Fig. 91A). Distribution: One known locality in southern Bolivia. The type specimens were collected in a small stony stream at 500 m altitude. The watershed is covered by mountain cloud forest. Nymphs were found among rocks and sand in riffle zones. Imagos were attracted to light traps just before dawn.

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B

261

C

D F

H

E

G

I

J L

K

M

N

O

Q

R

S

T

P

U

V

Plate 92. Haplohyphes, nymphs. Figs. 92A-V. H. aquilonius: 92A, color pattern of head. H. furtiva: 92B, color pattern of head. H. yanahuicsa: 92C, color pattern of head; 92D, hind leg; 92E, middle leg; 92F, fore leg; 92G, marginal spines on fore legs, detail; 92H, tarsal claw; 92I, left mandible, d.v.; 92J, hypopharynx, v.v.; 92K, labial palp, detail; 92L, labrum, d.v.; 92M, labium, v.v.; 92N, right mandible, d.v.; 92O, maxillary palpi, detail; 92P, right maxillae, v.v.; 92Q, abdominal gill II, d.v.; 92R-V, abdominal gills II-VI, v.v. Figs. 92A-V modified from MOLINERI, 2003b.

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Genus Leptohyphes EATON Leptohyphes EATON, 1882: 208; EATON, 1883-1888: 140; NEEDHAM & MURPHY, 1924: 32; TRAVER, 1958a: 497; EDMUNDS et al., 1963: 17; ALLEN, 1967: 350; ALLEN, 1978: 541; ULMER, 1933: 206; MOLINERI, 2003a: 49. (Type-species: Leptohyphes eximius EATON, original designation). Bruchella NAVÁS, 1920e: 56; DEMOULIN, 1952d: 281; TRAVER, 1958a: 494. (Type-species: Bruchella nigra NAVÁS, original designation). Cotopaxi MAYO, 1968b: 301; WIERSEMA & MCCAFFERTY, 2000: 340; MCCAFFERTY & WANG, 2000: 58; DOMÍNGUEZ et al., 2001: 18; MOLINERI, 2003a: 51. (Type-species: Cotopaxi macuchae MAYO, original designation).

Taxonomy: Leptohyphes was established by EATON (1882) and is one of the most common genera in South America. It contains several species and is found in a great variety of habitats. It is difficult to determine species within the genus, and there are no keys to all known species. The adults possess few useful characters, and genitalia in general have few variations. Numerous species have been described from nymphs, many times utilizing only a single immature nymph. ALLEN (1967, 1978), the author of the majority of the species, did not provide a key or include adequate discussion to make a reliable determination. MOLINERI (2003a) revised the South American species of this genus, proposing some synonymies. The genus Cotopaxi MAYO (1968b) was recently proposed as a junior synonym of Leptohyphes EATON (MOLINERI, 2003a). The type species, Cotopaxi macuchae MAYO (1968b) was described for a single specimen from Ecuador, apparently a gynandromorph with teratological genitalia. Adult Characteristics: 1) Male fore wings without extended cubitoanal lobe (Figs. 93C, I, Q, U); 2) hind wings present in male, either without (Fig. 93V), or with 2 (Figs. 93F, R) to 3 longitudinal veins (Fig. 93K), in females, hind wings present (Figs. 93B, E), reduced or absent; 3) female caudal filaments longer than fore wings; 4) membranous filaments of mesoscutellum long and slender; 5) male forceps 3-segmented (Figs. 93G, N, T); 6) penes divided approximately one half from apex, apex of penes lobes divergent (45°-180°); 7) male gonopore associated with a subapical triangular spine (Fig. 93M); 8) penes without subapical dorsal projections. Nymphal Characteristics: 1) Hind wingpads present in male, absent or present in female; 2) operculate gill oval, with a dorsal ridge and, ventral lobe with a basal spine (Figs. 95H-I); 3) gill formula: 3/5-10/5-8/5-8/2-5, with oval lobes (Figs. 95I-M); 4) fore femora with a transverse row of short spines (Figs. 95P, U, W, Y); 5) dorsum of middle and hind femora with a transverse row of spines at base (Fig. 95Q); 6) maxillary palpi relatively large, 3-segmented (Figs. 95C, G); 7) labrum with broad anteromedian emargination (Fig. 95A), rest of mouthparts as in Figs. 95B, D-F; 8) tarsal claws with numerous marginal denticles and 1 submarginal subapical denticle (Figs. 96Q-T), rarely this denticle absent (Figs. 96P, U) or replaced by a row of denticles (Figs. 96O, Y-Z); 9) genal projections present, frontal projections absent; 10) caudal filaments with darker annulations basally in females; in males, darker annulations in median zone of caudal filaments. Distribution: From South to North America.

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G

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H ICu2

N

K

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ICu1

O I

P

L

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Y

Plate 93. Leptohyphes, adults. Figs. 93A-Y. L. cornutus: 93A, female fore wing; 93B, female hind wing; 93C, male fore wing; 93D, male hind wing; 93E, female hind wing, enlarged; 93F, male hind wing, enlarged; 93G, male genitalia. L. eximius: 93H, female fore wing; 93I, male fore wing; 93J, male hind wing; 93K, same, enlarged; 93L, variation in Cu venation; 93M, detail of penes, lv.v.; 93N, male genitalia, v.v. L. petersi: 93O-P, details of penes, v.v. L. peterseni: 93Q, male fore wing; 93R, male hind wing; 93S, genitalia. L. plaumanni: 93T, male genitalia, v.v. L. mollipes: 93U, male fore wing; 93V, male hind wing, enlarged. Abdominal color pattern: 93W, L. petersi (terga IV-V); 93X, L. maculatus (terga VII-X); 93Y, L. setosus (terga VII-X). (MI = Marginal intercalaries; DL = Dorsal lobe; PA = Penean arm; VS = Ventral spine). Figs. 93A-P, modified from MOLINERI, 2003a; 93Q-S ULMER, 1920a; 93U-V from NEEDHAM & MURPHY, 1924 and 93W-Y from ALLEN, 1967.

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Biology: The adults of Leptohyphes generally perform the nuptial flight at early morning, emerging as subimagos the previous day (generally at dusk). They molt to imagos during the night or early morning. The swarms are relatively small, ranging from 5 to 50 males, but some huge swarms, of thousands of males, were occasionally observed. The swarms are more or less ovoid in form, located above the water or near the shore above some landmarks. Females fly in a straight pattern directly above the swarm, until detected and grabbed by a male. Some females were observed to enter the same swarm 24 times, each time abandoning it for a moment with a different male. Males were observed to return to the swarm after each mating. Only one undescribed species from Colombia was observed to fly at dusk. Nymphs are relatively poor swimmers inhabiting rivers and streams of variable conditions. In some silty rivers they were found the most abundant component of the mayfly fauna. The darker segments on the caudal filaments function as weak points where the filaments can break. The operculate gills of each side move independently; if disturbed they are pressed against the abdomen protecting the functional gills, and after a few seconds they are lifted slightly to allow the respiratory movements of the remaining gills. Analysis of gut content only showed detritus, suggesting the idea that they are collectors of particulate matter. KEYS

TO

Adults 1 – 2(1) – 3(2) – 4(3) – 5(4) – 15

SOUTH AMERICAN LEPTOHYPHES

15

Vein ICu2 of fore wings absent (Fig. 93U); hind wings without longitudinal veins (Fig. 93V) .............................................................................................. Leptohyphes mollipes Vein ICu2 of fore wings present (Figs. 93A, C, H-I, Q); hind wings with 2 or 3 longitudinal veins (Figs. 93E, F, K, R) ........................................................................... 2 Attached marginal intercalaries present in fore wings (Figs. 93A, C); hind wings present in both sexes (Figs. 93B, D); apical lobes of penes separated by an angle of 90º (Fig. 93G) ................................................................................. Leptohyphes cornutus Attached marginal intercalaries absent (Figs. 93H-I, Q); hind wings present in male only (Figs. 93J, R); apical lobes of penes separated by an angle of less than 80º (Figs. 93M-N, O-P, S-T) ............................................................................................ 3 Two pairs of dark marks present on each abdominal terga (Fig. 93Y) .................... ................................................................................................................... Leptohyphes setosus Abdominal color pattern not as above, terga generally darker medially with medial single or double bands (Figs. 93W-X) ............................................................................ 4 Hind wings with 3 longitudinal veins (Fig. 93K) .......................... Leptohyphes eximius Hind wings with 2 longitudinal veins (Fig. 93R) ......................................................... 5 Abdominal color pattern as in Fig. 93W; apical spines of penes longer than rounded lobes (Figs. 93O-P) ............................................................... Leptohyphes petersi Abdomen and genitalia not as above ............................................................................. 6

Leptohyphes nigripunctum TRAVER (1943), described from a single male subimago, is not included in the present key because the original description and figures are not sufficient to distinguish it.

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Abdomen with blackish medial marks (Fig. 93X) ..................... Leptohyphes maculatus Abdominal color pattern not as above; genitalia as in Figs. 93S-T ........................... ...................................................................... Leptohyphes peterseni / Leptohyphes plaumanni

Nymphs 1

Conspicuous and pointed tubercles present on occiput, pronotum, mesonotum and fore- and middle coxae (Fig. 96M) .......................................... Leptohyphes cornutus – Tubercles generally absent, or if present, small and blunt (Figs. 96B, D), not as above ..................................................................................................................................... 2 2(1) Hind femora with a distal tubercle as in Figs. 95N-O ............................................... 3 – Hind femora without distal tubercles ............................................................................ 4 3(2) Hind femora relatively thin (Fig. 95N); tarsal claw without subapical submarginal denticles (Fig. 96P) ............................................................................... Leptohyphes ecuador – Hind femora relatively wide (Fig. 95O); tarsal claw with a row of 3 subapical submarginal denticles (Fig. 96Z) ................................................ Leptohyphes tuberculatus 4(2) Compound eyes of male nymphs divided (Fig. 96I) .................... Leptohyphes populus – Compound eyes of both sexes normal, undivided (Fig. 96E) ................................. 5 5(4) Tarsal claws with a row of 3 or more subapical submarginal denticles (Figs. 96O, Y) ..... 6 – Tarsal claws with 1 or no subapical submarginal denticle (Figs. 96Q, R-X) ......... 7 6(5) Tubercles present on abdominal terga (similar to Fig. 96N); denticles on tarsal claws: 3 marginal and 4 submarginal near the apex (Fig. 96O) ................. Leptohyphes carinus – Abdomen without tubercles, and tinged with black as in Fig. 96L; denticles on tarsal claws: 4 marginal and 5 submarginal near the apex (Fig. 96Y) ...... Leptohyphes setosus 7(5) Small and blunt tubercles present on mesonotum (Fig. 96D); denticles on tarsal claws: 2 marginal and 1 submarginal near the apex (Fig. 96S) ... Leptohyphes invictus – Without tubercles on mesonotum (Figs. 96C, F-H); tarsal claw with at least 3 marginal denticles on tarsal claw (Figs. 96T, X) .......................................................... 8 8(7) Spines on dorsal margin of femora II and III inserted in elevated sockets (Figs. 95V, Z) .................................................................................................................................. 9 – Spines on dorsal margin of femora II and III not inserted in conspicuous elevations as above (Figs. 95Q, S-T, X) ....................................................................... 12 9(8) Plumose setae present on dorsum of labrum .................................... Leptohyphes liniti – Without plumose setae on labrum ............................................................................... 10 10(9) Dorsal surface of middle and hind femora with 7 or less spines (Fig. 95Z); tarsal claw denticulation: 3 or 4 marginal denticles and none subapical (Fig. 96W) ........ .................................................................................................................... Leptohyphes petersi – Dorsal surface of middle and hind femora with 10 or more spines (Fig. 95V); tarsal claw denticulation: 3 or 4 marginal denticles and 1 subapical (Figs. 96T, X) ......... 11 11 (10) Dorsal surface of middle and hind femora with relatively big spines located in a median longitudinal row (Fig. 95V) .............................................. Leptohyphes jodiannae – Dorsal surface of middle and hind femora with shorter spines more scattered over the dorsal surface ................................................................... Leptohyphes plaumanni 12(8) Hind wingpads present in females ................................................... Leptohyphes tacajalo – Hind wingpads absent in females ................................................................................. 13

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13(12) Middle and hind femora with short spines on dorsal margin and without spines on ventral margin (Figs. 95S-T) ..................................................................Leptohyphes illiesi – Middle and hind femora with medium size spines on dorsal margin and short spines on ventral margin (Figs. 95Q, X) ..................................................................... 14 14(13) Blackish median macula present on abdominal terga II-IX (Fig. 96K); femora relatively wide (Figs. 95W-X); tarsal claw denticulation: 4 marginal denticles and 0 or 1 submarginal near the apex (Figs. 96U-V) ........................... Leptohyphes maculatus – Abdominal color pattern generally not as above; femora slender (Figs. 95P-Q); tarsal claw denticulation: 5 to 7 marginal denticles and 1 submarginal near the apex (Fig. 96Q) .................................................................................... Leptohyphes eximius CLAVES

PARA LOS

Adultos 1 – 2(1) – 3(2) – 4(3) – 5(4) – 6(5) –

LEPTOHYPHES

SUDAMERICANOS

16

Vena ICu2 de las alas anteriores ausente (Fig. 93U); alas posteriores sin venas longitudinales (Fig. 93V) .................................................................... Leptohyphes mollipes Vena ICu2 de las alas anteriores presente (Figs. 93A, C, H-I, Q); alas posteriores con 2 ó 3 venas longitudinales (Figs. 93E, F, K, R) ................................................... 2 Intercalares marginales unidas a las venas principales presentes en alas anteriores (Figs. 93A, C); alas posteriores presentes en ambos sexos (Figs. 93B, D); brazos peneanos separados por un ángulo de 90º (Fig. 93G) ................ Leptohyphes cornutus Intercalares marginales ausentes (Figs. 93H-I, Q); alas posteriores presentes en machos solamente (Figs. 93J, R); brazos peneanos separados por un ángulo menor a 80º (Figs. 93M-P, S-T) .................................................................................................... 3 Dos pares de marcas oscuras en cada tergo abdominal (Fig. 93Y) ....... Leptohyphes setosus Coloración abdominal no como arriba, generalmente con marcas oscuras mediales, simples o dobles (Figs. 93W-X) ..................................................................... 4 Alas posteriores con 3 venas longitudinales (Fig. 93K) .............. Leptohyphes eximius Alas posteriores con 2 venas longitudinales (Fig. 93R) ............................................. 5 Coloración abdominal como en la Fig. 93W; espinas apicales de los penes más largas que los lóbulos membranosos (Figs. 93O-P) ....................... Leptohyphes petersi Abdomen y genitalia no como arriba ............................................................................ 6 Abdomen con marcas medias negruzcas (Fig. 93X) ................ Leptohyphes maculatus Abdomen no como arriba; genitalia como en las Figs. 93S-T ................................... ...................................................................... Leptohyphes peterseni / Leptohyphes plaumanni

Ninfas 1 –

16

Tubérculos conspicuos y aguzados presentes en occipucio, pronoto, mesonoto y coxas I-II (Fig. 96M) .......................................................................... Leptohyphes cornutus Tubérculos generalmente ausentes, si presentes son pequeños y romos (Figs. 96B, D), no como en el caso anterior ..................................................................................... 2 Leptohyphes nigripunctum TRAVER (1943), conocida solo del subimago macho, no fue incluida en la clave porque la descripción y las figuras originales no son suficientes para distinguirla.

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Fémures posteriores con un tubérculo distal como en las Figs. 95N-O ............... 3 Fémures posteriores sin tubérculo distal ...................................................................... 4 Femures posteriores relativamente delgados (Fig. 95N); uña tarsal sin dentículos subapicales submarginales (Fig. 96P) ............................................... Leptohyphes ecuador – Femures posteriores relativamente anchos (Fig. 95O); uña tarsal con una hilera de 3 dentículos subapicales submarginales (Fig. 96Z) ................ Leptohyphes tuberculatus 4(2) Ojos compuestos de los machos divididos (Fig. 96I) .................. Leptohyphes populus – Ojos compuestos normales en ambos sexos, no divididos (Fig. 96E) ................... 5 5(4) Uñas tarsales de todas las patas con una hilera de 3 o más dentículos subapicales submarginales (Figs. 96O, Y) ........................................................................................... 6 – Uñas tarsales con 1 o ningún dentículo subapical submarginal (Figs. 96Q-X) .... 7 6(5) Tergos abdominales con tubérculos medianos (similar a Fig. 96N); dentículos en las uñas tarsales: 3 marginales y 4 submarginales subapicales (Fig. 96O) ................ ...................................................................................................................Leptohyphes carinus – Abdomen sin tubérculos y con marcas negruzcas como en la Fig. 96L; dentículos en las uñas tarsales: 4 marginales y 5 submarginales subapicales (Fig. 96Y) ........... ................................................................................................................... Leptohyphes setosus 7(5) Mesonoto con un par de tubérculos romos pequeños (Fig. 96D); dentículos de la uña tarsal: 2 marginales y 1 submarginal subapical (Fig. 96S) .... Leptohyphes invictus – Mesonoto sin tubérculos (Figs. 96C, F-H); al menos con 3 dentículos marginales en las uñas tarsales (Figs. 96T, X) ................................................................................... 8 8(7) Espinas de los fémures medios y posteriores insertadas en bases elevadas (Figs. 95V, Z) .................................................................................................................................. 9 – Espinas de los fémures medios y posteriores no insertadas en bases elevadas conspicuas (Figs. 95Q, S-T, X) ...................................................................................... 12 9(8) Setas plumosas en el dorso del labro ................................................... Leptohyphes liniti Sin setas plumosas en el dorso del labro .................................................................... 10 10(9) Dorso de los fémures medios y posteriores con 7 o menos espinas (Fig. 95Z); denticulación de las uñas tarsales anteriores: 3 o 4 dentículos marginales y ningún submarginal (Fig. 96W) ......................................................................... Leptohyphes petersi – Dorso de los fémures medios y posteriores con 10 o más espinas (Fig. 95V); denticulación de las uñas tarsales anteriores: 3 o 4 marginales y 1 submarginal (Figs. 96T, X) ..................................................................................................................... 11 11 (10) Dorso de los fémures medios y posteriores con espinas relativamente grandes, ubicadas en una hilera mediolongitudinal dorsal (Fig. 95V) .... Leptohyphes jodiannae – Dorso de los fémures medios y posteriores con espinas relativamente más cortas y diseminadas ................................................................................... Leptohyphes plaumanni 12(8) Pterotecas posteriores presentes en hembras ................................ Leptohyphes tacajalo – Pterotecas posteriores ausentes en hembras .............................................................. 13 13(12) Fémures medios y posteriores con espinas cortas en el margen dorsal y sin espinas en el margen ventral (Figs. 95S-T) .......................................................Leptohyphes illiesi – Fémures medios y posteriores con espinas medianas en el margen dorsal y espinas cortas en el margen ventral (Figs. 95Q, X) ................................................................ 14

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14(13) Tergos abdominales II-IX generalmente con una mácula media negruzca (Fig. 96K); fémures relativamente anchos (Figs. 95W-X); denticulación de las uñas tarsales: 4 marginales y 0 o 1 submarginal subapical (Figs. 96U-V) .......................... .............................................................................................................. Leptohyphes maculatus – Coloración abdominal generalmente no como en el dilema anterior; fémures más delgados (Figs. 95P-Q); denticulación de las uñas tarsales: 5 a 7 marginales y 1 submarginal subapical (Fig. 96Q) .................................................... Leptohyphes eximius

A

Plate 94. Leptohyphes eximius. Fig. 94A, nymphal habitus. Drawing by C. Molineri.

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Leptohyphes carinus ALLEN Leptohyphes carinus ALLEN, 1973b: 365; HUBBARD, 1982a: 273; MOLINERI, 2003a: 59.

Taxonomy: Only the nymphal stage is known, and can be separated from the other species of the genus by the following combination of characters: 1) Abdomen with dorsal tubercles (similar to Fig. 96N); 2) tarsal claws with 3 marginal denticles and 4 submarginal denticles near apex (Fig. 96O); 3) spines on dorsal surface of middle and hind femora absent. Distribution: Peru (Tingo María). Leptohyphes cornutus ALLEN Leptohyphes cornutus ALLEN, 1967: 357 (nymph); HUBBARD, 1982a: 273; MOLINERI, 2003a: 55 (adults).

Taxonomy: Leptohyphes cornutus was originally described from a very immature nymph (ALLEN, 1967) and later MOLINERI (2003a) described adults of both sexes and presented SEM photographs of the eggs. This species can be differentiated from the other species of the genus by the following combination of characters, in the imagos: 1) Membrane of wings tinged with yellowish brown; 2) wing veins yellowish brown shaded with black; 3) head widely shaded with black; 4) abdominal segments IX-X and base of tails bright orange-brown in vivo; 5) genitalia as in Fig. 93G; 6) base of CuP present and parallel to base of CuA (Figs. 93A, C); 7) attached marginal intercalaries of fore wings present (Figs. 93A, C). Nymph: 1) Occiput widely washed with black; 2) tarsal claws with 9-11 irregular marginal denticles and 1 unpaired submarginal denticle near apex; 3) head, pronotum and mesonotum with pairs of tubercles as in Fig. 96M; 4) operculate lamellae widely shaded with black; 5) fore- and middle coxae with a dorsal projection, projection larger on middle coxa; 5) maxillary palp with setae on apex of segment I; 6) gill formula 3/10/8/8/5. Distribution and Biology: Argentina (Misiones) and Brazil (Paraná, Rio de Janeiro, Santa Catarina). The nymphs were found on a substrate composed mainly by a rock floor covered with submerged vegetation. Nuptial flight was observed before sunrise from 5:30 to 6:30 a.m. Swarms were small (7-10 males) and located about 1 m above surface of water, in riffle zones. Each male made a vertical zig-zag flight of about 40-60 cm, and females displayed a straight flight above the swarm. Leptohyphes ecuador MAYO Leptohyphes ecuador MAYO, 1968b: 305; HUBBARD, 1982a: 273; MOLINERI, 2003a: 61.

Taxonomy: Only nymphs are known. This species can be separated from the other species of the genus by the following combination of characters: 1) Occiput with a pair of blunt tubercles, pro- and mesonotum each with a pair of smaller tubercles (Fig. 96B), all tubercles with spicules on apex; 2) inner margin of fore wingpads with a row of a few spicules; 3) hind wingpads absent in female; 4) tarsal claws with 5 marginal denticles but without submarginal denticles near apex (Fig. 96P); 5) hind femora with a distal tubercle (Fig. 95N); 6) spines on dorsal surface of femora absent; 7) elevated sockets absent. Distribution: Ecuador (Cotopaxi).

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Leptohyphes eximius EATON Leptohyphes eximius EATON, 1882: 208 (female); EATON, 1884: 140; ULMER 1920c: 122; NEEDHAM & MURPHY, 1924: 32; NAVÁS; 1930e: 130; LESTAGE, 1931b: 60; TRAVER, 1958a: 497; TRAVER, 1958b: 81; KIMMINS, 1960: 305; EDMUNDS et al., 1976: 254; HUBBARD, 1982a: 273; DOMÍNGUEZ, 1984: 103; KLUGE & NARANJO, 1990; KLUGE, 1992a: 6-9 (nymph); HOFMANN et al., 1999: 66; MOLINERI, 2001b: 129; MOLINERI, 2003a: 52 (male). Leptohyphes bruchi NAVÁS, 1931b: 322; HUBBARD, 1982a: 273; DOMÍNGUEZ, 1984: 103; MOLINERI, 2003a: 52. Bruchella nigra NAVÁS, 1920e: 56; DEMOULIN, 1952d: 281; TRAVER, 1958a: 494. Leptohyphes nigra; EDMUNDS et al,. 1963: 17. Leptohyphes niger; HUBBARD, 1982a: 274; DOMÍNGUEZ, 1984: 103; MOLINERI, 2003a: 52.

Taxonomy: Leptohyphes eximius is the type species of the genus, and was first described from a female adult by EATON (1882). After more than a century KLUGE (1992a) described the probable nymphs and later MOLINERI (2003a) described for the first time the male imago. This species can be differentiated from the other species of the genus by the following combination of characters, in the imagos: 1) Membrane of wings almost hyaline; 2) wing veins yellowish; 3) occiput whitish-yellow with black markings similar to Fig. 96E; 4) abdominal segments IX-X and base of caudal filaments without bright orange-brown coloration in vivo. Nymph: 1) Occiput whitish or with an arc-shaped blackish mark as in Fig. 96E; 2) tarsal claws with 5-7 marginal denticles and 1 unpaired submarginal denticle near apex (Fig. 96Q); 3) basal half of operculate lamellae shaded with black; 4) gill formula 3/6/6/6/2 (Figs. 95I-M); 6) basal 1/3 of caudal filaments with whorls of long setae at each articulation. Distribution and Biology: Western Argentina and Bolivia. Leptohyphes eximius is a very common species encountered in a wide variety of streams and rivers at altitudes ranging from 300 to 2300 m. The species can tolerate low levels of dissolved oxygen and relatively high levels of suspended solids. Nymphs of different sizes and adults are found throughout the year, except at the upper altitudinal limits, where adults are rare in the coldest months. Adults from different seasons vary in size, those growing as nymphs in the winter being 2 times bigger than those of the faster summer generations. The nymphs prefer the rocks in riffles, but also are found on other substrates such as plants, algae and submerged wood. Leptohyphes illiesi ALLEN Leptohyphes illiesi ALLEN, 1967: 368; HUBBARD, 1982a: 273; MOLINERI, 2003a: 61.

Taxonomy: Only the nymph is known and it can be separated from the other species of the genus by the following combination of characters: 1) Tarsal claws with 4-5 marginal denticles and 1 submarginal denticle near apex (Fig. 96R); 2) hind wingpads absent in female; 3) developing wings whitish; 4) spines on dorsal surface of middle and hind femora present near dorsal margin (Figs. 95S-T); 5) elevated sockets on femora absent; 6) anterolateral corners of pro- and mesonotum projected as in Fig. 96C; 7) occiput apparently without color markings. Distribution: Peru.

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Plate 95. Leptohyphes, nymphs. Figs. 95A-Z. L. eximius: 95A, labrum; 95B, hypopharynx; 95C, maxillae; 95D, right mandible; 95E, left mandible; 95F, labium. L. setosus: 95G, maxilla. L. eximius: 95H, gill II, d.v.; 95I-M, gills II-VI, v.v. 95N, L. ecuador, hind leg; 95O, L. tuberculatus, hind leg; 95P-Q, fore and hind leg, L. eximius; 95R-T, fore, middle and hind legs, L. illiesi; 95U-V, fore and hind legs, L. jodiannae; 95W-X: fore and hind legs, L. maculatus; 95Y-Z, fore and hind legs, L. petersi. (BR = Basal row; BS = Basal spine; TR = Transversal row). Figs. 95A-Z modified from MOLINERI, 2003a.

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Leptohyphes invictus ALLEN Leptohyphes invictus ALLEN, 1973b: 368; HUBBARD, 1982a: 274; MOLINERI, 2003a: 61.

Taxonomy: This species, known only from nymphs, can be differentiated from the other species of the genus by the following combination of characters: 1) Denticles on tarsal claws: 2 marginal and 1 submarginal subapical (Fig. 96S); 2) small tubercles on mesonotum (Fig. 96D); 3) anterolateral corners of pro- and mesonotum projected as in Fig. 96D; 4) occiput apparently without color markings (Fig. 96D). Distribution: Peru (Tingo María). Leptohyphes jodiannae ALLEN Leptohyphes jodiannae ALLEN, 1967: 358; HUBBARD, 1982a: 274; MOLINERI, 2003a: 61.

Taxonomy: The nymphs of this species (only stage known) can be separated from the other species of the genus by the following combination of characters: 1) Denticles on tarsal claws: 3 marginal and 1 submarginal subapical (Fig. 96T); 2) mesonotum with spicules; 3) mesonotum with a small acute projection on anterolateral corners (Fig. 96F); 4) hind wingpads absent in female; 5) developing wings whitish; 6) femoral spines relatively long and located on elevated sockets (Figs. 95U-V); 7) spines on anterior face of middle and hind femora present (as a median longitudinal row of 12-13 spines, Fig. 95V); 8) inner margin of fore tibiae somewhat expanded (Fig. 95U). Distribution: Peru (Tingo María). Leptohyphes liniti WANG, SITES & MCCAFERTY Leptohyphes liniti WANG, SITES & MCCAFERTY, 1998: 69; MOLINERI, 2003a: 61.

Taxonomy: This species was described from nymphs and can be distinguished by: 1) Head, body and legs without tubercles; 2) tarsal claw: 4-5 marginal and without submarginal subapical denticles; 3) developing wings blackish; 4) elevated sockets present; 5) anterolateral corners of pronotum projected anteriorly; 6) occiput with a reticulated color pattern; 7) branched setae present on dorsum of labrum. Distribution: Ecuador. Leptohyphes maculatus ALLEN Leptohyphes sp. 3 ROBACK [partim], 1966: 151. Leptohyphes maculatus ALLEN, 1967: 360; ALLEN & ROBACK, 1969: 376; HUBBARD, 1982a: 274; MOLINERI, 2003a: 63. Leptohyphes sp. ILLIES, 1964. Leptohyphes comatus ALLEN, 1967: 368; HUBBARD, 1982a: 273; MOLINERI, 2003a: 63. Leptohyphes sp. 4 ROBACK , 1966: 151. Leptohyphes hirsutus ALLEN & ROBACK, 1969: 374; HUBBARD, 1982a: 273; MOLINERI, 2003a: 63. Leptohyphes sp. 2 ROBACK , 1966: 151. Leptohyphes myllonotus ALLEN & ROBACK, 1969: 375; HUBBARD, 1982a: 274; MOLINERI, 2003a: 63.

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Taxonomy: Molineri (2003a) proposed a list of synonyms, all of which were described from the same locality. Leptohyphes maculatus is known from subimagos and nymphs, and can be distinguished from the other species of the genus by the following combination of characters. In the adults: 1) Female hind wings absent; 2) membrane of wings not strongly tinged with brown or black; 3) abdominal color pattern as in Fig. 93X. In the nymphs: 1) Body and legs completely covered with thin whitish setae; 2) occiput with variable marks; 3) tarsal claw denticulation: 3-5 marginal and 0-1 submarginal subapical denticles (Figs. 96U-V); 4) developing wings whitish. Distribution: Peru (Tingo María). Leptohyphes mollipes NEEDHAM & MURPHY Leptohyphes mollipes NEEDHAM & MURPHY, 1924: 32; LESTAGE, 1931b: 60; TRAVER, 1958a; HUBBARD, 1982a: 274; MOLINERI, 2003a: 63.

Taxonomy: This species is known from adults of both sexes. The characterization of this species is difficult but following original drawings (NEEDHAM & MURPHY, 1924): 1) Male genitalia seems typical of the genus; 2) in fore wings, vein ICu2 absent (Fig. 93U); and 3) hind wings without veins (Fig. 93V). Distribution: Brazil (Cordisboro). Leptohyphes nigripunctum TRAVER Leptohyphes nigripunctum TRAVER, 1943: 82 (male); HUBBARD, 1982a: 274; MOLINERI, 2003a: 63.

Taxonomy: Leptohyphes nigripunctum was described from a male subimago. The genitalia was damaged during the process of mounting (TRAVER, 1943) but appears typical for the genus. Other characters given by TRAVER (1943) are: head and thorax grayish, abdominal segments II-VI yellowish and remaining segments reddish brown. The length of the body and of the fore wings is 4 mm. Distribution: Venezuela (Antimano, 900 m) and Mexico. Leptohyphes peterseni ULMER “Genus Nov” ESBEN-PETERSEN, 1909: 553. Leptohyphes peterseni ULMER, 1920a: 46; ULMER, 1920c: 122; LESTAGE, 1924c: 45; NEEDHAM & MURPHY, 1924: 32; LESTAGE, 1931b: 60; HUBBARD, 1982a: 274; DOMÍNGUEZ, 1984: 103; MOLINERI, 2003a: 63.

Taxonomy: This species is known from subimagos of both sexes. The male genitalia (Fig. 93S) drawn by ULMER (1920a) is quite similar to that described for the males of L. plaumanni (Fig. 93T), and at the moment both species are indistinguishable. Distribution: Brazil (Santa Catarina), Argentina and Bolivia. ULMER (1920a) also cited this species from Central and North America, but this probably is due to a misidentification.

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Plate 96. Leptohyphes, nymphs. Figs. 96A-Z. 96A, head and pronotum, L. carinus; 96B, head and thorax, L. ecuador; 96C, pro- and mesonota, L. illiesi; 96D, head and thorax, L. invictus; 96E, head, L. eximius; 96F, head and thorax, L. jodiannae; 96G, head and thorax, L. tacajalo; 96H, pro- and mesonota, L. petersi; 96I, head, L. populus. Abdominal color pattern: 96J, L. petersi (terga IV-V); 96K, L. maculatus (terga VII-X); 96L, L. setosus (terga VII-X); 96M, body, l.v., L. cornutus; 96N, abdomen, l.v., L. tuberculatus. Tarsal claws: 96O, L. carinus; 96P, L. ecuador; 96Q, L. eximius; 96R, L. illiesi; 96S, L. invictus; 96T, L. jodiannae; 96U-V, L. maculatus; 96W, L. petersi; 96X, L. plaumanni; 96Y, L. setosus; 96Z, L. tuberculatus. Figs. 96A, C-I and 96OZ modified from MOLINERI, 2003a; 96B MAYO, 1968b and 96J-L, N from ALLEN, 1967.

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Leptohyphes petersi ALLEN Leptohyphes nymph no. 2 NEEDHAM & MURPHY, 1924: 34. Leptohyphes petersi ALLEN, 1967: 364; HUBBARD, 1982a: 274; MOLINERI, 2003a: 64.

Taxonomy: Leptohyphes petersi can be separated from the other species of the genus by the following combinations of characters. In the adults: 1) Female hind wings absent; 2) penes as in Figs. 93O-P, distal spines longer than inner membranous lobes; 3) membrane of wings not strongly tinged with black or brown; 4) hind wings with 2 longitudinal veins; 5) abdominal color pattern as in Fig. 93W. In the nymphs: 1) Femoral spines of moderate length (Figs. 95Y-Z); 2) elevated sockets present on femora (Figs. 95Y-Z); 3) tarsal claw denticulation: 3-4 marginal and without submarginal subapical denticles (Fig. 96W); 4) maxillary palpi long and slender; 5) hind wingpads absent in female; 6) developing wings blackish or whitish; 7) pro- and mesonotum with projected anterolateral corners (Fig. 96H). Distribution: Peru. Leptohyphes plaumanni ALLEN Leptohyphes sp. EDMUNDS et al., 1963. Leptohyphes plaumanni ALLEN, 1967: 355 (nymph); HUBBARD, 1982a: 274; DA-SILVA, 1993b: 314; MOLINERI, 2003a: 57 (male, female). Leptohyphes pereirae DA-SILVA, 1993b: 313; MOLINERI, 2003a: 57.

Taxonomy: Leptohyphes plaumanni was first described from nymphs (ALLEN, 1967) and later MOLINERI (2003a) described the adults of both sexes. It can be differentiated from the other species of the genus by the following combination of characters. In the imagos: 1) Membrane of wings tinged with yellowish light brown; 2) wing veins grayish-brown; 3) occiput broadly shaded with black; 4) abdominal segments IX-X and base of caudal filaments bright orange-brown in vivo; 5) genitalia as in Fig. 93T. In the nymph: 1) Occiput widely washed with black; 2) tarsal claws with 3-4 marginal denticles and 1 submarginal denticle near apex (Fig. 96X); 3) operculate lamellae widely shaded with gray; 4) basal 1/3 of caudal filaments with whorls of short spines at each intersegmental joint; 5) gill formula 3/5-6/5-6/5/1. Distribution and Biology: Argentina (Misiones) and Brazil (Rio de Janeiro, Nova Teutonia). DA-SILVA (1993b) reported that the nymphs were collected from sandy bottoms or deposits of allochthonous organic material in marginal areas of a stream with a width between 2 and 4 m; and that the water temperatures at times of collection were 18°C and 25°C. Nymphs collected in Argentina (MOLINERI, 2003a) were found in the stony substrate of warm streams at low altitude (300-500 m). Adults were captured in the nuptial flight during early morning. Swarms occurred 2-3 m above water level, and were composed of 20-30 males flying in a typical up and down pattern. Leptohyphes populus ALLEN Leptohyphes populus ALLEN, 1973b: 366; HUBBARD, 1982a: 274; MOLINERI, 2003a: 64.

Taxonomy: This species is only known from the nymphal stage and can be distinguished from the other species of the genus by the following combination of characters: 1) Eyes of

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male divided (Fig. 96I); 2) without tubercles on body or legs; 3) tarsal claw denticulation: 3-4 marginal and without submarginal subapical denticles; 4) spines on dorsal surface of femora absent; 5) elevated sockets present on femora. This is the only species of Leptohyphes known to have divided eyes in the male, situation that should facilitate its identification. Distribution: Brazil (Amazonas). Leptohyphes setosus ALLEN Leptohyphes sp. 3 ROBACK [partim], 1966: 151. Leptohyphes setosus ALLEN, 1967: 362; ALLEN & ROBACK, 1969: 378; HUBBARD, 1982a: 274; MOLINERI, 2003a: 64. Leptohyphes sp. 5 ROBACK, 1966: 151. Leptohyphes echinatus ALLEN & ROBACK, 1969: 375; HUBBARD, 1982a: 273; MOLINERI, 2003a: 64.

Taxonomy: Leptohyphes setosus is known from male subimagos and nymphs and it can be distinguished from the others of the genus by the following combination of characters. In the adults: 1) Genitalia of “peterseni” type; no more data except for the abdominal color pattern (Fig. 93Y). In the nymphs: 1) Body and legs completely covered with thin and whitish setae; 2) without projections on legs or abdomen; 3) maxillae stout with enlarged apical canines, recurved setae of distal brush short and scarce, dentisetae small and reduced in number, palpi short and robust (Fig. 95G); 4) tibiae wide, ventral margin extended; 5) tarsal claw denticulation: 3-4 marginal and 4-5 submarginal subapical denticles (Fig. 96Y); 6) femora wide, femoral spines relatively short; 7) hind wingpads absent in females; 8) abdominal color pattern as in Fig. 96L. Distribution: Peru. Leptohyphes tacajalo MAYO Leptohyphes tacajalo MAYO, 1968b: 307; HUBBARD, 1982a: 274; MOLINERI, 2003a: 67. Leptohyphes albus MAYO, 1968b: 305; HUBBARD, 1982a: 273; MOLINERI, 2003a: 67.

Taxonomy: The nymphs of this species, the only known stage, can be separated from the others of the genus by the following combination of characters: 1) Tarsal claw denticulation: 4-5 marginal and 1 submarginal subapical; 2) pronotum with a small lateral and somewhat acute projection as in Fig. 96G; 3) hind wingpads present in female; 4) developing wings whitish; 5) without spines on dorsal surface of middle and hind femora; 6) elevated sockets absent on femora; 7) mesonotum with projected anterolateral corners (Fig. 96G). Distribution: Ecuador (Cotopaxi). Leptohyphes tuberculatus ALLEN Leptohyphes sp. 6 ROBACK, 1966: 152. Leptohyphes tuberculatus ALLEN, 1967: 369; ALLEN & ROBACK, 1969: 378; HUBBARD, 1982a: 274; MOLINERI, 2003a: 67.

Taxonomy: This species is only known from nymphs and can be separated from the other of the genus by: 1) Hind femora with a distal tubercle (Fig. 95O); 2) hind wingpads

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absent in female; 3) tarsal claw denticulation: 6 marginal and 3 submarginal subapical denticles (Fig. 96Z); 4) without spines on dorsal surface of middle and hind femora (Fig. 95O); 5) elevated sockets absent on femora; 6) pro- and mesonotum without projected anterolateral corners. Only L. ecuador shares with L. tuberculatus the presence of a tubercle on apex of hind femora, but they can be separated by: tarsal claw denticulation, absence of tubercles on the head of L. tuberculatus, and form of the femora. Distribution: Peru. Genus Leptohyphodes ULMER Leptohyphodes ULMER, 1920a; ULMER, 1920c; ULMER, 1933; TRAVER, 1944; TRAVER, 1958a: 498; ALLEN, 1967: 369; DOMÍNGUEZ et al., 1992; DOMÍNGUEZ et al., 2001; MOLINERI, 2005: 248. (Type-species: Potamanthus inanis PICTET, original designation).

Taxonomy: Leptohyphodes was established by ULMER (1920a) for the species Potamanthus? inanis PICTET. In 1921 ULMER placed in this genus the species Tricorythus australis BANKS, which was later (1958a) transferred by TRAVER to Tricorythodes. Presently, Leptohyphodes is monotypic. MOLINERI (2005) redescribed adults and nymphs and presented SEM photographs of the eggs. A

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Adult Characteristics: 1) Fore wings without extended cubito-anal lobe (Figs. 97AB); 2) hind wings absent in both sexes; 3) female caudal filaments relatively long; 4) membranous filaments of mesoscutellum long and slender; 5) male forceps 2-segmented, distal segment leaf-like (Fig. 97F); 6) penes plate-like, fused except on apical incision (Fig. 97G); 7) eyes of male large and divided (Figs. 97C-D). Nymphal Characteristics: 1) Hind wingpads absent in both sexes; 2) operculate gills subtriangular, dorsally with 2 ridges and a median weaker band (Fig. 99L); 3) gills on abdominal segments II-V, gill formula: 2/3/3/1, lamellae subtriangular (Figs. 99N-Q); 4) fore femora with a transverse row of long setae on 2/3 distance from base (Fig. 99J); 5) middle and hind femora with a transverse row of setae at base (Figs. 99H-I); 6) maxillary palp small, setiform (Fig. 99G); 7) labrum with a deep median cleft (Fig. 99B); 8) tarsal

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claw with 5-6 slightly marked and blunt marginal denticles on basal half and with a double row of 2-3 submarginal denticles on apical 1/3 (Fig. 99K); 9) frontal and genal projections present (Fig. 98A); 10) ) body elongated (Fig. 98A), not heavily sclerotized and covered by numerous long setae; 11) 3-4 small imbricated lobes on ventral lamellae of gills II-IV (Figs. 99N-P). Mouthparts as in Figs. 99A-G. Distribution: Brazil (São Paulo). Biology: Unknown. Leptohyphodes inanis (PICTET) Potamanthus? inanis PICTET, 1843: 232; EATON, 1886: 296 (male). Potamanthus inanis; WALKER, 1853: 544, 547; EATON, 1871: 91 (male). Leptohyphodes inanis; ULMER, 1920a: 51; LESTAGE, 1931a: 74; LESTAGE, 1931b: 60; TRAVER, 1958a: 496 (male, female, nymph); HUBBARD, 1982a: 274; MOLINERI, 2005: 250. Leptohyphodes sp. TRAVER, 1944; MOLINERI, 2005: 250.

Taxonomy: Leptohyphodes inanis is the only named species in this genus, and can be distinguished by the same characters listed above for the genus. Genus Lumahyphes MOLINERI Lumahyphes MOLINERI, in MOLINERI & ZÚÑIGA, 2004: 20; MOLINERI, 2004: 215. (Type-species: Lumahyphes guacra MOLINERI, original designation).

Taxonomy: Lumahyphes is known from all the stages and is represented by 3 species: Lumahyphes guacra MOLINERI (MOLINERI & ZÚÑIGA, 2004), L. yagua MOLINERI & ZÚÑIGA (2004), and L. pijcha MOLINERI (2004). MOLINERI & ZÚÑIGA (2004) reported an additional unnamed species from Mexico. Adult Characteristics: 1) Vein CuP on fore wings with basal portion paralleling CuA, both directed towards wing base (arrow in Fig. 100A); 2) hind wings present in male (Figs. 100B-C), absent in female; 3) hind wings reduced, with a large costal projection (0.42-0.56 total length of hind wing); 4) membranous filaments on mesoscutellum present and relatively long (reaching abdominal tergum II); 5) eyes of male small and remote; 6) styliger plate with median projections (MPS in Figs. 100E-H); 7) forceps 3-segmented, segment I relatively long (Figs. 100E, H, K); 8) penes almost completely fused, with a pair of long spines arising near the apex (PS in Figs. 100G, L); 9) penes with sclerotized lateral margins and basal ring (SLM and BR in Fig. 100G); 10) 2 pairs of distal membranous lobes present on apex of penes (ML in Fig. 100G); 11) with a pair of hook-like projections between penes and cerci (DS in Figs. 100F, I, L). Nymphal Characteristics: 1) Fore femora with a transversal row of long, generally bifid spines (Figs. 101I-J, O-P); 2) gills present on abdominal segments II-VI; 3) gill formula: 3/4/4/3/2, oval lobes (Figs. 101R-W); 6) maxillary palpi 3-segmented and relatively large (Fig. 101D); 7) prementum as in Fig. 101B; 8) tarsal claws with a basal row of marginal denticles and a double row (or single pair) of submarginal denticles near apex (Figs. 101K-L, Q). Distribution: Argentina, Bolivia, Ecuador, Colombia and Mexico.

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Plate 100. Lumahyphes, adults. Figs. 100A-L. L. guacra: 100A, male fore wing; 100B, male hind wing; 100C, same, enlarged; 100D, female fore wing; 100E, male genitalia, v.v.; 100F same, l.v.; 100G, detail of penes, v.v. L. pijcha: 100H, male genitalia, v.v.; 100I, same, l.v.; 100J, detail of penes, d.v. L. yagua: 100K, male genitalia, v.v.; 100L, same, l.v. (BR = basal ring; DS = dorsal structure; BR = basal ring; ML = membranous lobes; MP = membranous dorsal projection; MPS = median projections of styliger plate; PS = penis spines; SLM = sclerotized lateral margin). Figs. 100A-G and K-L modified from MOLINERI & ZUÑIGA, 2004; 100H-J from MOLINERI, 2004.

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Biology: Nymphs of this genus are generally found in medium to large sized streams (>10 m wide), inhabiting stony substrates or submerged wood. Nymphs are mainly crawlers and rarely swim. The nuptial flight occurs early in the morning, 1-3 m above the shore. Male imagos perform the typical up and down movements. KEYS

TO

SOUTH AMERICAN LUMAHYPHES

Adult Males 1 – 2(1) –

Penes relatively long and slender, spines of penes short, straight, not curved but directed dorsally (Figs. 100H-J) .......................................................... Lumahyphes pijcha Penes relatively wider and more robust, spines of penes longer, curved medially (Figs. 100E-G, K) ............................................................................................................... 2 First forceps segment much darker than other segments; in lateral view, apex of median projections of styliger plate strongly curved ventrally (MPS in Fig. 100F) ................................................................................................................... Lumahyphes guacra First forceps segment as dark as remaining segments; in lateral view, apex of median projections of styliger plate not as above (MPS in Fig. 100L) .................... .................................................................................................................... Lumahyphes yagua

Nymphs 1 – 2(1) –

CLAVES

Color markings of head as in Fig. 101X; abdominal terga completely shaded with light gray ................................................................................................... Lumahyphes pijcha Color markings of head not as above; abdominal terga completely shaded with light gray except on a median longitudinal line ........................................................... 2 Abdominal terga with a relatively broad median longitudinal pale line; tarsal claws with double row of 2 or 3 submarginal denticles near apex (Fig. 101K) ................ ................................................................................................................... Lumahyphes guacra Abdominal terga with narrow median longitudinal pale line; tarsal claws with 1 pair of submarginal denticles near apex (Fig. 101L) ...................... Lumahyphes yagua PARA LOS

LUMAHYPHES

SUDAMERICANOS

Machos Adultos 1 – 2(1) –

Penes relativamente largos y delgados, espinas peneanas cortas, no curvadas pero dirigidas dorsalmente (Figs. 100H-J) .................................................. Lumahyphes pijcha Penes relativamente anchos y robustos, espinas peneanas más largas, curvadas hacia la línea media (Figs. 100E-G, K) .......................................................................... 2 Primer segmento de los fórceps mucho más oscuro que el resto; en vista lateral el ápice de las proyecciones medias de la placa estilígera es fuertemente curvado ventralmente (MPS en Fig. 100F) ...................................................... Lumahyphes guacra Primer segmento de los fórceps tan oscuro como los restantes segmentos; en vista lateral el ápice de las proyecciones medias de la placa estilígera no como arriba (MPS en Fig. 100L) ................................................................................ Lumahyphes yagua

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Ninfas 1 – 2(1) –

Patrón de coloración en cabeza similar a Fig. 101X; tergos abdominales completamente sombreados con gris claro ...................................... Lumahyphes pijcha Patrón de coloración en cabeza no como arriba; tergos abdominales completamente sombreados con gris claro excepto sobre una línea mediolongitudinal ............................................................................................................... 2 Abdomen con una línea mediolongitudinal relativamente ancha en los tergos abdominales; uñas tarsales con hilera doble de 2 ó 3 dentículos submarginales cerca del ápice (Fig. 101K) .................................................................. Lumahyphes guacra Línea mediolongitudinal clara del abdomen más delgada; uñas tarsales con 1 par de dentículos submarginales cerca del ápice (Fig. 101L) ............... Lumahyphes yagua

Lumahyphes guacra MOLINERI Lumahyphes guacra MOLINERI, in MOLINERI & ZÚÑIGA, 2004: 25.

Taxonomy: Lumahyphes guacra can be separated from the other species of the genus by the following combination of characters. In the adults: 1) Total length of hind wing (Fig. 100B) 0.18-0.19 total length of fore wing; 2) costal projection of hind wings 0.42 total length of wing (Fig. 100C); 3) segment I of forceps much darker than remaining segments; 4) spines of penes almost as long as penes (Figs. 100E-G); 5) median projections of styliger plate, at base of forceps, as in Fig. 100F; 6) female abdominal sternum VII with a conspicuous black mark. In the nymph: 1) Median longitudinal pale line on abdominal terga relatively wide; 2) tarsal claws with 7-8 marginal denticles and double row of 2-3 submarginal ones (Fig. 101K). Distribution and Biology: Argentina and Bolivia. Males and females where collected at nuptial flight at 9:30 a.m. The swarm was approximately 2-4 m high near the margin of the river. The individuals displayed the typical up and down flight movements. Lumahyphes pijcha MOLINERI Lumahyphes pijcha MOLINERI, 2004: 215.

Taxonomy: Lumahyphes pijcha can be distinguished from the other species of the genus by the following combination of characters. In the imago: 1) Abdominal terga shaded with yellowish gray dorsally, lighter on median longitudinal line; 2) penes relatively long and slender (Figs. 100H, J); 3) spines of penes relatively short and straight, about 1/ 3 total length of penes (Figs. 100H-J); 4) median projection of styliger plate blunt (Fig. 100I); 5) female abdominal sternum VII without black macula. In the nymph: 1) Color markings of head as in Fig. 101X; 2) abdominal terga completely shaded with light gray; 3) operculate gill lightly shaded with gray; 4) tarsal claw long and slender, with 5-7 marginal and 4-5 submarginal denticles (Fig. 101Q). Distribution and Biology: Bolivia and Colombia. The nymphs of this species were collected in small stony streams at 500-1000 m of altitude. The watershed is covered by

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F

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V P W K

Q X

Plate 101. Lumahyphes, nymphs. Figs. 101A-X. L. guacra: 101A: labrum, d.v.; 101B, labium, v.v; 101C, hypopharynx, v.v.; 101D, maxillae, v.v.; 101E, right mandible, d.v.; 101F, left mandible, d.v.; 101G, hind leg; 101H, middle leg; 101I, fore leg; 101J detail of fore femoral spines; 101K, detail of fore tarsal claw. L. yagua: 101L, detail of fore tarsal claw; 101R, operculate gill, d.v.; 101S-W, abdominal gills II-VI, v.v. L. pijcha: 101M, hind leg; 101N, middle leg; 101O, fore leg; 101P, detail of fore femoral spines; 101Q, detail of fore tarsal claw; 101X, head color pattern. Figs. 101G-L modified from MOLINERI & ZUÑIGA, 2004; 101M-X from MOLINERI, 2004.

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mountain rain forest (Yungas). Nymphs were found among rocks and sand in riffle zones. Imagos were attracted to light traps just before dawn. Lumahyphes yagua MOLINERI & ZÚÑIGA Lumahyphes yagua MOLINERI & ZÚÑIGA, 2004: 27.

Taxonomy: Lumahyphes yagua can be separated from the other species of the genus by the following combination of characters. In the imago: 1) Total length of hind wing 0.12-0.15 total length of fore wing; 2) costal projection of hind wings 0.56 total length of wing; 3) FMI with a pair of submedian darker marks; 4) all segments of forceps similar in color; 5) spines of penes almost as long as penes (Fig. 100K); 6) median projections of styliger plate at base of forceps as in Fig. 100L; 7) female abdominal sternum VII without marks or with a small gray mark. In the nymph: 1) Median longitudinal pale line on dorsum of abdomen relatively thin; 2) tarsal claw with 7 marginal denticles and a pair of submarginal denticles (Fig. 101L). Distribution and Biology: Colombia and Peru. Imagos were collected before sunrise, and subimagos were collected at night. Nuptial flight was observed early in the morning, 13 m above riverbanks. The only nymph reported was collected in submerged wood. Genus Traverhyphes MOLINERI Traverhyphes MOLINERI, 2001b: 130; MOLINERI, 2004: 201. (Type-species: Leptohyphes indicator NEEDHAM & MURPHY, original designation).

Taxonomy: Traverhyphes was established by MOLINERI (2001b) for Leptohyphes indicator NEEDHAM & MURPHY (1924) and Traverhyphes pirai MOLINERI (2001b). Leptohyphes indicator NEEDHAM & MURPHY had been previously recognized as a strange species of Leptohyphes, because of its atypical male genitalia. MOLINERI (2001b) reared nymphs of this species, describing this stage and the female adults and eggs. Recently, MOLINERI (2004) proposed 3 subgenera for this group (Traverhyphes s.s., Byrsahyphes, and Mocoihyphes). Adult Characteristics: 1) Fore wings of male without developed cubito-anal lobe (Fig. 102B); 2) hind wings present in males, with 2 longitudinal veins (Figs. 102C-D), hind wings absent in females; 3) female caudal filaments 1.6-2.0 times length of fore wings; 4) membranous filaments of mesoscutellum long and slender; 5) male forceps 3-segmented; 6) penes almost completely fused and apical lobes parallel (Figs. 102G, 103J); 7) male gonopore associated with a hollow penean spine (somewhat hyaline) (Figs. 102H, 103D); 8) penes with subapical-dorsal membranous projections (Figs. 102H, 103D); 9) posterior margin of styliger plate, between forceps, with a pair of small median projections (Figs. 102E, I); 10) with a dorsal structure between penes and cerci, with varied shape: semicircular, hook-like or pyramidal (Figs. 102F, J, 103B-C). Nymphal Characteristics (Fig. 104): 1) Hind wingpads present in males, absent in females; 2) operculate gill suboval or subquadrate, with 2 well marked dorsal ridges (Fig. 106A); 3) gill formula 3/3-4/3/3/2-1 (e.g. Figs. 106B-F); 4) femora I with transverse row of long spines (Fig. 107B); 5) middle and hind femora without transverse basal row of spines at dorsum (Fig. 107A, D-E); 6) maxillary palpi small, 2 or 3-segmented with or

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without apical seta (Figs. 105F-I); 7) labrum with broad anteromedian emargination (Fig. 105A); 8) mandibles as in Figs. 105D-E, labium with enlarged submentum (Fig. 105B), hypopharynx as Fig. 105C; 9) tarsal claw with a double or single row of subapical denticles (Figs. 107C, Q); 9) genal projections present, frontal projections absent. Distribution: From Costa Rica to Northern Argentina. Biology: Nymphs inhabit rocky streams and rivers at low altitudes (100-500 m), and were collected mainly in riffle zones. Swarm activity of male imagos occurred early in the morning. KEYS

TO

TRAVERHYPHES

Adult Males 1 – 2(1) – 3(2) – 4(1) – 5(4) – 6(4) –

Posterolateral margins of styliger plate posteriorly projected (LP in Figs. 102E, I, L); penes with a deep apical incision (Figs. 102G, K, N) ............................................ ........................................................................................... Traverhyphes (Traverhyphes)........2 Posterolateral margins of styliger not projected as above (Figs.103A, E, H); penes almost completely fused (Figs.103D, G, J) ................................................................... 4 Spines of penes inserted dorsally (Figs. 102G-H) ........................................................ ...................................................................................... Traverhyphes (Traverhyphes) indicator Spines of penes inserted laterally or dorsolaterally (Figs. 102K, N) ...................... 3 Penes long and slender, with a similar width along their length (Fig. 102K) .......... .................................................................................... Traverhyphes (Traverhyphes) chiquitano Penes wider distally (Fig. 102N) ................................. Traverhyphes (Traverhyphes) pirai First segment of forceps very short, almost globular (Figs. 103E, H); lateral margins of penes entire (Figs. 103G, J); dorsal accessory structure single (Figs. 103F, I) ............................................................................. Traverhyphes (Byrsahyphes)........5 First segment of forceps longer and rectangular, length almost twice width (Fig. 103A); lateral margins of penes with a subapical constriction (Fig. 103D); accessory dorsal structure double (Fig. 103C) ............................. Traverhyphes (Mocoihyphes)........6 Penes broadly notched apically (Fig. 103G) and with a subapical ventral swelling (VS in Fig. 103F) ............................................................ Traverhyphes (Byrsahyphes) nanus Penes very slightly notched (Fig. 103J) and without subapical ventral swelling (Fig. 103I) ................................................................................... Traverhyphes (Byrsahyphes) yuqui Spines of penes long, about 1/2 length of penes (Fig. 103C) ................................... ............................................................................................. Traverhyphes (Mocoihyphes) yuati Spines of penes short, 1/4 length of penes (Fig. 103B) ............................................. ...................................................................................... Traverhyphes (Mocoihyphes) edmundsi

Nymphs 1 – 2(1) –

Maxillary palpi 3-segmented and relatively large (Fig. 105I) ...................................... ............................................................................................ Traverhyphes (Byrsahyphes) nanus Maxillary palpi 2-segmented and reduced in size (Figs. 105F-H) ........................... 2 Gill formula 3/4/3-4/3/2 (Figs. 106B-F) ............... Traverhyphes (Traverhyphes)........3 Gill formula 3/3/3/3/1-2 (Figs. 106H-L, N-R) ....... Traverhyphes (Mocoihyphes)........4

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B A MP2

D

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I MPS

E

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M

P LP

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G MP

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Plate 102. Traverhyphes (Traverhyphes), adults. Figs. 102A-N. T. (T.) indicator: 102A, female fore wing; 102B, male fore wing; 102C, male hind wing; 102D, same, enlarged; 102E, male genitalia, v.v.; 102F, same, l.v.; 102G, detail of penes, d.v. at left, v.v. at right; 102H, penes, l.v. T. (T.) chiquitano: 102I, male genitalia, v.v.; 102J, same, l.v.; 102K, penes, enlarged, v.v. T. (T.) pirai: 102L, male genitalia, v.v.; 102M, same, l.v.; 102N, detail of penes, d.v. at right, v.v. at left. (DS = dorsal structure; LP = lateral projections; MP = membranous projection; MPS = median projections of styliger plate; P = penes; PS = penean spines). Figs. 102A-H and L-N modified from MOLINERI, 2001b; 102I-K from MOLINERI, 2004.

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3(2) – 4(2)

–

CLAVES

Operculate gill shaded with blackish on anteromedian corner (Fig. 106A) ............ ...................................................................................... Traverhyphes (Traverhyphes) indicator Operculate gill shaded with blackish on the articulation point with the abdomen (basal macula) and transverse ridge .................... Traverhyphes (Traverhyphes) chiquitano Body heavily shaded with purplish gray, large marks present subapically on femora (Figs. 107M-O); abdomen uniformly shaded, rarely with a median longitudinal lighter line; maxillary palpi without apical seta (Fig. 105H) ................. ............................................................................................. Traverhyphes (Mocoihyphes) yuati Body shaded slightly with gray, subapical marks on femora small (Figs. 107I-K); abdomen with a median longitudinal pale line, broader on terga VII-IX (Fig. 105K); maxillary palpi with apical seta (Fig. 105G) ...................................................... ...................................................................................... Traverhyphes (Mocoihyphes) edmundsi PARA

TRAVERHYPHES

Machos Adultos 1 – 2(1) – 3(2) – 4(1) – 5(4) – 6(4) –

Márgenes posterolaterales de la placa estilígera proyectados posteriormente; (LP en Figs. 102E, I, L); penes con una incisión apical profunda (Figs. 102G, K, N) . ........................................................................................... Traverhyphes (Traverhyphes)........2 Márgenes de la placa estilígera no proyectados de esa forma (Figs. 103A, E, H); penes casi completamente fusionados (Figs. 103D, G, J) .......................................... 4 Espinas de los penes insertadas dorsalmente (Figs. 102G-H) ................................... ...................................................................................... Traverhyphes (Traverhyphes) indicator Espinas de los penes insertadas lateralmente (Figs. 102K, N) ................................. 3 Penes largos y delgados, de un ancho similar en toda su longitud (Fig. 102K) ..... .................................................................................... Traverhyphes (Traverhyphes) chiquitano Penes más anchos distalmente (Fig. 102N) .............. Traverhyphes (Traverhyphes) pirai Primer segmento de los fórceps muy corto, casi globular (Figs. 103E, H); márgenes laterales de los penes enteros (Figs. 103G, J); estructura accesoria dorsal simple (Figs. 103F, I) ..................................................... Traverhyphes (Byrsahyphes)........5 Primer segmento de los fórceps mas largo, rectangular (Fig. 103A); márgenes laterales de los penes con una constricción subapical (Fig. 103D); estructura accesoria dorsal doble (Fig. 103C) .............................. Traverhyphes (Mocoihyphes)........6 Penes con ranura apical amplia (Fig. 103G) y con una hinchazón ventral subdistal (VS en Fig. 103F) ........................................................... Traverhyphes (Byrsahyphes) nanus Penes casi sin ranura apical (Fig. 103J); hinchazón ventral ausente (Fig. 103I) ...... ............................................................................................. Traverhyphes (Byrsahyphes) yuqui Espinas de los penes largas, 1/2 del largo total de los penes (Fig. 103C) ............... ............................................................................................. Traverhyphes (Mocoihyphes) yuati Espina de los penes cortas, 1/4 o menos de la longitud total de los penes (Fig. 103B) ........................................................................... Traverhyphes (Mocoihyphes) edmundsi

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Plate 103. Traverhyphes (Mocoihyphes), adults. Figs. 103A-J. T. (M.) edmundsi: 103A, male genitalia, v.v.; 103B, same, l.v.; 103D, penes, d.v. T. (M.) yuati: 103C, male genitalia, l.v. Traverhyphes (Byrsahyphes), adults. T. (B.) nanus: 103E, male genitalia, v.v.; 103F, penes, l.v.; 103G, penes, d.v. T. (B.) yuqui: 103H, male genitalia, v.v.; 103I, same, l.v.; 103J, penes, v.v. (AL = Apical lobes; DL = Dorsal lobes; DS = dorsal structure; P = penes; PS = penean spines; VS = Ventral swelling). Figs. 103A-J modified from MOLINERI, 2004.

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Ninfas 1 – 2(1) – 3(2) – 4(2)

–

Palpo maxilar 3-segmentado y relativamente grande (Fig. 105I) ............................... ............................................................................................ Traverhyphes (Byrsahyphes) nanus Palpo maxilar 2-segmentado y reducido en tamaño (Figs. 105F-G) ....................... 2 Fórmula branquial 3/4/3-4/3/2 (Figs. 106B-F) ........................................................... .................................................................................. Traverhyphes (Traverhyphes).................3 Fórmula branquial 3/3/3/3/1-2 (Figs. 106H-L, N-R) ................................................ ...................................................................................Traverhyphes (Mocoihyphes).................4 Branquia opercular sombreada con negruzco en el borde anteromedial (Fig. 106A) .......................................................................... Traverhyphes (Traverhyphes) indicator Branquia opercular sombreada con negruzco en el punto de articulación con el abdomen (mácula basal) y sobre la costilla transversal ................................................ .................................................................................... Traverhyphes (Traverhyphes) chiquitano Cuerpo fuertemente sombreado con gris purpúreo, marcas subapicales de los fémures relativamente grandes (Figs. 107M-O); abdomen sombreado más o menos uniformemente, raramente con linea mediolongitudinal mas clara; palpo maxilar sin seta apical (Fig. 105H) .............................. Traverhyphes ( Mocoihyphes) yuati Cuerpo suavemente sombreado con gris, marcas femorales más pequeñas (Figs. 107I-K); abdomen con banda longitudinal media más clara que se ensancha en los tergos VII-IX (Fig. 105K); palpo maxilar con seta apical (Fig. 105G) ..................... ..................................................................................... Traverhyphes ( Mocoihyphes) edmundsi

Subgenus Traverhyphes (Traverhyphes) MOLINERI Traverhyphes MOLINERI, 2001b; MOLINERI, 2004: 201. (Type-species: Leptohyphes indicator NEEDHAM & MURPHY, original designation).

Taxonomy: Traverhyphes (Traverhyphes) is currently known from 3 species and all the stages. This subgenus can be distinguished from the other subgenera of the genus by the following combination of characters. In the imago: 1) Posterolateral corners of styliger plate projected posteriorly (Figs. 102E, I, L); 2) penes with deep apical incision, generally not visible (Figs. 102G, K, N); 3) paired membranous dorsal lobes on penes (Fig. 102G). In the nymphs: 1) Operculate gills subquadrate (Fig. 106A); 2) gill formula: 3/4/3-4/3/2 (Figs. 106B-F); and 3) maxillary palpi small with 2 segments and an apical seta (Fig. 105F). Distribution: Argentina, Bolivia, Brazil, Uruguay. Probably, the distribution range will prove to be more widespread because subimagos of an undescribed species from Central America were studied. Traverhyphes (Traverhyphes) chiquitano MOLINERI Traverhyphes chiquitano MOLINERI, 2004: 201.

Taxonomy: Traverhyphes (Traverhyphes) chiquitano is known from all the stages and can be distinguished from the other species of the subgenus by the following combination of characters. In the imago: 1) Abdominal terga shaded with gray, darker

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Plate 104. Traverhyphes (T.) indicator. Fig. 104A, nymphal habitus. Drawing by C. Molineri.

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at lateral margins; 2) penes with a similar width along their length (Fig. 102K); 3) spines of penes inserted laterally (Fig. 102K); 4) in lateral view posterolateral projections of styliger plate short and broad apically (Fig. 102J); 6) veins IMP and MP2 of fore wings not connected by cross veins. In the nymph: 1) Operculate gill shaded with blackish on a basal macula and transverse ridge. Distribution and Biology: Bolivia (Santa Cruz). Nymphs were collected in rivers with stony substrate at 200 m of altitude. Adults were captured with light traps before sunrise. Traverhyphes (Traverhyphes) indicator (NEEDHAM & MURPHY) Leptohyphes indicator NEEDHAM & MURPHY, 1924: 33; LESTAGE, 1931b: 60; NAVÁS, 1931b: 322; TRAVER, 1958a: 500; HUBBARD, 1982a: 274; DOMÍNGUEZ, 1984: 103; DOMÍNGUEZ et al., 1994: 99; MOLINERI, 2001b: 201. Traverhyphes (Traverhyphes) indicator; MOLINERI, 2004: 201.

Taxonomy: Traverhyphes (Traverhyphes) indicator is known from all stages and can be distinguished from the other species of the subgenus by the following combination of characters. In the imago: 1) Abdominal terga shaded with black on lateral margins, with a wide median pale band; 2) penes with a similar width along their length (Fig. 102G); 3) spines of penes inserted dorsally (Figs. 102G-H); 4) in lateral view, posterolateral projections of styliger plate long and relatively uniform in width (Fig. 102F); 6) veins IMP and MP2 of fore wings connected by 2-4 cross veins (Figs. 102A-B). In the nymph (Fig. 104A): 1) Operculate gill shaded with blackish on anteromedian corner (Fig. 106A). Distribution and Biology: Argentina (Misiones), Uruguay (Lavalleja), Southern Brazil. This species occurs in a wide variety of rivers and streams, preferentially with stony substrate. The nymphs were collected on rock substrate, in reaches with moderate to high current velocity. Subimagos emerge during the night, and male imagos fly at early morning, performing the usual up and down flight movements. Swarms are variable in number of individuals but generally are located above the water or shores, at a height of 1-2 m. To oviposit, female imagos extrude a single mass of eggs that is retained in the tip of the abdomen, until the female touches the water surface. Traverhyphes (Traverhyphes) pirai MOLINERI Traverhyphes pirai MOLINERI, 2001b: 138. Traverhyphes (Traverhyphes) pirai; MOLINERI, 2004: 201.

Taxonomy: This species is only known from male subimagos and can be distinguished from the other species of the subgenus by the following combination of characters: 1) Abdominal terga shaded uniformly with gray, darker on a pair of submedian longitudinal lines on terga I-VI; 2) apical half of penes wider than basal half (Fig. 102N); 3) spines of penes inserted laterally (Fig. 102N); 4) posterolateral projections of styliger plate as in Figs. 102L-M. Distribution: Brazil (Rio de Janeiro).

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Subgenus Traverhyphes (Byrsahyphes) MOLINERI Byrsahyphes MOLINERI, 2004: 209. (Type-species: Leptohyphes nanus ALLEN, original designation).

Taxonomy: The subgenus Traverhyphes (Byrsahyphes) can be distinguished from the other subgenera by the following combination of characters. In the imago: 1) Posterolateral projections of styliger plate absent (Figs. 103E, H); 2) penes almost completely fused with 2 pairs of apical lobes (Figs. 103G, J); 3) penes with a pair of dorsal membranous projections

A

B

C

D E G

F

H

I

K J

Plate 105. Traverhyphes, nymphs. Figs. 105A-K. Mouthparts (T. (M.) yuati): 105A, labrum, d.v. at left, v.v. at right; 105B, labium; 105C, hypopharynx; 105D, right mandible; 105E, left mandible. Maxilla and detail of palpi: 105F, T. (T.) indicator; 105G, T. (M.) edmundsi; 105H, T. (M.) yuati; 105I, T. (B.) nanus. Abdomen (right gills omitted): 105J, T. (B.) nanus; 105K, T. (M.) edmundsi. Figs. 105A-K modified from MOLINERI, 2004.

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(Figs. 103G, J); 4) penes with a pair of spines arising at base of dorsal projections; 5) accessory dorsal structure associated with penes, single and acute (Figs. 103F, I). In the nymph: 1) Operculate gill elongate ovoid (Fig. 106S); 2) gill formula: 3/3/3/3/1 (Figs. 106T-X); 3) maxillary palpi larger than in other subgenera, with 3 segments and an apical seta (Fig. 105I). Distribution: Mountain rain forests from Bolivia, Colombia, Costa Rica and Panama. Traverhyphes (Byrsahyphes) nanus (ALLEN) Leptohyphes nanus ALLEN, 1967: 355; ALLEN, 1978: 550. Allenhyphes nanus; WIERSEMA & MCCAFFERTY, 2000: 343. Traverhyphes (Byrsahyphes) nanus; MOLINERI, 2004: 211.

Taxonomy: This species was originally described from nymphs (ALLEN, 1967). WIERSEMA & MCCAFFERTY (2000) transferred it to Allenhyphes. Later MOLINERI (2004), described the adults and, based on a cladistic analysis of the group, included this species in Traverhyphes (Byrsahyphes). T. (Byrsahyphes) nanus can be distinguished from the other species of the subgenus by the following combination of characters. In the adults: 1) Gray subapical marks on femora weak or absent; 2) abdominal terga lightly shaded with gray; 3) spines of penes short, less than 1/4 total length of penes (Figs. 103F-G); 4) fore wing veins yellowish brown, lighter toward hind margin; 5) dorsal structure associated with penes apically acute as in Fig. 103F. In the nymphs: 1) Gray subapical marks on femora weak or absent (Figs. 107DE; 2) abdominal terga lightly shaded with gray (Fig. 105J); 3) operculate gills shaded with yellowish gray (Fig. 106S); 4) maxillary palpi 3-segmented with apical seta (Fig. 105I). Distribution: Costa Rica, Colombia. Traverhyphes (Byrsahyphes) yuqui MOLINERI Traverhyphes (Byrsahyphes) yuqui MOLINERI, 2004: 212.

Taxonomy: Traverhyphes (Byrsahyphes) yuqui is known from imagos of both sexes, and can be distinguished from the other species of the subgenus by the following combination of characters: 1) Abdominal terga uniformly and lightly shaded with gray, median line paler on terga I-V; 2) spines of penes very short, less than 1/6 total length of penes (Figs. 103I-J); 3) penes without ventral membranous extension, and without deep apicomedial notch (Fig. 103J); 4) veins of fore wings yellowish brown; and 5) accessory dorsal structure associated with penes single and relatively blunt (Fig. 103I). Distribution: Bolivia, Panama. Subgenus Traverhyphes (Mocoihyphes) MOLINERI Mocoihyphes MOLINERI, 2004: 203. (Type-species: Leptohyphes edmundsi ALLEN, original designation).

Taxonomy: Traverhyphes (Mocoihyphes) was established for 2 species from Argentina and Brazil; it is known from all stages. This subgenus can be distinguished from the other subgenera of the genus by the following combination of characters. In the imago: 1) Posterolateral projections of styliger plate absent (Figs. 103A-C); 2) penes almost

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Plate 106. Traveryphes, nymphs. Figs. 106A-X. Abdominal gills II d.v., II-VI v.v.: 106A-106F, T. (T.) indicator; 106G-L, T. (M.) edmundsi; 106M-R, T. (M.) yuati; 106S-X, T. (B.) nanus. (UZ = unpigmented zone). Figs. 106A-X modified from MOLINERI, 2004.

completely fused (Fig. 103D); 3) penes with 2 pairs of membranous apical lobes (Fig. 103D); 4) penes with a pair of dorsal membranous projections (Fig. 103D); 5) penes with a pair of spines that arise at base of dorsal projections (Fig. 103D); 5) accessory dorsal structure, double (Figs. 103B-C); 6) lateral margins of penes with a lateral notch at apex (Fig. 103D). In the nymph: 1) Operculate gill ovoid to subquadrate (Figs. 106G, M); 2) gill formula: 3/3/3/3/2 (Figs. 106H-L and N-R); and 3) maxillary palpi small with 2 segments with or without apical seta (Figs. 105G-H). Distribution: Southern Brazil, NE Argentina. Traverhyphes (Mocoihyphes) edmundsi (ALLEN) Leptohyphes edmundsi ALLEN, 1973b: 363; HUBBARD, 1982a: 272; MOLINERI, 2001b: 134. Allenhyphes edmundsi; WIERSEMA & MCCAFFERTY, 2000: 343. Traverhyphes (Mocoihyphes) edmundsi; MOLINERI, 2004: 203.

Taxonomy: Traverhyphes (M.) edmundsi was originally described from nymphs (ALLEN, 1973b). WIERSEMA & MCCAFFERTY (2000) transferred it to Allenhyphes. Later MOLINERI (2004), described the adults and eggs and included this species in a subgenus of Traverhyphes. Traverhyphes (Mocoihyphes) edmundsi can be distinguished from the other species of the subgenus by the following combination of characters. In the imago: 1) Subapical small gray marks on femora; 2) abdominal terga with a pale median band, wider on terga VII-IX (see Fig. 105K); 3) spines of penes short, less than 1/4 length of penes (Figs. 103B, D); 4) fore wing veins translucent yellowish; 5) dorsal structure associated with penes with blunt apex (Fig. 103B). In the nymphs: 1) Subapical small gray marks on femora (Figs. 107I-K); 2) abdominal terga with a pale median band, wider on terga VII-IX

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(Fig. 105K); 3) operculate gill yellowish brown, shaded with gray at base (Fig. 106G); 4) maxillary palpi with apical seta (Fig. 105G). Distribution and Biology: Brazil (Rio Grande do Sul, Santa Catarina, Paraná), Argentina (Misiones). A gynandromorph (male subimago), showing normal appearance and sexual characteristics, had an abdomen filled with eggs. Swarming activity occurred early in the morning. Swarms were small to medium sized (10-100 insects) located 2-3 m above the river, each male flying in a vertical zig-zag pattern. Traverhyphes (Mocoihyphes) yuati MOLINERI Traverhyphes (Mocoihyphes) yuati MOLINERI, 2004: 207.

Taxonomy: Traverhyphes (M.) yuati is known from all the stages and can be distinguished from the other species of the subgenus by the following combination of characters. In the imago: 1) Large and well defined subapical marks present on femora; 2) abdominal terga uniformly shaded with purplish gray; 3) spines of penes long, more than 1/2 total length of penes (Fig. 103C); 4) fore wing veins yellowish brown; 5) dorsal structure associated with penes with hook-like apex (Fig. 103C). In the nymphs: 1) Large and well marked subapical marks present on femora (Figs. 107M-O); 2) abdominal terga uniformly shaded with purplish gray; 3) operculate gills yellowish brown (Fig. 106M); 4) general coloration with purplish tones; 5) maxillary palpi very small, without apical seta (Fig. 105H). Distribution and Biology: Argentina (Misiones). This species is sympatric with its sister species, T. (M.) edmundsi, inhabiting the same rivers and streams. Male imagos were collected at low numbers in the same swarms with T. (T.) indicator and T. (M.) edmundsi. Genus Tricorythodes ULMER Tricorythodes ULMER, 1920a: 51; NEEDHAM et al., 1935: 630; TRAVER, 1958a: 501; ALLEN, 1967: 369; ALLEN & BRUSCA, 1973: 94; ALLEN, 1977: 431; ALLEN & MURVOSH, 1987: 36; WIERSEMA & MCCAFFERTY, 2000: 353; MOLINERI, 2002: 305. (Type-species: Tricorythus explicatus EATON, original designation). Tricorythodes (Tricoryhyphes) ALLEN & MURVOSH, 1987: 38. (Type-species: Tricorythodes condylus ALLEN, original designation). Tricorythodes (Homoleptohyphes) ALLEN & MURVOSH, 1987: 39. (Type-species: Tricorythodes dimorphus ALLEN, original designation). Tricoryhyphes; WIERSEMA & MCCAFFERTY, 2000: 351. (Type-species: Tricorythodes condylus ALLEN, original designation). Asioplax WIERSEMA & MCCAFFERTY, 2000: 347. (Type-species: Tricorythodes edmundsi ALLEN, original designation). Epiphrades WIERSEMA & MCCAFFERTY, 2000: 349. (Type-species: Tricorythodes undatus LUGO-ORTIZ & MCCAFFERTY, original designation). Homoleptohyphes; WIERSEMA & MCCAFFERTY, 2000: 350. (Type-species: Tricorythodes dimorphus ALLEN, original designation).

Taxonomy: ULMER (1920a) established Tricorythodes for Tricorythus explicatus EATON (1892) known from adults from Mexico. The nymphal stage had already been

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described by NEEDHAM (1905) as Caenis allecta. TRAVER (1958a, 1959a) characterized the adult stage in detail. MOLINERI (2002) proposed a phylogeny for the South American species of the genus. A

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Plate 107. Traverhyphes, nymphs. Figs. 107A-Q. T. (T.) indicator: 107A, hind leg; 107B, fore leg; 107C, fore tarsal claw. T. (B.) nanus: 107D, hind leg; 107E, middle leg; 107F, fore leg; 107G, femoral spines; 107H, fore tarsal claw. T. (M.) edmundsi: 107I, hind leg; 107J, middle leg; 107K, fore leg; 107L, fore tarsal claw. T. (M.) yuati: 107M, hind leg; 107N, middle leg; 107O, fore leg; 107P, femoral spines; 107Q, fore tarsal claw. Figs. 107A-Q modified from MOLINERI, 2004.

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This Pan-American genus shows great specific diversity, and the description of several new species from the Neotropics is expected. The morphological diversity is more marked at the nymphal stage, with various groups of different body form and size, the adult stage being much more uniform. This may be the reason for some nymphs of Tricorythodes have been described in other genera (commonly in Leptohyphes). This great nymphal diversity led WIERSEMA & MCCAFFERTY (2000) to divide this group in many genera (Asioplax, Epiphrades, Homoleptohyphes, Tricoryhyphes). MOLINERI (2002) showed that some of these groups are not supported in a phylogenetic context, and thus we prefer to treat the group as a unit. Currently, there are 15 valid South American species. DEMOULIN (1966a) described a Tricorythodes sp. from nymphs from Surinam, but these nymphs do not show diagnostic characters of Tricorythodes. Adult Characteristics: 1) Male fore wings with well developed cubito-anal lobe (CL in Figs. 108A, C); 2) hind wings absent in both sexes; 3) female terminal filament covered with small setae and longer than cerci, cerci glabrous and shorter than body; 4) membranous filaments of mesoscutellum short and wide; 5) male forceps 3segmented, second segment with a basal swelling (Figs. 108K-L); 6) penes almost completely fused (except T. ocellus ALLEN), apical lobes of penes not divergent; 7) male gonopore associated with a blunt apical spine in T. ocellus ALLEN (Fig. 108O), but spine reduced or absent in remaining species; 8) penes without projections (except T. australis, Fig. 108H); 9) styliger plate with a pair of small median projections on apical margin, between forceps (Fig. 108L). Nymphal Characteristics (Figs. 109A, 112A): 1) Hind wingpads absent in both sexes; 2) operculate gill of abdominal segment II triangular (Figs. 110Q, 111A-C, E), suboval (Fig. 111D) or subquadrate (Fig. 111F), with 2 dorsal ridges (Fig. 110Q); 3) gill formula 3/3/3/3/2 (Figs. 110R-V), number of lamellae reduced in some species, gill lamellae generally subtriangular; 4) fore femora with transverse row of long setae, position of row variable (Figs. 111G, I, K, M); 5) middle and hind femora with transverse dorsal row of setae at base (Figs. 111H, J, L, N); 6) maxillary palpi reduced (Figs. 110N-P), with 3, 2 or 1 segment, generally with apical seta; 7) labrum with deep to broad anteromedian emargination (Fig. 110K); 8) tarsal claws with a marginal row of denticles and 2 submarginal rows near apex (Figs. 111P-R), rarely some of these rows absent (Fig. 111O); 9) genal and frontal projections present (sometimes frontal projections reduced). Distribution: Neotropical and Nearctic (from central Argentina to Canada). Biology: The nymphs of Tricorythodes are encountered in a wide variety of clean flowing water habitats, from small stony streams to medium-sized, sandy bottomed rivers. They mainly dwell among banks or patches of sand, but may also be found in coarser substrate or among Cladophora filaments. The adults emerge throughout the year, but the greater swarms can be seen in spring-summer. The mating flight takes place along the shore of the water course, at a height of 0.5-3 m, and sometimes ovoid swarms may form. Females of 1 species (T. popayanicus) were observed to enter repeatedly the same swarm, each time mating with a male. To oviposit, females release the eggs in a single mass that remains attached to the tip of the abdomen until they fly down and touch the water.

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Plate 108. Tricorythodes, adults. Figs. 108A-S. Fore wings: 108A, T. yura (male); 108B, T. yura (female); 108C, T. bullus (male). Male mesothoracic legs: 108D, T. bullus; 108E, T. santarita; 108F, T. arequita; 108G, T. mirca. Male genitalia: 108H, T. australis; 108I, T. arequita; 108J, T. mirca; 108K, T. bullus; 108L, T. hiemalis; 108M, T. quizeri; 108N, T. ocellus; 108O, idem, detail of penes; 108P, T. popayanicus; 108Q, T. yura; 108R, T. santarita; 108S, T. zunigae. (AS = Apical spine; BS = Basal swelling; CL = Cubito-anal lobe; DF = Distal furrow; FB = Femoral bands; G = Gonopore; MP = Median projection of styliger plate; SB = Subapical band; SC = Subapical constriction; SM = Subapical mark; VP = Ventral projection; I, II, III = forceps segments). Figs. 108A-S modified from MOLINERI, 2002.

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TO

SOUTH AMERICAN TRICORYTHODES

Adults 1 – 2(1) 3(2) 4(2) – 5(4) – 6(4) – 7(6) – 8(6) – 9(8) – 10(9) –

Penes without ventral projections; first segment of forceps variable and second segment with basal swelling (Figs. 108K-108L) ........................................................... 2 Penes with a ventral projection covered with setae (Fig. 108H); first segment of forceps with apicomedial projection and second segment without basal swelling (Fig. 108H) .......................................................................................... Tricorythodes australis Tibiae whitish with conspicuous blackish subapical marks (SM in Figs. 108F-G) ................................................................................................................................................ 3 Tibiae without subapical blackish marks (Figs. 108D-E) .......................................... 4 Blackish subapical marks present on tibiae and tarsi (Fig. 108F) ..... Tricorythodes arequita Blackish subapical marks present only on tibiae (Fig. 108G) ...... Tricorythodes mirca Femora with 2 or 3 reddish or grayish transverse bands (FB in Fig. 108E) ......... 5 Femora without strong marks or with a subapical grayish band, sometimes more extended but never as above (Fig. 108D) ..................................................................... 6 Fore wings shaded with gray or black on basal 1/3; genitalia as in Fig. 108R ....... .............................................................................................................. Tricorythodes santarita Fore wings shaded with black only on costal margin; genitalia as in Fig. 108S ..... ................................................................................................................ Tricorythodes zunigae Penes abruptly narrowed on distal half, as in Figs. 108L-M ..................................... 7 Penes pyramidal or plate-like (Figs. 108K, N-S), never as above ............................ 8 Penes divided on apical 1/5; penes 1.5 times longer than its basal width (Fig. 108L) .................................................................................................... Tricorythodes hiemalis Penes divided on apical 1/6; penes 2 times longer than its basal width (Fig. 108M) ................................................................................................................. Tricorythodes quizeri Penes plate-like with a strongly marked distal furrow, distal arms of penes well separated from each other (Figs. 108N-O) .................................... Tricorythodes ocellus Penes almost completely fused with a small to medium distal furrow, distal arms of penes not separated from each other (Figs. 108K, P-S) ..................................... 9 Vein CuP of fore wings entire, reaching posterior margin of wing (Figs. 108A-B) .............................................................................................................................................. 10 Vein CuP of fore wings generally incomplete, not reaching hind margin of wing (Figs. 108C) ............................................................................................. Tricorythodes bullus Abdominal segments shaded with gray except on intersegmental membranes, penes without marked constrictions (Fig. 108P) ................... Tricorythodes popayanicus Abdominal segments almost without shading; penes with a subapical constriction (SC in Fig. 108Q) .................................................................................... Tricorythodes yura

Nymphs 1 –

Body very depressed and discoidal; femora almost ovoid and fringed with very long setae (Figs. 111K-N); operculate gills ovoid or subquadrate (Figs. 111D, F) .............. 2 Body not so depressed; femora slender (Figs. 111G-J); operculate gills triangular (Figs. 110Q, 111A-C, E) ................................................................................................... 4

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2(1)

Abdominal tubercles absent, abdominal color pattern as in Fig. 110D ................... ................................................................................................................ Tricorythodes zunigae – Abdominal tubercles present (Fig. 110H); color pattern not as above .................. 3 3(2) Abdomen with median tubercles on terga VII-IX (Figs. 110H, 112A) ................... .............................................................................................................. Tricorythodes santarita – Abdomen with median tubercles on terga III-IX, larger on terga VI-IX ............... .............................................................................................................. Tricorythodes nicholsae 4(1) Dorsal tubercles present on head, pronotum and mesonotum (Figs. 110E-F) .... 5 – Head and thorax without tubercles ................................................................................ 6 5(4) Tubercles large and acute in lateral view (Fig. 110E); abdominal color pattern as in Fig. 110C ................................................................................................. Tricorythodes bullus – Tubercles small and blunt, slightly elevated above head and thorax (Fig. 110F) ... ........................................................................................................... Tricorythodes cristatus 17 6(4) Operculate gill whitish or yellowish shaded with gray or black only at base (Figs. 111B, E) ................................................................................................................................ 7 – Operculate gill almost completely shaded with gray or black (Figs. 110Q, 111A, C) ................................................................................................................................................ 8 7(6) Operculate gill shaded with black on basal 1/4 (Fig. 111B); tarsal claws with more than 1 pair of submarginal denticles near apex (Fig. 111Q) ...................................... ..........................................................................................................Tricorythodes popayanicus – Operculate gill shaded with gray only on a transverse basal line (Fig. 111E); tarsal claws with 1 pair of submarginal denticles near apex (Fig. 111R) ............... Tricorythodes yura 8(6) Tibiae and tarsi whitish with conspicuous subapical blackish marks at least on tibiae (Figs. 111G-H); color pattern formed by small pigmented marks irregularly distributed (Fig. 110A) ...................................................................................................... 9 – Tibiae and tarsi without these marks (Figs. 111I-J); color pattern formed by pigmentation uniformly distributed (Fig. 110B) ........................................................ 10 9(8) Maxillary palpi 3-segmented (Fig. 110O); big blackish marks present on tibiae and tarsi (Figs. 111G-H) .......................................................................... Tricorythodes arequita – Maxillary palpi 2-segmented with an apical setae (Fig. 110P); blackish marks present on tibiae only ........................................................................... Tricorythodes mirca 10(8) Lateral ocelli large, almost as large as compound eyes (Fig. 110G) .......................... .................................................................................................................. Tricorythodes ocellus – Lateral ocelli small, at least 40% smaller than compound eyes (Fig. 109A) ....... 11 11(10) Abdominal segments III-VI expanded laterally (LE in Fig. 110B), posterolateral spines present on abdominal segment VII; tarsal claws without marginal denticles, with only 1 pair of submarginal denticles near apex (Fig. 111O) ............................. ..................................................................................................................Tricorythodes barbus – Abdominal segments III-VII expanded laterally (similar to Fig. 110A), posterolateral spines present on abdominal segment VII-VIII; tarsal claws with 4 or 5 marginal denticles and 1 pair of submarginal denticles near apex (Fig. 111P) ...................... 12 17

T. cristatus: abdominal color pattern was not adequately described by Allen, and bodies of the paratypes were not available for study.

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12(11) Operculate gill shaded completely with blackish (Fig. 111A); frontal projection present ................................................................................................. Tricorythodes hiemalis – Operculate gill shaded completely with blackish except at anterolateral corner (Fig. 111C); frontal projection absent ............................................. Tricorythodes quizeri CLAVES

PARA LOS

TRICORYTHODES

SUDAMERICANOS

Adultos 1 – 2(1) – 3(2) – 4(2) – 5(4)

6(4) – 7(6) – 8(6) – 9(8) –

Penes sin proyecciones ventrales; segmento I del fórceps variable, segmento II con engrosamiento basal (Figs. 108K-L) ...................................................................... 2 Penes con proyección ventral (Fig. 108H); segmento I del fórceps con proyección distomedial, segmento II sin engrosamiento basal (Fig. 108H) ................................. ............................................................................................................... Tricorythodes australis Tibias blanquecinas con notorias marcas subapicales negruzcas (SM en Figs. 108F-G) ................................................................................................................................ 3 Tibias sin marcas subapicales (Figs. 108D-E) .............................................................. 4 Marcas subapicales negruzcas presentes en tibias y tarsos (Fig. 108F) .................... ............................................................................................................... Tricorythodes arequita Marcas subapicales negruzcas presentes solo en las tibias (Fig. 108G) ................... ................................................................................................................... Tricorythodes mirca Fémures de todas las patas con 2 ó 3 bandas transversas grisáceas o rojizas (FB en Fig. 108E) ............................................................................................................................. 5 Fémures sin marcas fuertes o con una banda subapical grisácea, algunas veces más extendida pero nunca como en el caso anterior (Fig. 108D) ................................... 6 Alas anteriores con el 1/3 basal negruzco; genitalia como en la Fig. 108R ............ .............................................................................................................. Tricorythodes santarita Alas anteriores sombreadas con gris o negro solo en el margen costal; genitalia como en la Fig. 108S ......................................................................... Tricorythodes zunigae Penes afinándose bruscamente en la mitad distal, como en las Figs. 108L-M ..... 7 Penes piramidales o en forma de placa (Figs. 108K, N-S), nunca como en el caso anterior ................................................................................................................................. 8 Penes divididos en el 1/5 apical; penes 1,5 veces más largos que su ancho basal (Fig. 108L) ........................................................................................... Tricorythodes hiemalis Penes divididos en el 1/6 apical; penes 2 veces más largos que su ancho basal (Fig. 108M) ..................................................................................................... Tricorythodes quizeri Penes en forma de placa, con una profunda incisión distal media (Figs. 108N-O); brazos de los penes separados distalmente .................................... Tricorythodes ocellus Penes fusionados casi completamente, sin incisión tan marcada (Figs. 108K, P-S); brazos no separados distalmente .................................................................................... 9 Vena CuP de las alas anteriores entera, alcanzando el margen posterior de las alas (Figs. 108A-B) ................................................................................................................... 10 Vena CuP generalmente incompleta, no alcanzando el margen posterior de las alas (Fig. 108C) .............................................................................................. Tricorythodes bullus

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Plate 109. Tricorythodes popayanicus. Fig. 109A, nymphal habitus. Drawing by C. Molineri.

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10(9) –

Segmentos abdominales sombreados con gris, excepto en las membranas intersegmentales; penes sin constricciones marcadas (Fig. 108P) ............................. ..........................................................................................................Tricorythodes popayanicus Segmentos abdominales casi sin sombreado, penes con una constricción subapical (SC en Fig. 108Q) ................................................................................... Tricorythodes yura

Ninfas 1 – 2(1) – 3(2) – 4(1) – 5(4) – 6(4) – 7(6) – 8(6) –

18

Cuerpo discoidal y muy deprimido; fémures casi ovoides y orlados por largas espinas (Figs. 111K-N); branquias operculares ovales o subcuadradas (Figs. 111D, F) ................................................................................................................................................ 2 Cuerpo no tan deprimido; fémures más delgados (Figs. 111G-J); branquia opercular triangular (Figs. 110Q, 111A-C, E) .............................................................. 4 Tubérculos abdominales ausentes, patrón de coloración como en la Fig. 110D ... ................................................................................................................ Tricorythodes zunigae Tubérculos abdominales presentes (Fig. 110H); patrón de coloración no como en el caso anterior .................................................................................................................... 3 Abdomen con tubérculos medianos en los tergos VII-IX (Figs. 110H, 112A) ...... .............................................................................................................. Tricorythodes santarita Abdomen con tubérculos medianos en los tergos III-IX, más grandes en VI-IX .............................................................................................................. Tricorythodes nicholsae Tubérculos dorsales presentes en cabeza, pronoto y mesonoto (Figs. 110E-F) .. 5 Sin tubérculos en la cabeza o tórax ................................................................................ 6 En vista lateral los tubérculos son grandes y aguzados (Fig. 110E); patrón de coloración abdominal como en la Fig. 110C .................................. Tricorythodes bullus Tubérculos pequeños y romos, elevados apenas sobre la cabeza y el tórax (Fig. 110F) ................................................................................................ Tricorythodes cristatus 18 Branquia opercular blanquecina o amarillenta sombreada con gris o negro solo en la base (Figs. 111B-E) ........................................................................................................ 7 Branquia opercular sombreada con gris o negro casi completamente (Figs. 110Q, 111A, C) ............................................................................................................................... 8 Branquia opercular sombreada con negruzco en el 1/4 basal (Fig. 111B); uñas tarsales con más de 1 par de dentículos submarginales cerca del ápice (Fig. 111Q) ..........................................................................................................Tricorythodes popayanicus Branquia opercular sombreada con gris solo en una línea transversa basal (Fig. 111E); uñas tarsales con 1 par de dentículos submarginales cerca del ápice (Fig. 111R) ......... ..................................................................................................................... Tricorythodes yura Tibias y tarsos blanquecinos con notorias marcas subapicales negruzcas al menos en las tibias (Figs. 111G-H); patrón de coloración formado por pequeñas máculas de pigmento distribuidas irregularmente (Fig. 110A) ................................................ 9 Tibias y tarsos sin tales marcas (Figs. 111I-J); pigmentos distribuidos uniformemente (Fig. 110B) ............................................................................................ 10

T. cristatus: el patrón de coloración abdominal no está descripto adecuadamente por Allen y los cuerpos de los paratipos no estuvieron asequibles al estudio.

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Plate 110. Tricorythodes, nymphs. Figs. 110A-V. Abdominal color pattern: 110A, T. arequita; 110B, T. barbus, 110C, T. bullus; 110D, T. zunigae. 110E, T. bullus, head and thorax, l.v.; 110F, T. cristatus, head and thorax, l.v.; 110G, T. ocellus, head, d.v.; 110H, T. santarita, abdomen, l.v. Mouthparts, T. hiemalis: 110I, right mandible; 110J, left mandible; 110K, labrum, dorsal at left, ventral at right; 110L, hypopharynx; 110M, labium, ventral at left, dorsal at right; 110N, maxillae. 110O, T. arequita, detail of maxillary palp; 110P, T. mirca, detail of maxillary palp. Abdominal gills, T. barbus: 110Q, gill II, d.v.; 110R-V, gills II-VI, v.v. (AE = Anteromedian emargination; LE = Laterally expanded segments; LO = Lateral ocelli; P = Maxillary palp; PS = Posterolateral spine; R = Dorsal ridge; T = Tubercles). Figs. 110A-D, H and O-V modified from MOLINERI, 2002; 110G from MOLINERI (2005); 110I-N from MOLINERI, 2001c.

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9(8)

Palpo maxilar trisegmentado (Fig. 110O); marcas negruzcas grandes presentes en tibias y tarsos (Figs. 111G-H) ......................................................... Tricorythodes arequita – Palpo maxilar bisegmentado con seta apical (Fig. 110P); marcas negruzcas presentes solo en tibias ........................................................................................... Tricorythodes mirca 10(8) Ocelos laterales grandes, casi del mismo tamaño que los ojos compuestos (Fig. 110G) .................................................................................................................. Tricorythodes ocellus – Ocelos laterales pequeños, al menos 40% más chicos que los ojos compuestos (Fig. 109A) ......................................................................................................................... 11 11(10) Segmentos abdominales III-VI expandidos lateralmente (Fig. 110B), espinas posterolaterales presentes en el segmento abdominal VII; uñas tarsales sin dentículos marginales, solo con un par de dentículos submarginales cerca del ápice (Fig. 111O) ..................................................................................................................Tricorythodes barbus – Segmentos abdominales III-VII expandidos lateralmente (similar to Fig. 110A); espinas posterolaterales presentes en los segmentos abdominales VII-VIII; uñas tarsales con 4 ó 5 dentículos marginales y un par de dentículos submarginales cerca del ápice (Fig. 111P) ............................................................................................. 12 12(11) Branquia opercular sombreada completamente con negruzco (Fig. 111A); proyección frontal presente ............................................................ Tricorythodes hiemalis – Branquia opercular sombreada completamente con negruzco, excepto en el ángulo anterolateral (Fig. 111C); proyección frontal ausente .................... Tricorythodes quizeri Tricorythodes arequita TRAVER Tricorythodes arequita TRAVER, 1959a: 128 (male); HUBBARD, 1982a: 274; MOLINERI, 2002: 275 (male, female, nymph).

Taxonomy: Tricorythodes arequita was described by TRAVER (1959a) for male subimagos from Uruguay. Recently MOLINERI (2002) described imagos of both sexes and nymphs from Northeast Argentina. This species can be distinguished from the others of the genus by the following combination of characters. In the imago: 1) Abdomen shaded with black almost completely, with small whitish dots (similar to Fig. 110A); 2) tibiae and tarsi whitish with conpicuous subapical blackish bands (Fig. 108F); 3) penes broad and flattened (Fig. 108I); 4) length of segment I/ length of segment II of forceps: 0.8; 5) vein CuP present. In the nymph: 1) Abdominal color pattern as in Fig. 110A; 2) operculate gill shaded with black as in Fig. 110A; 3) maxillary palp 3-segmented and short (Fig. 110O); 4) position of transverse row of setae on fore femora: submedian; 5) legs with subapical blackish marks on femora, tibiae and tarsi (Figs. 111G-H); 6) tarsal claws with 11-13 basal marginal denticles and a pair of submarginal denticles near apex. Distribution and Biology: Argentina (Misiones), Brazil (Rio Grande do Sul) and Uruguay (Lavalleja). Adults were collected flying in the early morning, but swarms were not observed. The nymphs were found in warm stony streams at 900 m of altitude.

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Tricorythodes australis (BANKS) Tricorythus australis BANKS, 1913: 85 (male). Leptohyphodes australis; ULMER, 1920a: 50; LESTAGE, 1931b: 60. Tricorythodes australis; TRAVER, 1958a: 501 (male, female); HUBBARD, 1982a: 274; MOLINERI, 2002: 278.

Taxonomy: This species originally described by BANKS (1913) was adequately redescribed by TRAVER (1958a) from imagos of both sexes. As TRAVER noted, the basal swelling of the segment II of forceps, so typical of Tricorythodes, is absent. This situation, along with the absence of the longitudinal vein CuP on male fore wings and the form and structure of penes, led TRAVER to suggest that T. australis was an aberrant member of Tricorythodes. The position of T. australis in the phylogeny proposed by MOLINERI (2002) is basal to all other species of Tricorythodes. Male imagos of T. australis can be distinguished from the other species of the genus by the following combination of characters: 1) Abdomen shaded uniformly with gray only on tergum I and median zone of terga II-VI; 2) tibiae and tarsi without blackish marks; 3) penes with a ventral projection covered with setae (Fig. 108H); 4) forceps: first segment with an apicomedial projection and second segment without basal swelling (Fig. 108H); 5) vein CuP absent in males, sometimes present but rudimentary in females. Distribution: Argentina (Misiones), Brazil (Pará, Paraná, Mato Grosso) and Guyana. Tricorythodes barbus ALLEN Tricorythodes barbus ALLEN, 1967: 373 (nymph); HUBBARD, 1982a: 274; MOLINERI, 2002: 278. Tricorythodes (Tricorythodes) barbus; ALLEN & MURVOSH, 1987: 36. Tricoryhyphes barbus; WIERSEMA & MCCAFFERTY, 2000: 353.

Taxonomy: Tricorythodes barbus is only known from nymphs and can be distinguished from the other species of the genus by the following combination of characters: 1) Abdomen with blackish marks as in Fig. 110B; 2) operculate gill triangular and blackish (Fig. 110Q); 3) maxillary palpi 3-segmented, with apical seta; 4) position of transverse row of setae on fore femora: subapical; 5) tibiae and tarsi without blackish marks (Figs. 111IJ); 6) tarsal claws with a pair of submarginal denticles (Fig. 111O); 7) very large body size (not fully grown nymphs have a length of 7 mm without cerci). Distribution: Argentina (Misiones), Brazil (Paraná). Tricorythodes bullus ALLEN Tricorythodes bullus ALLEN, 1967: 374 (nymph); HUBBARD, 1982a: 274; MOLINERI, 2002: 280 (male, female, nymph). Tricorythodes (Tricorythodes) bullus; ALLEN & MURVOSH, 1987: 36. Epiphrades bullus; WIERSEMA & MCCAFFERTY, 2000: 350.

Taxonomy: Tricorythodes bullus was described by ALLEN (1967) based on nymphs; later MOLINERI (2002) described male and female imagos. WIERSEMA & MCCAFFERTY (2000) included this species (and T. cristatus) in Epiphrades, a genus defined by the presence of dorsal tubercles on head and thorax, and operculate gills subrectangulate. In spite of this group

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represents probably a monophyletic group, we prefer to continue treating Tricorythodes as a unity, until a more detailed knowledge of the genus is attained. Accepting these “apical” groups (i.e. Epiphrades) as different taxa would assemble together the more basal species in non-natural groups (MOLINERI, 2002). Tricorythodes bullus can be distinguished from the other species of the genus by the following combination of characters: 1) Abdominal color pattern similar to Fig. 110C; 2) legs with grayish-black marks on apex of femora and base of tibiae (Fig. 108D); 3) penes more or less pyramidal (Fig. 108K); 4) length of segment I/ length of segment II of forceps: 1.0-1.1; 5) vein CuP incomplete (Fig. 108C). In the nymph: 1) Abdominal color pattern as in Fig. 110C; 2) operculate gills blackish (Fig. 110C); 3) maxillary palp 1-segmented, with apical seta; 4) position of transverse row of setae on fore femora: subbasal; 5) tibiae and tarsi without blackish marks; 6) tarsal claws with 8-11 marginal denticles; 7) tubercles present on head, pronotum and mesonotum (Fig. 110E). Distribution and Biology: Argentina (Misiones), Brazil (Paraná, Nova Teutonia). Adults and mature nymphs were collected in summer, in stony streams at 900 m. Tricorythodes cristatus ALLEN Tricorythodes cristatus ALLEN, 1967: 373 (nymph); HUBBARD, 1982a: 274; MOLINERI, 2002: 283. Tricorythodes (Tricorythodes) cristatus; ALLEN & MURVOSH, 1987: 38. Epiphrades cristatus; WIERSEMA & MCCAFFERTY, 2000: 350.

Taxonomy: Tricorythodes cristatus is presently known only from nymphs from Serra do Mar (SE of Brazil). It is related to T. bullus, and they can be separated because the tubercles on the thorax are much shorter in T. cristatus (Fig. 110F). It was included in Epiphrades by WIERSEMA & MCCAFFERTY (2000), classification not followed here for the reasons explained above for T. bullus. Nymphs of T. cristatus can be distinguished from the other species of the genus by the following combination of characters: 1) Abdominal color pattern “black with pale markings” (ALLEN, 1967: 373); 2) operculate gill shaded completely with black; 3) maxillary palp 1-segmented, with apical seta; 4) position of transverse row of setae on fore femora: subbasal; 5) legs without subapical blackish marks on tibiae or tarsi; 6) tarsal claws with 16-18 marginal denticles and without submarginal denticles. Distribution: Brazil (Serra do Mar). Tricorythodes hiemalis MOLINERI Tricorythodes hiemalis MOLINERI, 2001c: 61 (adults, nymph); MOLINERI, 2002: 285.

Taxonomy: Tricorythodes hiemalis is known from all the stages and can be distinguished from the other species of the genus by the following combination of characters. In the imago: 1) Abdominal segments I-II and VII-X shaded with heavier gray than that of segments III-VI; 2) femora without markings or with a grayish subapical band, tibiae and tarsi without blackish marks; 3) penes wide at base becoming abruptly narrower in distal half (Fig. 108L); 4) length of segment I/ length of segment II of forceps: 1.2; 5) vein CuP present. In the nymphs: 1) Abdominal color pattern as in imagos; 2) operculate gill triangular and blackish (Fig. 111A); 3) maxillary palp 2-segmented with apical setae (Fig. 110N); 4) position of transverse row of setae on fore femora: submedian; 5) tibiae

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B

D

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F

TR

J

I G

N

M

TR

L K

O SD

P

Q

R

MD SD

Plate 111. Tricorythodes, nymphs. Figs. 111A-R. Operculate gills, d.v.: 111A, T. hiemalis; 111B, T. popayanicus; 111C, T. quizeri; 111D, T. santarita; 111E, T. yura; 111F, T. zunigae. Fore- and hind legs: 111G-H, T. arequita; 111I-J, T. barbus; 111K-L, T. santarita; 111M-N, T. zunigae. Fore tarsal claws: 111O, T. barbus; 111P, T. quizeri; 111Q, T. popayanicus; 111R, T. yura. (MD = Marginal denticles; SD = Submarginal denticles; TR = Transversal row). Figs. 111H-R modified from MOLINERI, 2002.

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and tarsi without blackish marks; 6) tarsal claws with 3-4 marginal denticles and a single pair of submarginal denticles. Distribution and Biology: Argentina (Salta, Jujuy). Adults and mature nymphs of this species were collected in winter, when the population of the other species of the genus (T. popayanicus) inhabiting the same river were at a minimum. The nymphs of T. hiemalis preferred sandy patches downstream from large stones and showed whitish wingpads in both sexes when ready to molt to the subimago. Tricorythodes lichyi TRAVER Tricorythodes lichyi TRAVER, 1943: 83 (male, female); HUBBARD, 1982a: 274; MOLINERI, 2002: 305.

Taxonomy: Tricorythodes lichyi is known from imagos of both sexes, the original description is not sufficient to distinguish it from the other species, but because TRAVER described the presence of blackish marks on subapex of tibiae it may be related to T. arequita and T. mirca. Distribution: Venezuela. Tricorythodes mirca MOLINERI Tricorythodes mirca MOLINERI, 2002: 285 (adults, nymph).

Taxonomy: Tricorythodes mirca, known from all the stages, can be distinguished from the other species of the genus by the following combination of characters. In the imago: 1) Abdomen shaded almost completely with black, but with small unpigmented dots; 2) tibiae whitish with heavy subapical blackish bands (Fig. 108G); 3) penes broad and flattened (Fig. 108J); 4) length of segment I/ length of segment II of forceps: 0.8; 5) vein CuP present. In the nymph: 1) Abdominal color pattern formed by irregularly distributed pigments; 2) operculate gill shaded with black irregularly; 3) maxillary palp 2-segmented with apical seta (Fig. 110P); 4) position of transverse row of setae on fore femora: submedian; 5) legs with subapical blackish marks on femora and tibiae; 6) tarsal claws with 8-9 basal marginal denticles and 2 or 3 submarginal denticles near apex. Tricorythodes mirca is very closely related to T. arequita with a very similar coloration and male genitalia. The species can be distinguished in nymphs and adults because subapical blackish marks are present on the tibiae and tarsi in T. arequita but only on the tibiae in T. mirca. Distribution: Bolivia (Santa Cruz). Tricorythodes nicholsae WANG, SITES & MCCAFFERTY Leptohyphes nicholsae WANG, SITES & MCCAFFERTY, 1998: 69 (nymph). Asioplax nicholsae; WIERSEMA & MCCAFFERTY, 2000: 348. Tricorythodes nicholsae; MOLINERI, 2002.

Taxonomy: WIERSEMA & MCCAFFERTY (2000) included this species in Asioplax, a genus defined by: body flattened, posterolateral projections of abdominal terga VII and VIII longer than the medial length of respective terga, fore femora greatly expanded and bordered by setae, among other characters. Although Asioplax may represent a monophyletic group (MOLINERI, 2002), we prefer to continue treating Tricorythodes as a unit, to avoid the problems

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Plate 112. Tricorythodes santarita. Fig. 112A, nymphal habitus. Drawing by C. Molineri.

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discussed before (in taxonomic section of T. bullus). Tricorythodes nicholsae, known only from nymphs, can be distinguished from the other species of the genus by the following combination of characters: 1) Body very depressed and discoidal; 2) operculate gills suboval, with a pair of paler marks similar to Fig. 111D; 3) maxillary palp 1-segmented with apical seta; 4) position of transverse row of setae on fore femora: subbasal (similar to Fig. 111K); 5) tibiae and tarsi without blackish marks; 6) tarsal claws without marginal denticles, only with a pair of submarginal denticles near the apex; 7) femora very expanded, almost as long as wide (similar to Figs. 111K-L); 9) ventral lamellae of gill II present and longer than operculate lamellae; 10) median tubercles present on abdominal terga III-IX. This species is very similar to T. santarita but can be distinguished by characteristics 9 and 10 listed above. Distribution: Ecuador. Tricorythodes ocellus ALLEN & ROBACK Tricorythodes sp. ROBACK, 1966: 150 (nymph). Tricorythodes ocellus ALLEN & ROBACK, 1969: 378 (nymph); HUBBARD, 1982a: 274; MOLINERI, 2002: 288; MOLINERI, 2005. Tricorythodes (Tricorythodes) ocellus; ALLEN & MURVOSH, 1987: 38. Tricoryhyphes ocellus WIERSEMA & MCCAFFERTY, 2000: 353.

Taxonomy: Tricorythodes ocellus was described by ALLEN & ROBACK (1969) from nymphs and later MOLINERI (2005) described the probable male imagos. WIERSEMA & MCCAFFERTY (2000) included this species in Tricoryhyphes but we prefer to continue treating this species as Tricorythodes ocellus for the same reasons explained above for T. barbus. Tricorythodes ocellus is known from nymphs and adults and can be separated from the other species of the genus by the following combination of characters. In the imago: 1) Abdominal segments uniformly shaded with grayish-black, except the area previously covered by gills of nymph; 2) tibiae and tarsi without blackish subapical marks; 3) femora without markings; 4) length of segment I/ length of segment II of forceps: 1.5-1.7 (Figs. 108N-O); 5) vein CuP present. In the nymph: 1) Abdomen with irregular black markings on tergum VII and a median black macula on terga VIII to X (from original description); 2) operculate gill triangular and shaded almost completely with black; 3) maxillary palp 2-segmented; 4) position of transverse row of setae on fore femora: submedian; 5) tibiae and tarsi without blackish marks; 6) tarsal claws without marginal denticles, only with a pair of submarginal denticles near the apex; 7) lateral ocelli very large, similar in size to compound eyes (Fig. 110G). Distribution: Peru (Tingo María). Tricorythodes popayanicus DOMÍNGUEZ Tricorythodes popayanicus DOMÍNGUEZ, 1982: 331 (adults, nymph); DOMÍNGUEZ, 1984: 103; DOMÍNGUEZ et al., 1994: 101; MOLINERI, 2002: 288. Tricorythodes (Tricoryhyphes) popayanicus; ALLEN & MURVOSH, 1987: 38. Tricoryhyphes popyanicus [sic]; WIERSEMA & MCCAFFERTY, 2000: 353.

Taxonomy: Tricorythodes popayanicus is known from all the stages and can be distinguished from the other species of the genus by the following combination of characters. In the imago:

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1) Abdominal segments uniformly shaded with grayish-black, except on intersegmental membranes; 2) tibiae and tarsi without blackish subapical marks; 3) femora without markings or with a grayish subapical band; 4) length of segment I/ length of segment II of forceps: 0.81.2; 5) vein CuP present; 6) genitalia as in Fig. 108P. In the nymph: 1) Abdominal color pattern as in imago; 2) operculate gill triangular shaded with black on basal 1/4 (Fig. 111B); 3) maxillary palp 2-segmented with apical setae; 4) position of transverse row of setae on fore femora: submedian; 5) tibiae and tarsi without blackish subapical marks; 6) tarsal claws with 23 marginal denticles and 2-3 pairs of submarginal denticles (Fig. 111Q). Distribution and Biology: Argentina (Jujuy, Salta, Tucumán, Catamarca, San Luis, Córdoba), Bolivia (Bermejo). This is a very common species in the stony and sandy bottoms of unpolluted rivers and streams from western Argentina. Nymphs of different sizes and adults are encountered all around the year, but the main part of the population emerges in spring and summer. Tricorythodes quizeri MOLINERI Tricorythodes quizeri MOLINERI, 2002: 290 (adults, nymph).

Taxonomy: Tricorythodes quizeri is known from adults and nymphs. It can be distinguished from the other species of the genus by the following combination of characters. In the imago: 1) Abdominal segments I-II and VII-X shaded with gray heavier than segments III-VI; 2) femora without markings or with a grayish subapical band, tibiae and tarsi without blackish marks; 3) penes broad at base and narrowing in distal half (Fig. 108M); 4) length of segment I/ length of segment II of forceps: 0.9; 5) vein CuP present; 6) third segment of forceps with a median projection. In the nymphs: 1) Abdominal color pattern as in imagos; 2) operculate gills triangular and blackish, except whitish zone at base (Fig. 111C); 3) maxillary palp 2-segmented with apical seta; 4) position of transverse row of setae on fore femora: submedian; 5) tibiae and tarsi without blackish marks; 6) tarsal claws with 3-4 marginal denticles and a single pair of submarginal denticles. Distribution: Bolivia (Santa Cruz). Tricorythodes santarita TRAVER Tricorythodes santarita TRAVER, 1959a: 130 (female); PETERS, 1981: 216; HUBBARD, 1982a: 274; MOLINERI, 2002: 293 (male, nymph).

Taxonomy: Tricorythodes santarita, known from all the stages, can be distinguished from the other species of the genus by the following combination of characters: 1) Abdomen shaded with blackish and reddish; 2) without blackish marks on tibiae or tarsi but with notorious reddish bands on femora and tibiae (Fig. 108E); 3) penes wide at base and becoming thinner towards apical 1/3 (Fig. 108R); 4) length of segment I/ length of segment II of forceps: 1.5-1.6; 5) vein CuP may be present, incomplete or absent; 6) basal 2/3 of wings shaded with blackish gray (in some males this can be reduced to C and Sc areas). In the nymph (Fig. 112A): 1) Body very depressed and discoidal; 2) operculate gills suboval, with a pair of paler marks (Fig. 111D); 3) maxillary palp 1-segmented with apical seta; 4) transversal row of spines on fore femora subbasal (Fig. 111K); 5) tibiae and tarsi

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without blackish marks; 6) tarsal claws without marginal denticles, only with a pair of submarginal denticles near the apex; 7) very expanded femora, almost as long as wide (Figs. 111K-L); 9) ventral lamellae of gill II absent (only a small rest of the inferior lamella is present near the base of the operculate lamella); 10) median tubercles present on abdominal terga VI-IX (Fig. 110H). Distribution and Biology: Uruguay (Paysandú), Argentina (Misiones). Adults of this species were collected in high numbers at light traps, in summer season. The few nymphs collected were encountered in fast-flowing areas of a warm stony stream. Tricorythodes yura MOLINERI Tricorythodes yura MOLINERI, 2002: 296 (adults, nymph).

Taxonomy: Tricorythodes yura can be distinguished from the other species of the genus by the following combination of characters. In the imago: 1) Abdomen shaded very slightly with gray, mainly on terga I and VII-IX; 2) femora, tibiae and tarsi without blackish marks; 3) penes with a subapical constriction (Fig. 108Q); 4) length of segment I/ length of segment II of forceps: 1.1-1.2; 5) vein CuP present (Figs. 108A-B). In the nymph: 1) Abdominal coloring as in imago; 2) operculate gill triangular shaded with gray only at base (Fig. 111E); 3) maxillary palp 2-segmented with apical seta; 4) position of transverse row of setae on fore femora: submedian; 5) tibiae and tarsi without blackish marks; 6) tarsal claws with 3-4 marginal denticles and 1 pair of submarginal denticles near apex (Fig. 111R). T. yura is similar to T. popayanicus DOMÍNGUEZ but they can be separated by color pattern and tarsal claw denticulation. Distribution and Biology: Bolivia. The nymphs were entirely covered by sediment attached to the setae, giving them a soiled appearance. They were collected in a mountain rain forest stream at 500 m of altitude, and appeared mainly in sand patches among rocks in riffle areas. Imagos were attracted to light traps before sunrise. Tricorythodes zunigae MOLINERI Tricorythodes zunigae MOLINERI, 2002: 298 (male, nymph).

Taxonomy: Tricorythodes zunigae can be distinguished from the other species of the genus by the following combination of characters. In the imago: 1) Abdomen shaded with blackish on a broad median longitudinal band; 2) without blackish marks on tibiae or tarsi but with reddish pink bands on femora and tibiae; 3) penes wide at base and becoming thinner on apical 1/3 (Fig. 108S); 4) length of segment I/ length of segment II of forceps: 1.1; 5) vein CuP present; 6) membrane of wings shaded with blackish gray in C and Sc areas. In the nymph: 1) Abdomen shaded with blackish as in Fig. 110D; 2) operculate gills subquadrate (Fig. 111F); 3) maxillary palp 2-segmented with apical seta; 4) position of transverse row of setae on fore femora: subbasal (Fig. 111M); 5) tibiae and tarsi without blackish marks; 6) tarsal claws with 3-4 marginal denticles and with a pair of submarginal denticles near the apex; 7) body discoidal, very depressed; 8) femora very expanded, almost as long as wide (Figs. 111M-N). Tricorythodes zunigae is related to the T. santarita species group (MOLINERI, 2002) and can be distinguished from the other species of this

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group by its characteristic color pattern (Fig. 110D), absence of abdominal tubercles (Fig. 110D), subquadrate operculate gill (Fig. 111F) and longer maxillary palpi. Distribution: Colombia. Genus Tricorythopsis TRAVER Tricorythopsis TRAVER, 1958a: 491; HUBBARD, 1990: 88; DOMÍNGUEZ et al., 1992: 24; PETERS & PETERS, 1993: 46; MOLINERI, 1999b; WIERSEMA & MCCAFFERTY, 2000; MOLINERI, 2001d. (Type-species: Tricorythopsis artigas TRAVER, original designation).

Taxonomy: Tricorythopsis was established from imagos by TRAVER (1958a), nymphs were described by MOLINERI (1999b, 2001d). It is represented by 8 South American species. Adult Characteristics: 1) Fore wings of male with large cubito-anal lobe (Figs. 113A, C); 2) hind wings absent in both sexes; 3) female caudal filaments reduced, shorter than the last 3 abdominal segments combined; 4) membranous filaments of mesoscutellum short and relatively wide; 5) male forceps 2-segmented, apical segment outwardly directed (Figs. 113D-K); 6) penes almost completely fused, except for distal furrow, apical lobes of penes not strongly divergent; 7) male gonopore not associated with a spine; 8) penes without projections; 9) intercalary veins and longitudinal veins from sectors Cu and MP forming 2 consecutive triads (T1 and T2 in Fig. 113B); 10) styliger plate projected forming a columnar base for each forceps (SP in Fig. 113D); 11) female imagos generally without fore legs (except for coxae and trochanters), fore legs generally are lost with the subimaginal skin. Nymphal Characteristics: 1) Hind wingpads absent in both sexes; 2) operculate gill oval, without ridges, with a weaker transverse line near middle (WL in Figs. 115H-I); 3) gill formula 5/4/4/2/1 (Figs. 115I-M); 4) fore femora with a transverse row of short to long spines on dorsum (Fig. 116B); 5) middle and hind femora with transverse row of spines at base (Figs. 116C-D); 6) maxillary palpi reduced, 2-segmented with apical seta (Fig. 115G); 7) labrum with shallow anteromedian emargination (Fig. 115A); 8) tarsal claws with 2 parallel rows of submarginal denticles near the apex and almost always with a marginal row of denticles more basally (Fig. 116A); 9) small genal projections present, frontal projections absent; 9) nymphs relatively small and robust, generally less than 3 mm in length; 10) labium as in Fig. 115D, labial palpi straight. Distribution: Argentina, Uruguay, Brazil, Venezuela, Ecuador, Colombia, Guyana, French Guiana and Costa Rica. Biology: Nymphs of this genus were collected in various swift-current subtropical streams. They are mainly crawlers inhabiting among different sized sediments. The morphology of mouth parts as well as gut contents suggest they are gatherers of fine particulate organic matter, as is common in the family. Young nymphs, although in small number, appeared in hyporheic samples. Much care is needed to collect the hardly visible nymphs in the field, both because of their small size and their lack of movement in the net. The adults are so tiny that they may be overlooked in light traps, where they are attracted by the hundreds (at dusk and dawn). Nuptial flights of 2 species were observed (T. artigas and T. yacutinga). Both showed the typical up and down movement of individual males. Swarms were small but numerous, with no more than 50 males in each individual swarm. Males were observed to fly among different neighboring swarms. Swarms

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commonly were located 0.5-3 m above river banks. The males were active during the day time, mainly around noon. Females fly for a longer time than males, being observed to fly mainly upstream until dusk. Oviposition appears to be similar to other genera of Leptohyphidae, the eggs being extruded in a single mass that is retained below the apex of abdomen until the female drop to the water. FRANCISCHETTI et al. (2002) reported a similar nuptial flight of an undescribed species of Tricorythopsis, except that females also were actively present in the swarms and the flight period was later in the day (before dusk). KEYS

TO

TRICORYTHOPSIS

Adult Males 1 – 2(1) – 3(1) – 4(3) – 5(4) – 6(5) – 7(4) –

Penes relatively wide with lateral margins sclerotized, darker than the rest (Figs. 113F, I) ................................................................................................................................. 2 Penes slender, without lateral margins sclerotized (Figs. 113D-E, G-H, J-K) ...... 3 Hind femora (and sometimes middle femora) with a wide transverse blackish band (similar to Figs. 116S-T); genitalia as in Fig. 113I ......... Tricorythopsis undulatus Hind and middle femora with a wide transverse blackish band interrupted in the middle by a unpigmented zone (similar to femora in Fig. 114D); genitalia as in Fig. 113F ....................................................................................................... Tricorythopsis gibbus Forceps relatively short and robust, distal lobes of penes rounded, if acute never curved medially (Fig. 113D-E, G-H, K) ....................................................................... 4 Forceps long and slender; distal lobes of penes apically acute and curved medially (Fig. 113J) ............................................................................................Tricorythopsis volsellus Penes divided on apical 1/6 or less (Figs. 113E, G-H) ............................................. 5 Penes divided on apical 1/3 to 1/4 (Figs. 113D, K-L) .............................................. 7 Penes conical, with narrow distal zone (apical width 1/5 of basal width) (Figs. 113E, G); thoracic sterna shaded strongly with gray ................................................. 6 Penes cylindrical, with similar width along their length (Fig. 113H); thoracic sterna whitish, not shaded with gray ....................................................... Tricorythopsis sigillatus Abdominal and thoracic color pattern as in the nymph (Fig. 114C), generally with a well defined blackish mediolongitudinal line ....................... Tricorythopsis chiriguano Abdominal and thoracic color pattern as in the nymph (Fig. 114E), not as above .............................................................................................................. Tricorythopsis minimus Body almost completely shaded reddish, abdomen broadly shaded with reddish gray; penes relatively short and wide, with somewhat quadrangular base (Figs. 113K-L) ............................................................................................. Tricorythopsis yacutinga Body with reddish marks but less extensive; abdomen whitish with small blackish and reddish marks; penes slender with the base widening gradually (Fig. 113D) .................................................................................................................Tricorythopsis artigas

Nymphs 1

Body dorso-ventrally flattened (Fig. 114G); femora almost as wide as long and fringed by long spines (Figs. 116U-W) ....................................... Tricorythopsis yacutinga

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C

A

CuP

A CuP

ICu1

DL

MP MA2 CuA MP2 IMP 1

F II

B

FI SP

D

DF

T2 T1

E

DL SM

F

G

SM

I

H

J

K

L CB

Plate 113. Tricorythopsis, adults. Figs. 113A-L. T. artigas: 113A, male fore wing; 113B, female fore wing. T. gibbus: 113C, male fore wing. Male genitalia, v.v.: 113D, T. artigas; 113E, T. chiriguano; 113F, T. gibbus; 113G, T. minimus; 113H, T. sigillatus; 113I, T. undulatus; 113J, T. volsellus; 113K, T. yacutinga. 113L, detail of penes (T. yacutinga). (CB = Quadrangular base; DF = Distal furrow; DL = Distal lobes; FI, FII = Forceps segment; SM = Sclerotized margin; SP = Styliger plate; T1, T2 = triads). Figs. 113A-D, G-J, modified from MOLINERI, 1999b; 113E-F, K-L modified from MOLINERI, 2001d.

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–

5(4) –

Body not so depressed; femora longer than wide with short to long spines but never as above ..................................................................................................................... 2 Coxae with a dorsal projection (Figs. 116J-L, R-S); femoral spines relatively short (Figs. 116J, R); abdomen with median tubercles (Fig. 114D) or with undulated margins (Fig. 114F) ............................................................................................................ 3 Coxae without dorsal projections; femoral spines relatively long; abdomen without tubercles or undulated margins ........................................................................ 4 Distal part of mesonotum with median hump (Fig. 114D); median dorsal tubercles present on abdominal terga (Fig. 114D); legs as in Figs. 116J-L ............. ................................................................................................................. Tricorythopsis gibbus Mesonotum without a hump (Fig. 114F); without abdominal tubercles but with dorsal and lateral margins of abdominal terga undulated (Fig. 114F); legs as in Figs. 116R-T .................................................................................... Tricorythopsis undulatus Abdominal coloration as in Figs. 114C or E; femora almost without pigments; thoracic sterna shaded with gray .................................................................................... 5 Abdominal coloration as in Figs. 114A-B; femora tinged with yellowish as in Figs. 116B-D; thoracic sterna whitish ...................................................... Tricorythopsis artigas Abdominal and thoracic color pattern as in Fig. 114E ............ Tricorythopsis minimus Abdominal and thoracic color pattern as in Fig. 114C ......... Tricorythopsis chiriguano

CLAVES

PARA

2(1) – 3(2) – 4(2) –

TRICORYTHOPSIS

Machos Adultos 1 – 2(1) – 3(1) – 4(3) – 5(4) – 6(5)

Penes relativamente anchos con los márgenes laterales esclerosados, más oscuros que el resto (Figs. 113F-I) ................................................................................................ 2 Penes sin márgenes laterales esclerosados (Figs. 113D-E, G-H, J-K) .................... 3 Fémures posteriores y a veces los medios con una banda transversal ancha negruzca (similar a Figs. 116S-T); genitalia como en la Fig. 113I ........... Tricorythopsis undulatus Fémures medios y posteriores con una banda transversal ancha negruzca, interrumpida en el centro por una zona sin pigmento (similar a Fig. 114D); genitalia como en la Fig. 113F ........................................................................... Tricorythopsis gibbus Fórceps relativamente cortos y robustos, lóbulos distales de los penes redondeados o si aguzados nunca curvados medialmente (Figs. 113D-E, G-H, K) ................................................................................................................................................ 4 Fórceps largos y delgados; lóbulos distales de los penes aguzados apicalmente y curvados medialmente (Fig. 113J) ................................................. Tricorythopsis volsellus Penes divididos en el 1/6 apical o menos (Figs. 113E, G-H) .................................. 5 Penes divididos en 1/3 o 1/4 apical (Figs. 113D, K, L) ............................................ 7 Penes cónicos, con la zona distal delgada (1/5 del ancho basal) (Figs. 113E, G); esternos torácicos fuertemente sombreados con gris ................................................ 6 Penes cilíndricos, con un ancho similar en toda su longitud (Fig. 113H); esternos torácicos blanquecinos (sin sombreado) .................................... Tricorythopsis sigillatus Patrón de coloración abdominal y torácico como en la ninfa (Fig. 114C), generalmente con una línea mediolongitudinal negruzca .................................. Tricorythopsis chiriguano

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319

B

H

D

DT

C

G E

F UM

Plate 114. Tricorythopsis, nymphs, general aspect. Figs. 114A-G. 114A-B, T. artigas; 114C, T. chiriguano; 114D, T. gibbus; 114E, T. minimus; 114F, T. undulatus; 114G, T. yacutinga. (DT = Dorsal tubercles; H = Hump; UM = Undulated margins). Figs. 114A-G modified from MOLINERI, 2001d.

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– 7(4) –

Patrón de coloración abdominal y torácico como en la ninfa (Fig. 114E), no como arriba ................................................................................................... Tricorythopsis minimus Cuerpo casi completamente teñido de rojizo, abdomen ampliamente sombreado con gris rojizo; penes relativamente anchos y cortos, con base angulosa (Figs. 113K-L) ............................................................................................. Tricorythopsis yacutinga Cuerpo con manchas rojizas, pero más restringidas; abdomen blanquecino con algunas marcas negruzcas y rojizas; penes más largos y delgados con la base ensanchada paulatinamente (Fig. 113D) ........................................ Tricorythopsis artigas

Ninfas 1 – 2(1) – 3(2) – 4(2) – 5(4) –

Cuerpo muy achatado dorsoventralmente (Fig. 114G); fémures de todas las patas muy anchos (casi circulares) y bordeados por largas espinas (Figs. 116U-W) ........ ............................................................................................................. Tricorythopsis yacutinga Cuerpo no tan achatado; fémures más largos que anchos con espinas largas o cortas pero nunca dispuestas como en el caso anterior ............................................ 2 Coxas de todas las patas con proyecciones dorsales (Figs. 116J-L, R-S); espinas femorales relativamente cortas (Figs. 116J, R); abdomen con tubérculos (Fig. 114D) o márgenes ondulados (Fig. 114F) .................................................................... 3 Coxas sin proyecciones dorsales; espinas femorales relativamente largas; abdomen sin tubérculos ni márgenes ondulados .......................................................................... 4 Extremo del mesonoto elevado en forma de joroba (Fig. 114D); tubérculos dorsales presentes en los tergos abdominales (Fig. 114D); patas como en las Figs. 116J-L .................................................................................................... Tricorythopsis gibbus Extremo del mesonoto sin joroba (Fig. 114F); sin tubérculos en el abdomen, pero con los márgenes dorsales y laterales ondulados (Fig. 114F); patas como en las Figs. 116R-T .................................................................................... Tricorythopsis undulatus Coloración abdominal como en las Figs. 114C, E; fémures casi sin pigmentos; esternos torácicos sombreados con gris ........................................................................ 5 Coloración abdominal como en las Figs. 114A-B; fémures teñidos de amarillento como en las Figs. 116B-D; esternos torácicos blanquecinos ............. .................................................................................................................Tricorythopsis artigas Patrón de coloración abdominal y torácico como en la Fig. 114E ........................... .............................................................................................................. Tricorythopsis minimus Patrón de coloración abdominal y torácico como en la Fig. 114C ........................... ........................................................................................................... Tricorythopsis chiriguano

Tricorythopsis artigas TRAVER Tricorythopsis artigas TRAVER, 1958a: 492; HUBBARD, 1982a: 274; MOLINERI, 2001d: 221. Leptohyphes tinctus ALLEN, 1973b: 368; HUBBARD, 1982a: 274; HUBBARD, 1990: 88. Allenhyphes tinctus; WIERSEMA & MCCAFFERTY, 2000: 343. Tricorythopsis fictilis MOLINERI, 1999b: 290-294.

Taxonomy: This species is known from all the stages, and can be separated from the other species of the genus by the following combination of characters. In the imago: 1)

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Body with reddish and blackish marks; 2) posterolateral projections of styliger plate at forceps base long, similar in length to forceps segment I (Fig. 113D); 4) penes divided at apical 1/3- 1/4 (Fig. 113D); 5) lateral margins of penes not sclerotized; and 6) female cerci as long as last 3 abdominal segments combined. In the nymph: 1) Thorax and abdomen without tubercles (Figs. 114A-B); 2) coxae without projections, femora with relatively long spines (Figs. 116B-D); 3) tarsal claws with 8 marginal denticles and a double row of 3 + 6 submarginal denticles near apex (Fig. 116A); 4) abdominal coloration as in Figs. 114A-B. This species is similar to T. minimus, but can be distinguished because T. minimus has the mesothoracic sterna shaded heavily with gray. Distribution and Biology: Argentina (Misiones), Uruguay (Artigas), and Brazil (Rio Grande do Sul). Nymphs of T. artigas were collected between 500 and 1000 m, in small to medium-sized streams, with rocky substrate, shaded by the surrounding forest. Imagos were collected flying in the morning and at noon. Tricorythopsis chiriguano MOLINERI Tricorythopsis chiriguano MOLINERI 2001d: 223.

Taxonomy: Tricorythopsis chiriguano is known from all the stages and can be separated from the other species of the genus by the following combination of characters. In the imago: 1) Vein CuP generally fused with vein A at base; 2) coloration of body light, with some grayish marks; 3) meso- and metathoracic sterna shaded with gray; 4) legs whitish; 5) penes pyramidal (Fig. 113E). In the nymph: 1) Thorax and abdomen without tubercles; 2) coxae without projections, legs with long spines (Figs. 116E-G); 3) tarsal claws with 7-9 marginal denticles and a pair of subdistal rows of 3-4 + 4-6 submarginal denticles (Fig. 116H); 4) abdominal color pattern as in Fig. 114C. Distribution: Bolivia (Santa Cruz). Tricorythopsis gibbus (ALLEN) Leptohyphes gibbus ALLEN, 1967: 358; HUBBARD, 1982a: 273. Tricorythopsis gibbus; MOLINERI, 2001d: 225.

Taxonomy: Tricorythopsis gibbus is known from all the stages and can be separated from the other species of the genus by the following combination of characters. In the imago: 1) Vein CuP of fore wings sometimes free at base (arrow in Fig. 113C); 2) penes broad apically with darkened lateral margins (Fig. 113F); 3) body dark; 4) middle and hind femora tinged with blackish in a median band, interrupted medially by an unpigmented zone (similar to Fig. 114D). In the nymph: 1) Pair of submedian humps near apex of wingpads (Fig. 114D) and with median tubercles on abdominal terga V-IX; 2) coxae with a dorsal projection (Figs. 116J-L), femora with short spines (Fig. 116J); 3) tarsal claws with 4-5 marginal denticles and a double row of 1-2 + 2-3 submarginal denticles (Fig. 116I); 4) abdominal coloration as in Fig. 114D. Distribution and Biology: Argentina (Misiones), Brazil (Nova Teutonia). Nymphs of this species were collected in stony streams of Misiones (Northeastern Argentina). Stream beds were composed of rocks of different sizes, gravel and sand. Also a high

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content of clay (suspended in the water and forming river banks) was present. Nuptial flight was not observed, but females were commonly seen flying upstream. Tricorythopsis minimus (ALLEN) Leptohyphes minimus ALLEN, 1973b: 369; HUBBARD, 1982a: 274. Allenhyphes minimus; WIERSEMA & MCCAFFERTY, 2000: 343. Tricorythopsis minimus; MOLINERI, 2001d: 229. Leptohyphes viriosus ALLEN, 1973b: 369; HUBBARD, 1982a: 274; MOLINERI, 2001d: 229. Allenhyphes viriosus; WIERSEMA & MCCAFFERTY, 2000: 343. Tricorythopsis artigas TRAVER (partim); TRAVER 1958a: 492; MOLINERI, 1999b: 289.

Taxonomy: Tricorythopsis minimus is known from all the stages and can be separated from the other species of the genus by the following combination of characters. In the imago: 1) Vein CuP generally fused at base with vein A; 2) body coloration light with some grayish shading; 3) meso- and metathoracic sterna and abdominal sterna I-II shaded with gray; 4) legs whitish; 5) penes pyramidal (Fig. 113G). In the nymph: 1) Thorax and abdomen without tubercles (Fig. 114E); 2) coxae without projections, legs with relatively long spines (Figs. 116N-P); 3) tarsal claws with 7-9 marginal denticles and a pair of subdistal rows of 3-4 + 4-6 submarginal denticles (Fig. 116M); 4) abdominal color pattern as in Fig. 114E.

A

B

WL

H

I

E

F

J

G AS

K C

I

D

II

L

M Plate 115. Tricorythopsis, nymphs. Figs. 115A-M. Mouthparts, T. gibbus: 115A, labrum, left d.v., right v.v.; 115B, hypopharynx, v.v.; 115C, right mandible, d.v.; 115D, labium, right d.v., left v.v.; 115E, left mandible, d.v.; 115F, maxillae; 115G, detail of maxillary palp. Gills, T. chiriguano: 115H, gill II, d.v.; 115I-M, gills II-VI, v.v. (AS = Apical seta; I, II = Palp segments; WL = Weakness line). Figs. 115AM modified from MOLINERI, 2001d.

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Distribution and Biology: Argentina (Misiones), Uruguay (Artigas), and Brazil (Rio Grande do Sul). This species was present in the same habitats reported for T. artigas and T. gibbus. Tricorythopsis sigillatus MOLINERI Tricorythopsis sigillatus MOLINERI, 1999b: 296.

Taxonomy: Tricorythopsis sigillatus is only known from male imagos and can be distinguished from the other species of the genus by the following combination of characters: 1) Vein CuP generally fused at base with vein A; 2) body light colored with some grayish marks; 3) posterolateral projections of styliger plate at forceps base long, similar in length to forceps segment I (Fig. 113H); 4) penes divided on apical 1/6 or less (Fig. 113H); 5) penes cylindrical, tapering gently from base to apex (Fig. 113H); 6) penes without lateral margins sclerotized. Distribution: Brazil (Rio de Janeiro). Tricorythopsis undulatus (ALLEN) Leptohyphes undulatus ALLEN, 1967: 357; HUBBARD, 1982a: 274; MOLINERI, 2001d: 232. Tricorythopsis undulatus; MOLINERI, 2001d: 232. Tricorythopsis petersorum MOLINERI, 1999b: 298.

Taxonomy: Tricorythopsis undulatus is known from all the stages and can be separated from the other species of the genus by the following combination of characters. In the imago: 1) Vein CuP of fore wings sometimes free at base (similar to Fig. 113C); 2) body dark without heavy marks; 3) posterolateral projections of styliger plate at base of forceps short, as long as half length of forceps segment I (Fig. 113I); 4) penes divided on apical 1/ 3-1/2 (Fig. 113I); 5) lateral margins of penes darkened, more sclerotized than rest. In the nymphs: 1) Thorax and abdomen without tubercles, but with undulating lateral margins (Fig. 114F); 2) coxae with dorsal projections, femora with short spines (Figs. 116R-T); 3) tarsal claws with 4-6 marginal denticles and a pair of submarginal denticles near apex (Fig. 116Q); 4) abdominal coloration as in Fig. 114F. Distribution: Argentina (Misiones), and Brazil (Paraná). Tricorythopsis volsellus MOLINERI Tricorythopsis volsellus MOLINERI, 1999b: 294.

Taxonomy: Tricorythopsis volsellus is known from male and female imagos that can be distinguished from the other of the genus by the following combination of characters: 1) Vein CuP generally fused at base with vein A; 2) body with blackish marks; 3) lateral projections of styliger plate at base of forceps short, less than half length of forceps segment I (Fig. 113J); 4) forceps long and slender (Fig. 113J); 5) lobes of penes apically acute and medially curved (Fig. 113J); 6) lateral margin of penes not sclerotized; and 7) female cerci shorter than abdominal tergum X. Distribution: Venezuela.

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SR MR

A

TR

F

TR

E G

B C D

H

I

M

N

O

P

J K L Q

R U S

V

W X

T Plate 116. Tricorythopsis, nymphs. Figs. 116A-X. Fore, middle, hind legs, and fore tarsal claws: 116AD, T. artigas; 116E-H: T. chiriguano; 116I-L, T. gibbus; 116M-P, T. minimus; 116Q-T, T. undulatus; 116UX, T. yacutinga. (MR = Marginal row; SR = Submarginal rows; TR = Transversal row; lower arrows in 116J, R = details of fore femoral spines; upper arrows in 116J-L, R, S = coxal dorsal projections). Figs. 116A-X modified from MOLINERI, 2001d.

Tricorythopsis yacutinga MOLINERI Tricorythopsis yacutinga MOLINERI, 2001d: 233.

Taxonomy: Tricorythopsis yacutinga is known from all the stages and can be separated from the other species of the genus by the following combination of characters. In the imago: 1) Vein CuP generally fused at base with vein A; 2) body almost completely tinged with reddish gray; 3) penes divided on apical 1/3-1/4 and with base abruptly truncated (Figs. 113K-L); 4) lateral margin of penes without dark pigments. In the nymph: 1) Body depressed, disk-shaped (Fig. 114G); 2) femora wide, subcircular, fringed by long spines

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(Figs. 116U-W); 3) without tubercles on body or legs; 4) abdominal color pattern as in Fig. 114G; 5) tarsal claws without marginal denticles and with a double subdistal row of 4+6 submarginal denticles (Fig. 116X). The nymphs of this species may be confused with those of Tricorythodes santarita Traver, because their general aspect and their distribution are similar. As they pertain to different genera, use of the generic key will easily distinguish them. Distribution and Biology: Argentina (Misiones, Corrientes). This species shows an extremely depressed body form, but its microhabitat could not be determined because nymphs were collected in very low numbers. Bodies of some nymphs were completely covered with fine particles of clay. This species co-exists with the other 4 species recorded from Misiones. Adults perform a nuptial flight as already described in the generic section. Genus Vacupernius WIERSEMA & MCCAFFERTY Vacupernius WIERSEMA & MCCAFFERTY, 2000: 345 (adult and nymph); EMMERICH, 2004: 105. (Type-species: Leptohyphes packeri ALLEN, original designation).

This genus is known from 3 species which have been recorded from the Antilles and North and Central America (WIERSEMA & MCCAFFERTY, 2000). Recently, EMMERICH (2004) reported a male subimago from an undescribed species of the genus from Colombia, and we also saw subimagos from Bolivia (Cochabamba). Genus Yaurina MOLINERI Yaurina MOLINERI, 2001a: 337 (adults, nymph). (Type-species: Yaurina mota MOLINERI, original designation).

Taxonomy: MOLINERI (2001a) established the genus Yaurina for 3 new species: Y. mota, Y. yapa, and Y. yuta. Later this author (MOLINERI, 2004) included Yaurina ralla which had previously been described in Leptohyphes by ALLEN (1973b, as L. rallus), and known only from nymphs. This genus is more closely related to Allenhyphes than to any other genus of Leptohyphidae. They share a similar male genitalia and, in the nymphs, the presence of a single row of subapical denticles (generally double in others). The nymphs of 3 species are known but they are indistinguishable, thus only a key to the male adults is given. Adult Characteristics: 1) Fore wings of male without developed cubito-anal lobe (Fig. 117B); 2) hind wings present in males, with 2 longitudinal veins (Figs. 117C-D), hind wings absent in females; 3) female caudal filaments relatively long; 4) membranous filaments of mesoscutellum long and slender; 5) male forceps 3-segmented; 6) penes fused except for apical excavation (Figs. 117G, J, M) and with a lateral groove as in Fig. 117F; 7) male gonopore associated with a lateral hollow-filament attached at the ventral base of penes (PF in Figs. 117F, M); 8) penes without subapical and dorsal membranous projections; 9) apical hind margin of styliger plate, between forceps, with a pair of small median projections (MP in Figs. 117E-F); 10) base of male terminal filament without ventral spine. Nymphal Characteristics: 1) Hind wing pads present in males, absent in females; 2) operculate gill suboval, with 2 slightly marked dorsal ridges (Fig. 119L); 3) gill formula 3/3/

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A

B C

D

MP

I

E

L LG

F P PF PF

G

MP

H

M

J

K

Plate 117. Yaurina, adults. Figs. 117A-M. Y. mota: 117A, female fore wing; 117B, male fore wing; 117C, male hind wing; 117D, same, enlarged; 117E, male genitalia, v.v.; 117F, same, l.v.; 117G, detail of penes, v.v.; 117H, transverse cut of penes. Y. yuta: 117I, male genitalia, v.v.; 117J, detail of penes, v.v.; 117K, transverse cut of penes. Y. yapa: 117L, male genitalia, v.v.; 117M, detail of penes, v.v. (LG = Lateral groove of penes; MP = Medial projections; P = Penes; PF = Penes filaments). Figs. 117AM modified from MOLINERI, 2001a.

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3/3/1 (Figs. 119M-Q), lobes suboval; 4) fore femora with transverse row of long spines (Fig. 119H); 5) middle and hind femora without transverse basal row of spines at dorsum (Figs. 119I-J); 6) maxillary palpi small, 2-segmented (Fig. 119C); 7) labrum with broad anteromedian emargination (Fig. 119A); 8) mandibles as in Figs. 119D and G, labium as in Fig. 119E, hypopharynx as Fig. 119B; 9) tarsal claw with 4-7 marginal denticles and a single palisade of submarginal denticles (Fig. 119K); 9) genal projections present, frontal projections absent. Distribution: NW Argentina, Bolivia, Peru and Ecuador. Biology: The species of the genus have a rather restricted distribution along the eastern slopes of the Andes, from 500 to 1300 m of altitude. Nymphs were collected among gravel and rocks in riffle zones of rivers and streams. Adults were observed swarming at early morning. KEY

TO

YAURINA

Adult Males 1 – 2(1) –

CLAVE

Total length of filaments of penes longer than penes, and curled (Figs. 117L-M) ............................................................................................................................. Yaurina yapa Filaments of penes shorter than penes, apex of filaments slightly recurved at apex or straight (Figs. 117E, I) ................................................................................................. 2 Filaments of penes with apex acutely pointed and outwardly directed (Figs. 117I, J); penes with a marked ventral excavation (Fig. 117K) ......................... Yaurina yuta Filaments of penes with blunt apex, not curved as above (Figs. 117E-G); penes not as above (Fig. 117H) .............................................................................. Yaurina mota PARA

YAURINA

Machos Adultos 1 – 2(1) –

Filamentos esclerozados de los penes más largos que los penes (si estuviesen estirados) y formando un bucle distal Figs. 117L-M) ............................. Yaurina yapa Filamentos esclerozados más cortos que los penes, pueden estar levemente curvados en el ápice (Figs. 117E, I) ............................................................................... 2 Espinas con el ápice agudo y curvado hacia fuera (Figs. 117I, J); penes con una excavación ventral marcada (Fig. 117K) .................................................... Yaurina yuta Espinas con el ápice romo, no curvado como en el caso anterior (Figs. 117E-G); penes no como arriba (Fig. 117H) ............................................................. Yaurina mota

Yaurina mota MOLINERI Yaurina mota MOLINERI, 2001a: 342.

Taxonomy: This species is known from adults and nymphs of both sexes. Only the male can be distinguished from the other species of the genus, based on males of Y. mota have short filaments on penes, that are apically blunt (Figs. 117E-H). Distribution and Biology: Argentina (Jujuy). This species is known from a single locality in NW Argentina, a very small stream (1 m wide, 1250 m) that is completely

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A

Plate 118. Yaurina yuta. Fig. 118A, nymphal habitus, modified from MOLINERI, 2001a.

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covered by the canopy of a subtropical rain forest (Yungas). Substrate is composed of sand and rocks of different sizes, with numerous fallen trees. Adults performed the nuptial flight for a few minutes at early morning, before sunrise. Small swarms were observed at a height of 1-2 m above the stream, individual males flew up and down about 30 cm. Light traps attracted subimagos at dusk and imagos before sunrise. Yaurina ralla (ALLEN) Leptohyphes rallus ALLEN, 1967: 363; HUBBARD, 1982a: 274. Allenhyphes rallus; WIERSEMA & MCCAFFERTY, 2000: 343. Yaurina ralla; MOLINERI, 2004: 216.

Taxonomy: Leptohyphes rallus ALLEN (1967) was described from a single nymph from Tingo María (Huánuco, Peru). As the species did not have the diagnostic characters of Leptohyphes, WIERSEMA & MCCAFFERTY (2000) moved it to Allenhyphes. Later, MOLINERI (2004) stated that it better fits characters of Yaurina, mainly because of the characteristic maxillary palpi (2-segmented, distal segment basally widened as in Fig. 119C). Other characters shared with Yaurina are: 1) Submentum rounded and wide; 2) tarsal claw with 4 marginal denticles and a single palisade of 6 submarginal denticles; 3) abdominal gills II ovoid without marked dorsal ridges; 4) body as in Fig. 118A. Distribution: Peru (Tingo María). Yaurina yapa MOLINERI Yaurina yapa MOLINERI, 2001a: 344.

Taxonomy: This species is known from a single male subimago with remarkable genitalia which allow a rapid identification. The length of the filaments of the penes does not change with the molt to imago in the other species of the genus, and is expected that imaginal penes of Y. yapa will be similar to those of the subimago. Male adults of Y. yapa can be separated from the other species of the genus by the long curved filaments of the penes (total length greater than length of penes) and because they are curled as in Figs. 117L-M. Distribution: Ecuador. Yaurina yuta MOLINERI Yaurina yuta MOLINERI, 2001a: 341.

Taxonomy: This species is known from all stages but only male adults can be distinguished from the other species of the genus, by the following character: 1) Filaments of penes curved apically, and shorter than penes (Figs. 117I-K). Distribution and Biology: Argentina (Salta), Bolivia. Nymphs of this species were collected in small streams (2 m wide) and rivers (15 m wide). They were found in the stony substrate of riffles. Subimagos emerge in the afternoon, molting to imago during the night. Male imagos were captured while swarming during early morning. The swarm was located 1-2 m above the river, each male flew the usual up and down pattern.

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B

A

C

D

F

E

G

M H L N I

O

P SP MD

Q J

K

Plate 119. Yaurina, nymphs. Figs. 119A-Q. Y. mota: 119A, labrum, d.v.; 119B, hypopharynx, v.v.; 119C, detail of maxillary palp; 119D, right mandible; 119E, labium, v.v.; 119F, maxilla; 119G, left mandible; 119H, fore leg; 119I, middle leg; 119J, hind leg; 119K, detail of fore tarsal claw; 119L, gill II, d. v.; 119M-Q, gills II-VI, v.v. (MD = Marginal denticles; SP = Submarginal palisade). Figs. 119AQ modified from MOLINERI, 2001a.

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FAMILY LEPTOPHLEBIIDAE Taxonomy: This group was first recognized by EATON (1883-1888) as “Section 5 of Leptophlebia”, and the authorship of the family name was established by BANKS (1900), as Leptophlebiini. This is probably the most diverse family in the Neotropics, followed closely only by Baetidae. At present, the family includes in South America approximately 40 genera and 150 species, all belonging to the subfamily Atalophlebiinae. Of these genera, several are monotypic, while others are species-rich; similarly, some genera are known only from the type locality or have a very restricted distribution, while others extend from Central Argentina to central Canada. Adult Characteristics: 1) Eyes of male divided into dorsal and ventral portions (with one exception in Africa) (Figs. 137A, 165L-Q, 192A); 2) two to four long intercalaries between CuA and CuP (Figs. 131A, 154A); 3) hind wings present and well developed to small or absent (Figs. 120B, 123B, 137C); 4) tarsi of fore legs of male 5-segmented, segment 1 short; tarsi of middle and hind legs of male 4segmented; tarsi of all legs of female 4-segmented; 5) forceps of the male genitalia 2 to 4-segmented, segments II and III or segment II (if III absent) shorter than segment I (Figs. 120C, 121C, 137I) (segment IV present in one North American genus only); 6) penis lobes divided to completely fused (Figs. 120C, 159L, 187N); 7) female sternum IX entire to deeply cleft apically (Figs. 123O, 154G, 159E); 8) well developed egg-guide present (Figs. 121D, 140D) or absent; and 9) caudal filaments well developed, usually 3, but the median terminal filament sometimes reduced. The square facets of the dorsal portion of the male eyes (Figs. 140C, 162G-H, 165O-P), a defining character of the subfamily Atalophlebiinae, is another useful character for recognizing the family in South America; other subfamilies and other mayflies have typical hexagonal eye facets. Nymphal Characteristics: 1) Head hypognathous or prognathous; 2) hypopharynx well developed, usually with well developed lateral processes (Figs. 133E, 138B); 3) maxillary palpi 3-segmented (Figs. 186E, 198F); 4) labial palpi 3-segmented (Figs. 140I, 152D); 5) clypeus fused with frons, although fusion sometimes incomplete; 6) gills present on abdominal segments I-VII, I-VI or II-VII, gills formed by plate-like or slender dorsal and ventral lamellae (ventral portion sometimes absent) (Figs. 122E, 136D, 153G, 191FH); and 7) 3 well developed caudal filaments present. Distribution: Worldwide. Biology: The nymphs of this family are very common in South American rivers and streams. They can be found at altitudes from sea level to up to 2000 m. The nymphs are generally prognathous (Figs. 127A, 143A), although a few are hypognathous (Fig. 186A). The species are mainly clingers or sprawlers, although the hypognathous species seems to be slow swimmers. The nymphs have generalized herbivorous mouthparts, getting their food as collectors-gatherers, collector-filterers or shredders-detritivores. The subimagos generally emerge at dusk, molting within 24 hours, often less. Swarming takes place in early morning or late afternoon, although several species seem to fly during the night, as they have never been observed swarming, but have been attracted by lights.

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KEYS TO SOUTH AMERICAN GENERA Adults19 l – 2 (1)

– 3 (1) – 4 (3)

– 5 (4) – 6 (5) – 7 (4) – 8 (7) – 9 (8)

Hind wings absent ......................................................................................................... 2 20 Hind wings present ............................................................................................................ 3 Vein ICu1 of fore wings joined at base to CuA (Fig. 121A); male styliger plate subequal in length and width (Fig. 121C); each penis lobe lacking ventral spine; female egg guide (Fig. 121D) rounded apically, not reaching middle of sternum IX ...................................................................................................................................... Askola Vein ICu1 of fore wings joined at base to CuP (Fig. 140A); male styliger plate half as long as wide (Fig. 140B); each penis lobe with ventral spine; female egg guide acute apically, extending beyond middle of sternum IX ........................ Hagenulopsis Tarsal claws of each pair similar, both acute (Figs. 137H, 175H) ........................... 4 Tarsal claws of each pair dissimilar, one acute, other blunt (Figs. 164L, 187H) ....... 13 Vein Sc of hind wings ending close to apex of costal projection (Figs. 137D, 185B); penes short, triangular, lobes close together apically and fused basally (Figs. 137I, 185E); stem of vein MA sagged so forks sometimes slightly asymmetrical (Fig. 185A) .................................................................................................. 5 Vein Sc of hind wings extending well beyond weak costal projection (Figs. 171B, 175D); penes long, or broad (never triangular), partially or completely divided (Figs. 171F-J, 175J-L); fork of MA symmetrical (Figs. 171A, 175C) ...................... 7 Costal projection of hind wings blunt, rounded, barely projecting from costal margin ........................................................................................................................ Terpides Costal projection of hind wings long, thumb-like (Figs. 137C, D, F, G) ............... 6 Dorsal portion of male eyes on long stalk (Fig. 137A); forceps of males bisegmented (Fig. 137I) ......................................................................................... Fittkaulus Dorsal portion of male eyes on short, wide stalk (Fig. 192A) or stalk absent; forceps of males three-segmented (Fig. 192F) .............................................. Tikuna 21 Cross veins in apical 1/3 of costal area of fore wings anastomosed (Fig. 142A) ............................................................................................................................. Hapsiphlebia Cross veins in costal area of fore wings not anastomosed (Figs. 154A, 156A) ....... 8 Vein ICu1 joining CuA directly (Fig. 179A) .................................................................. 9 Base of vein ICu1 not as above; free in membrane (Fig. 171A), joining veins CuA and CuP by cross veins (Fig. 156A), or apparently joining CuP directly (Fig. 154A) .............................................................................................................................................. 10 Vein MP2 of fore wings strongly recurved, base of MP2 closer to vein CuA than to vein MP1, occasionally joining vein CuA (Fig. 175C); fore wings of male lacking spots on fork of MA; each lobe of penes with apical spine (Figs. 175J, O) .......... ............................................................................................................................... Penaphlebia

Although this key is intended for males, it can help in certain cases to determine females too. The adults of Bessierus and Perissophlebiodes are not known, but the nymphs do not have hind wing pads, and for this reason they should key here. 21 No species of this genus has been recorded from South America yet, but we have seen undescribed specimens from Colombia. 19 20

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10 (8) – 11 (10) – 12 (11) – 13 (3) – 14 (13) – 15 (14) – 16 (15) – 17 (16) – 18 (17) – 19 (18) –

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Vein MP2 of fore wings moderately recurved, base of vein MP2 between veins CuA and MP1 (Fig. 179A); fore wings of male with small spots on fork of MA (sometimes present also in the females); each lobe of penes with lateral spine (Fig. 179C) ...................................................................................................................... Rhigotopus Vein MP2 of fore wings strongly recurved; vein ICu1 connected to veins CuA and CuP at base by a cross vein (Fig. 154A) ........................................................ Massartella Vein MP2 of fore wings moderately recurved; vein ICu1 connected to veins CuA and CuP at base by cross veins (Fig. 156A), or base of vein ICu1 free in membrane (Fig. 171A) ......................................................................................................................... 11 Penes totally divided, tubular, with apical projection on each lobe (Fig. 156D); vein Sc of hind wings 9/10 total length of wing (Fig. 156B) ............ Massartellopsis Penes fused in basal 1/2 to 2/3 (Fig. 120C); vein Sc of hind wings 3/4 to 4/5 total length of wing (Fig. 120B) ............................................................................................. 12 Penes fused in basal 1/2, each lobe with long and thick spine medially directed (Fig. 120C) .................................................................................................... Archethraulodes Penes fused in basal 2/3, each lobe with short sclerotized process on internal margin, never shaped as spine (Figs. 171F-J) ......................................................Nousia Fork of vein MA of fore wings distinctly asymmetrical (Figs. 144A, G, 164A, D) .... 14 Fork of vein MA of fore wings symmetrical (Figs. 187B, D), or stem of MA sagged so fork appears slightly asymmetrical (Figs. 131A, D) ............................... 26 Costal projection of hind wings well developed, distal margin forming almost square angle; vein Sc ending in cross vein or at costal projection (Figs. 144C, I, 147C, 183C) ....................................................................................................................... 15 Costal projection of hind wings tent-shaped and projecting from costal margin basal to point where vein Sc meets vein C (Figs. 164B-C, E) ................................ 24 Second segment of genital forceps short, no more than 1.5 times as long as wide (Figs. 134D, 144M); male eyes frequently enlarged but not on stalk (Fig. 183D) ............... 16 Second segment of genital forceps long, at least 3 times as long as wide (Fig. 147G); male eyes on stalk; small species (body approximately 5 mm) ................ 25 Posterolateral angles of male styliger plate projected posteriorly; lobe of each penis with ventral appendix, generally laterally directed (Figs. 134D, F, J) ............ Farrodes Posterolateral angles of male styliger plate not projected as above; lobe of each penis with ventral projections not laterally directed (Figs. 144J-M, 148D, 174E) ............... 17 Male styliger plate lacking projections on posterior margin (Figs. 148D, 151D) ...... 18 Male styliger plate with one or two projections on posterior margin (Figs. 174E, 193F) ................................................................................................................................... 20 Penes divided in apical 1/4, with ventral projection shaped as curved spine, divergent apically (Fig. 151D) ................................................................................Hylister Penes divided in apical 1/2 or 2/3, ventral projection of each lobe not as above (Figs. 148D, 183G) ........................................................................................................... 19 Costal projection of hind wings strongly developed (Figs. 183B-C); part of costal area strongly tinged (Fig. 183C) ............................................................... Simothraulopsis Costal projection of hind wings not so developed; hind wings lacking strong pigmentation (Figs. 148B-C) ........................................................................Homothraulus

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20 (17) Male styliger plate with single median projection, dorsally projected between penes (Fig. 174F); each penis lobe with spine-like projection, anteromedially directed (Fig. 174E) ............................................................................................ Paramaka – Male styliger plate with paired projections (Figs. 150E, 169E); projections of penis lobes not as above (Figs. 150E, 169E, 193F) .................................................. 21 21 (20) Penes with ventral, straight spine-like projections, directed toward median line of body; male styliger plate with blunt projections (Figs. 150E, 193F) ..................... 22 – Penes with curved spine-like projections or lobe-like projections; projections of male styliger plate variable (Fig. 169E) ........................................................................ 23 22 (21) Projections of male styliger plate with wide base (South American species) (Fig. 193F) ......................................................................................................... Traverella (Zonda) – Projections of male styliger plate with narrow base (South American species) (Fig. 150E) .............................................................................................................. Hydrosmilodon 23 (21) Ventral projections of penes narrow, spine-like and divergent apically (Fig. 169E); projections of male styliger plate shaped as acute spines, and located close to base of forceps .......................................................................................................... Needhamella – Projections of penes and styliger plate variable, but not in combination mentioned above (Figs. 144J-M) .....................................................................Hermanella 24 (14) Tarsal claws of each fore leg of male different in shape than those of middle and hind legs; one fore claw blunt, other acute, spine-like (Fig. 164L) ............ Miroculis – Tarsal claws of each fore leg of male similar in shape to those of middle and hind legs; in all legs one claw blunt, other acute (Fig. 168D) ............................ Miroculitus 25 (15) Male styliger plate with two small median projections (Fig. 147G); costal projection of hind wings acute (Figs. 147B-C); shape of hind wings as in Figs. 147B-C .......... .......................................................................................................................... Hermanellopsis – Male styliger plate rounded (Figs. 162A-B); costal projection of hind wings rounded (Figs. 162D-E); shape of hind wings as in Figs. 162D-E ....... Microphlebia 26 (13) Hind wings very small, less than 1/10 length of fore wings, with strong costal projection that extends beyond apex of hind wing; posterior part of hind wings reduced (Figs. 141B-C) ...................................................................................... Hagenulus – Hind wings variable, 1/10 to 1/4 length of fore wings, costal projection variable, never extending beyond apex of wing (Figs. 123C, F, 128B, 187C, E); posterior part of hind wings not reduced .................................................................................... 27 27 (26) Lobes of penes fused except for apical notch, often with small apical spine (Figs. 159H, M) ............................................................................................................. Meridialaris – Lobes of penes totally divided, or fused in basal 1/2 or 1/3 (Figs. 123J, 196N, 199G) .................................................................................................................................. 28 28 (27) Basal segment of male forceps long and thin, with basal 1/6 forming strong angle on internal margin (Figs. 199C, G); fore wings with cross vein forming angle of approximately 45° between veins MA1 and R4+5 distal to fork of veins MA (Figs. 199A, E) ............................................................................................................................. 29 – Basal segment of male forceps not as above (Figs. 128D, 187J); fore wings with cross vein between veins MA1 and R4+5 variable, if present, angle usually greater than 45° (Figs. 123A, D, 187B, D) ............................................................................... 30

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29 (28) Vein Sc of hind wings 9/10 length of wing (Figs. 199B, F); membrane of wings with dark maculae at cross veins (Figs. 199A, E), apex of each penis round, lobe with ventral digitiform projection (Figs. 199C, G) ....................................... Ulmeritus – Vein Sc of hind wings 8/10 or less length of wing (Fig. 196B); wings without color pattern, except in costal membrane (Fig. 196A); apex of each penis rather straight, lobe with one or more spines (Figs. 196F, I, O, R) ................... Ulmeritoides 30 (28) Penes without ventral or ventrolateral projections (Figs. 128D, 181C) ................ 31 – Penes with strong ventral or ventrolateral projections (Figs. 123H-123M, 131F, 187J) .................................................................................................................................... 33 31 (30) Penes acute (Fig. 126C); fore wings with 3 or 4 intercalaries in cubital area (Fig. 126A) (Colombia) ...............................................................................................Choroterpes – Penes long, rounded, or somewhat quadrangular apically (Figs. 128D, 181C); fore wings with 2 long intercalaries in cubital area (Figs. 128A, 181A) ........................ 32 32 (31) Apex of each penis without spines, apex rounded, with semicircle of subapical setae dorsally (Fig. 128D) ............................................................................. Demoulinellus – Apex of each penis with short spines, apical setae and serrations on inner subapical margin (Fig. 181C) ............................................................................... Secochela 33 (30) Lobe of each penis with apical or subapical spine-like projection mesally directed (Figs. 187J, N, P); male styliger plate apically acute, sockets of forceps fused, forming single cavity (Figs. 187J, N, 188D) .................................................. Thraulodes – Lobe of each penis with subapical projection directed ventrolaterally (Figs. 123HM, 131F-G); male styliger plate rounded apically, sockets of forceps not fused (Figs. 123H-M, 131F) ...................................................................................................... 34 34 (33) Lobes of penes completely divided, with acute projections (Figs. 131F-G); wings with spots and pigmentation more pronounced at bullae (Figs. 131A, D) ......... Ecuaphlebia – Lobes of penes fused in basal 1/3, forming single round structure with blunt ventral projection (Figs. 123H-M); wings with spots on basal part of costal area or without spots (Figs. 123A, D) ...................................................................... Atopophlebia Nymphs 1 – 2 (1)

– 3 (2) –

Hind wing pads absent .................................................................................................... 34 Hind wing pads present .................................................................................................... 2 Body hydrodynamic (Fig. 186A); head hypognathous; caudal filaments with swimming setae (basal 2/5 to 3/5 of filaments with long setae on inner margin of cerci and lateral margins of terminal filament); lingua of hypopharynx without lateral projections (Figs. 138B, 186D ); glossae of labium curved over ventrally (Figs. 138D, 186G) ............................................................................................................. 3 Body rather flattened (Figs. 143A, 149A, 157A, 170A); head prognathous; caudal filaments with spines or setae not forming swimming setae; lingua of hypopharynx with lateral projections (Fig. 133E); glossae of labium variable (Fig. 133F) ............ 5 Gills plate-like, tapering gradually towards apex and ending in single filament (Fig. 186I) ........................................................................................................................... Terpides Gills plate-like, dorsal portion of gills II to VI ending in a filament and 1 or 2 lateral lobes (Fig. 138G) .................................................................................................... 4

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4 (3) – 5 (2)

– 6 (5) – 7 (6) – 8 (6) – 9 (8) – 10(9) – 11(10) – 12 (5) – 13 (12) – 14 (13) –

Ventral portion of gills II–VI without developed lobes (Fig. 138G) ...... Fittkaulus Ventral portion of gills II–VI with apical internal and external lobes (Fig. 192N) ...................................................................................................................................... Tikuna Labrum as wide as, or wider than, head (Figs. 170A, 151I, 174H); mouthparts of filtering type (third segment of maxillary palpi and second segment of labial palpi with rows of thick setae) (Figs. 151K, 174I); basal segment of maxillary palpi cupshaped, attached to side of maxilla (Figs. 146F, 195C) ............................................. 6 Labrum narrower than head (Figs. 149A, 157A); mouthparts not of filtering type (setae of maxillary and labial palpi not forming rows); basal segment of maxillary palpi not cup-shaped at base of maxilla (Figs. 173D, 191C) .................................. 12 Gills with 8 or more finger-like projections (Figs. 151M, 195F) ............................. 7 Gills without, or with 3 or fewer finger-like projections (Figs. 146K, 152F) ....... 8 Gills oval, fringed with filaments (Fig. 195F); all denticles of tarsal claws of equal size (Fig. 195E) ..................................................................................................... Traverella Gills rather quadrangular, truncated apically, with 8 to 10 finger-like projections (Fig. 151M); denticles of tarsal claws of equal size except for much larger apical denticle (Fig. 151L) ..................................................................................................Hylister Segment II of labial palpi elongated, approximately 6 times as long as segment I (Fig. 152D) .......................................................................................................... Leentvaaria Segment II of labial palpi no more than 3 times as long as segment I (Figs. 146H, 174I) ...................................................................................................................................... 9 Gills ending in lateral lobes and longer median process (Fig. 146K) ........ Hermanella Some or all gills with single median digitiform projection (Figs. 150N, 170G, 174L) ................................................................................................................................... 10 Apical tooth of maxilla very large (approximately 1/2 width of apex of maxilla) (Fig. 150K) ..................................................................................................... Hydrosmilodon Apical tooth of maxilla small (approximately 1/4 or less width of apex of maxilla) (Fig. 170D) ......................................................................................................................... 11 Gills present on abdominal segments I–VI, gills of segment VII vestigial; apical process of gills variable (Fig. 174L), absent on gill I, small or absent on some other gills; labrum recurved (Fig. 174H) ........................................................ Paramaka Gills present and similar in form on abdominal segments I–VII, dorsal and ventral portions with digitiform process (Fig. 170G); labrum not recurved (Fig. 170B) ...... .............................................................................................................................. Needhamella Each abdominal gill formed by single ovoid lamella, ending abruptly in a thick finger-like projection (Fig. 153G) ................................................................. Magallanella Gills on abdominal segments II–VI formed by two narrow to broad lamellae (one ventral and one dorsal portion) .................................................................................... 13 Gill margins with 6 or more filaments (Figs. 124H, 127G, 198I) ......................... 14 Gill margins with 0 to 3 filaments (Figs. 155J, 180H) .............................................. 17 Entire gill margins fringed with filamentous projections (Fig. 198I) .................... 15 Filamentous projections only present on the distal portion of the gill (Figs. 124H, 127G) .................................................................................................................................. 16

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15 (14) Row of long dorsal setae of labrum divided, without setae medially (Fig. 200A); anteromedian emargination of labrum with 5 flattened denticles (Fig. 200B) ...... .................................................................................................................................. Ulmeritus – Row of long dorsal setae of labrum entire, forming a single sinusoidal row (Figs. 198A, C); anteromedian emargination of labrum with 5 denticles, medial one sometimes very large (Figs. 198B, D) .......................................................... Ulmeritoides 16 (14) Gills I–VI with filamentous projections on apical margin (Fig. 124H), gill VII reduced to pair of filaments; margins of mandibles fringed with long setae, setae denser on apical 1/2 (Fig. 124D); anteromedian emargination of labrum broad, shallow (Figs. 124B-C) ................................................................................... Atopophlebia – Gills I–VII similar, with filamentous projections on apical margin (Fig. 127G); margins of mandibles with median tuft of setae, and some setae on basal 1/2 (Fig. 127D); anteromedian emargination of labrum with deep cleft (Figs. 127B-C) ..... ........................................................................................................................... Dactylophlebia 17 (13) Gills present on abdominal segments I–VI, gills broad, with (Figs. 155J-K) or without (Fig. 155L) an apical filament ........................................................... Massartella – Gills present on abdominal segments I–VII, gills variable but not as above ..... 18 18 (17) Gills on abdominal segment I reduced to a single filament, gills on segments II– VII with 3 projections, terminal filament of dorsal portion long and expanded (Fig. 126D) ...........................................................................................................Choroterpes – Gills on abdominal segment I similarly shaped as other gills ................................ 19 19 (18) Gills on abdominal segments VII or VI–VII vestigial, almost reduced to single filament (Fig. 143I) .......................................................................................................... 20 – Gills on abdominal segments VI and VII with same shape as gills on segments II–V .............................................................................................................................................. 21 20 (19) Gills on abdominal segments I–VI with 3 distal projections (Fig. 180H), median projection sometimes a little longer; smaller ventral portion, with similar projections; gill VII almost reduced to single filament ............................... Rhigotopus – Dorsal portion of gills on abdominal segments I–V wide, with narrow distal projection; small ventral portion, without terminal projection (Fig. 143H); gills VI and VII reduced to slender filaments (Fig. 143I) ..................................... Hapsiphlebia 21 (19) Gills on abdominal segments I–VII oval, dorsal portion 3/4 as long and 1/2 as wide as ventral portion (Fig. 163E) .............................................................. Microphlebia – Gills on abdominal segments I–VII various, dorsal portion as large as, or larger than, ventral portion (Figs. 167E, 191F–H) ............................................................... 22 22 (21) Gills with long and narrow terminal projection, 4/5 to 1/2 as long as basal part; basal part wider, usually with lateral lobes (Figs. 167E–K) ......................... Miroculis – Distal projection of gills, if present, equal or less than 1/3 as long as basal part (Figs. 177I, 191G); basal part not as above ................................................................ 23 23 (22) Posterolateral projections present on abdominal segments II or III–IX; lateral margins of clypeus divergent apically (Figs. 158A, 191A) ...................................... 24 – Posterolateral projections present on abdominal segments VI or VII or VIII– IX; lateral margins of clypeus parallel or convergent apically (Figs. 149B, 177A) .................................................................................................................................. 27

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24 (23) Lateral margins of labrum divergent, forming lateral angles (Fig. 191A) ..... Thraulodes 22 – Lateral margins of labrum rounded (Figs. 158A, 161A–B) .................................... 25 25 (24) External margin of mandibles glabrous, except for thin submedian tuft (Fig. 182C); basal denticles of tarsal claws of unequal size, remaining denticles gradually larger towards the apex (Fig. 182F) ................................................................................ Secochela – External margin of mandibles with setae on at least half its length (Figs. 158C, 161F); all denticles of tarsal claws gradually larger from the base to apex (Figs. 158E, 161H) ...................................................................................................................... 26 26 (25) Gills with main tracheal trunk and tracheal branches similarly pigmented (Fig. 158F) .......................................................................................................................... Massartellopsis – Gills with tracheal branches less pigmented than main tracheal trunk (Figs. 161M–O) ............................................................................................................................... Meridialaris 27 (23) Abdominal gills wide and oval (width approximately 1/3 total length of gill) with apical process narrowed apically (Figs. 177I–J); labium with glossa ventral to paraglossa (Fig. 177F) ....................................................................................... Penaphlebia – Abdominal gills various, lanceolate (width less than 1/4 total length of gill) (Fig. 149E) to narrow (Fig. 136D); labium with glossa on same plane as, or dorsal to, paraglossa (Fig. 184G) ..................................................................................................... 28 28 (27) Posterolateral projections present on abdominal segments VIII–IX ................... 29 – Posterolateral projections present on abdominal segments VI or VII–IX .......... 31 29 (28) Gills very narrow (maximum width approximately 1/10 the length), as in Fig. 136D ..................................................................................................................................... Farrodes – Gills wider (maximum width 1/5 to 1/3 the length), as in Figs. 149E and 184I .............................................................................................................................................. 30 30 (29) Widest section of gills approximately 1/5 of length (Fig. 184I) ...... Simothraulopsis – Widest section of gills approximately 1/3 of the length (Fig. 149E) ...................... ............................................................................................................................Homothraulus 31 (28) Antennae 2.5 times as long as head (Northern South America) ............ Ecuaphlebia – Antennae up to 1.5 times as long as head (Andes of central and southern Argentina and Chile) ....................................................................................................... 32 32 (31) Maxilla with tusk on apex of galea-lacinia (Fig. 139E) .............................. Gonserellus – Maxilla without tusk on apex of galea-lacinia (Figs. 130D, 173D) ....................... 33 33 (32) Labrum with lateral margins strongly rounded, anteromedian emargination wide and deep (Fig. 130A) with all denticles small and of equal size (Fig. 130B) .......... ............................................................................................................................ Demoulinellus – Labrum with the lateral margins weakly rounded, anteromedian emargination shallow (Fig. 173A) with central denticle larger than lateral denticles (Fig. 173B) ....................................................................................................................................... Nousia 34 (1) Gills wide, apical 1/2 fringed with finger-like processes (Fig. 122E) ........... Askola – Gills narrow, tapering gradually towards apex (Figs. 125F, 140K) ........................ 35 22

Nymphs of the group of genera containing Thraulodes, Meridialaris, Massartellopsis and Secochela are very similar and some variations occur within each genus. Adults are needed for confirmation of new records.

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35 (34) Tarsal claws with two rows of denticles (Figs. 125E, 178J), mandibles strongly curved (Figs. 125B, 178D) .............................................................................................. 36 – Tarsal claws with one row of denticles, apical denticle much larger than others (Fig. 140J); mandibles weakly curved (Fig. 140H) ............................................... Hagenulopsis 36 (35) Tarsal claws with 3 large denticles on median area (Fig. 178J); median area of mandible with round protuberance (Fig. 178D) ................................. Perissophlebiodes – Tarsal claws with 1 denticle near median area (Fig. 125E); median area of mandible without distinct protuberances (Fig. 125B) ........................................................ Bessierus CLAVES

PARA LOS

Adultos l – 2 (1)

–

3 (1) – 4 (3)

–

5 (4) – 6 (5)

GÉNEROS

SUDAMERICANOS

23

Alas posteriores ausentes .............................................................................................. 2 24 Alas posteriores presentes ................................................................................................ 3 Vena ICu1 de las alas anteriores unida a CuA en la base (Fig. 121A); placa estilígera del macho casi tan larga como ancha (Fig. 121C); lóbulos de los penes sin espinas ventrales; guía de huevos de la hembra (Fig. 121D) redondeado apicalmente, no alcanzando la mitad del esterno IX ...................................................................... Askola Vena ICu1 de las alas anteriores unida a la vena CuP en la base (Fig. 140A); placa estilígera del macho dos veces más ancha que larga; lóbulos de los penes con una espina ventral; guía de los huevos de la hembra agudos apicalmente, extendiéndose mas allá de la mitad del esternito IX ............................... Hagenulopsis Uñas tarsales de un par iguales entre sí, ambas agudas (Figs. 137H, 175H) ......... 4 Uñas tarsales de un par diferentes entre sí, una aguda y la otra roma (Figs. 164L, 187H) .................................................................................................................................. 13 Vena Sc de las alas posteriores termina distalmente cerca de la proyección costal (Figs. 137D, 185B); penes cortos, triangulares, lóbulos cercanos apicalmente y fusionados basalmente (Figs. 137I, 185E); rama basal de la vena MA combada, por lo que la horquilla es a menudo levemente asimétrica ....................................... 5 Vena Sc de las alas posteriores se extiende claramente mas allá de la débil proyección costal (Figs. 171B, 175D); penes largos o anchos (nunca triangulares), divididos parcial o totalmente, nunca triangulares (Figs. 171F-J, 175J-L); horquilla de MA simétrica (Figs. 171A, 175C) .............................................................................. 7 Proyección costal del ala posterior redondeada, apenas sobresaliendo del margen costal .......................................................................................................................... Terpides Proyección costal del ala posterior larga, en forma de dedo pulgar (Figs. 137C-D, 137F-G) ................................................................................................................................ 6 Porción dorsal de los ojos compuestos del macho sobre un pedúnculo largo (Fig. 137A); fórceps de los machos bi-segmentados (Fig. 137I) ........................ Fittkaulus

Aunque esta clave está orientada a la determinación de machos, puede ayudar en ciertos casos a la determinación de hembras también. 24 Los adultos de Bessierus y Perissophlebiodes no se conocen, pero las ninfas no poseen pterotecas posteriores, por lo que deberían separarse aquí. 23

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– 7 (4) – 8 (7) – 9 (8)

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10 (8) – 11 (10) – 12 (11) – 13 (3) – 14 (13)

25

Porción dorsal de los ojos compuestos del macho sobre un pedúnculo corto y ancho (Fig. 192A), fórceps de los machos tri-segmentados (Fig. 192F) ................. Tikuna 25 Venas transversas en el 1/3 apical del área costal del ala anterior anastomosadas (Fig. 142A) ........................................................................................................ Hapsiphlebia Venas transversas del área costal del ala anterior no anastomosadas (Figs. 154A, 156A) .................................................................................................................................... 8 Vena ICu1 unida directamente a la vena CuA (Fig. 179A) ........................................ 9 Base de la vena ICu1 no como arriba; encontrándose libre en la membrana (Fig. 171A), o uniéndose a las venas CuA y CuP por venas transversas (Fig. 156A), o aparentemente uniéndose directamente a CuP (Fig. 154A) .................................... 10 Vena MP2 del ala anterior fuertemente recurvada, base de MP2 mas cercana a la vena CuA que a la vena MP1, ocasionalmente uniéndose a la vena CuA (Fig. 175C); alas anteriores sin manchas en la horquilla de MA; cada lóbulo de los penes con una espina apical (Figs. 175J-O) ............................................................ Penaphlebia Vena MP2 del ala anterior moderadamente recurvada, base de la vena MP2 entre las venas CuA y MP1 (Fig. 179A); ala anterior del macho con pequeñas manchas en la horquilla de MA (a veces presentes también en las hembras); cada lóbulo de los penes con una espina lateral (Fig. 179C) ................................................. Rhigotopus En el ala anterior vena MP2 fuertemente recurvada; la vena ICu1 unida en la base a las venas CuP y CuA por una vena transversa; (Fig. 154A) .................. Massartella En el ala anterior la vena MP2 moderadamente recurvada; vena ICu1 unida en la base a las venas CuA y CuP por medio de venas transversas (Fig. 156A), o base de la vena ICu1 libre en la membrana (Fig. 171A) ......................................................... 11 Penes divididos en toda su extensión, tubulares, con una proyección apical en cada uno de ellos (Fig. 156D); Sc del ala posterior 9/10 de la longitud total del ala (Fig. 156B) ..................................................................................................... Massartellopsis Penes fusionados en 1/2 o 2/3 basales (Fig. 120C), Sc de las alas posteriores 3/4 a 4/5 de la longitud total del ala (Fig. 120B) ............................................................. 12 Penes fusionados en 1/2 basal, cada lóbulo con una espina subapical larga y gruesa dirigida hacia la línea media (Fig. 120C) ................................... Archethraulodes Penes fusionados en sus 2/3 basales, cada lóbulo con un corto proceso esclerotizado en el margen interno, nunca en forma de espina (Figs. 171F-J) ....................................................................................................................................... Nousia Horquilla de la vena MA del ala anterior claramente asimétrica (Figs. 144A, G, 164A, D) ............................................................................................................................ 14 Horquilla de la vena MA del ala anterior simétrica (Figs. 187B, D), o rama basal de MA combada, por lo que la horquilla parece levemente asimétrica (Figs. 131A, D) .............................................................................................................................................. 26 La proyección costal bien desarrollada en el ala posterior, su borde distal formando un ángulo casi recto con este; la vena Sc termina en una vena transversa o en la proyección costal (Figs. 144C, I, 147C, 183C) ............................................. 15

No han sido descriptas especies de este género de Sud América todavía, pero hemos visto especímenes no descriptos aún de Colombia.

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– 21 (20) – 22 (21) –

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La proyección costal en el ala posterior con forma de V invertida, y sobresaliendo del margen costal basalmente al punto donde la vena Sc se une a la vena C (Figs. 164B-C, E) ......................................................................................................................... 24 Segundo segmento de los fórceps genitales corto, longitud no más de 1,5 veces el ancho (Figs. 134D, 144M); ojos del macho frecuentemente agrandados pero no sobre pedúnculo (Fig. 183D) ......................................................................................... 16 Segundo segmento de los fórceps genitales largo, longitud por lo menos 3 veces el ancho (Fig. 147G); ojos del macho sobre pedúnculo; especies pequeñas (cuerpo aproximadamente 5 mm) ................................................................................................ 25 Angulos posterolaterales de la placa estilígera del macho proyectados posteriormente; cada lóbulo de los penes con un apéndice ventral, generalmente dirigido lateralmente (Figs. 134D, F, J) ...............................................................Farrodes Angulos posterolaterales de la placa estilígera del macho no proyectados como en el caso anterior; cada lóbulo de los penes con proyecciones ventrales no dirigidas lateralmente (Figs. 144J-M, 148D, 174E) .................................................................... 17 Placa estilígera del macho sin proyecciones sobre su margen posterior (Figs. 148D, 151D) .................................................................................................................................. 18 Placa estilígera del macho con una o dos proyecciones sobre su margen posterior (Figs. 174E, 193F) ............................................................................................................ 20 Penes divididos en su 1/4 apical, cada lóbulo con una proyección ventral en forma de espina curva, divergentes apicalmente (Fig. 151D) ........................ Hylister Penes divididos en su 1/2 o 2/3 apicales, proyecciones ventrales de cada lóbulo no como se menciona arriba (Figs. 148D, 183G) ..................................................... 19 Proyección costal de las alas posteriores fuertemente desarrollada (Figs. 183B-C); parte del área costal fuertemente teñida (Fig. 183C) ........................... Simothraulopsis Proyección costal de las alas posteriores no tan desarrollada; ala posterior sin fuerte pigmentación (Figs. 148B-C) ...........................................................Homothraulus Placa estilígera del macho con una proyección media única, que se proyecta dorsalmente entre los penes (Fig. 174F); cada lóbulo de los penes con una proyección en forma de espina, dirigida anteromedialmente (Fig. 174E) ......... ................................................................................................................................. Paramaka Placa estilígera del macho con proyecciones pares (Figs. 150E, 169E); proyecciones de los lóbulos de los penes no como arriba (Figs. 150E, 169E, 193F) .............................................................................................................................................. 21 Penes con proyecciones ventrales en forma de espinas rectas, dirigidas hacia el eje del cuerpo, con proyecciones romas en la placa estilígera del macho (Figs. 150E, 193F) ................................................................................................................................... 22 Penes con proyecciones en forma de espinas curvadas, o en forma de lóbulos; proyecciones de la placa estilígera del macho variadas (Fig. 169E) ...................... 23 Proyecciones de la placa estilígera del macho con base ancha (especies sudamericanas) (Fig. 193F) ................................................................. Traverella (Zonda) Proyecciones de la placa estilígera del macho con base angosta (especies sudamericanas) (Fig. 150E) ........................................................................ Hydrosmilodon

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23 (21) Proyecciones ventrales de los penes delgadas, en forma de espinas y divergentes apicalmente (Fig. 169E); proyecciones de la placa estilígera del macho en forma de espinas agudas cercanas a la base de los fórceps ...................................... Needhamella – Proyecciones de los penes y la placa estilígera variadas pero no en la combinación arriba mencionada (Figs. 144J-M) ..................................................................Hermanella 24 (14) Par de uñas tarsales de la pata anterior del macho de forma diferente a las de las patas medias y posteriores, una de las uñas anteriores roma, la otra aguzada en forma de espinas (Fig. 164L) .............................................................................. Miroculis – Par de uñas tarsales de cada pata anterior de igual forma que las de las patas medias y posteriores; en todas las patas una uña roma, la otra aguzada (Fig. 168D) ................................................................................................................................ Miroculitus 25 (15) Placa estilígera del macho con dos pequeñas proyecciones medianas (Fig. 147G); proyección costal del ala posterior aguda (Figs. 147B-C); forma de las alas posteriores como en las Figs. 147B-C ..................................................... Hermanellopsis – Placa estilígera de los machos redondeada (Figs. 162A-B); proyección costal de las alas posteriores redondeada (Figs. 162D-E); forma de las alas posteriores como en las Figs. 162D-E ............................................................................................... Microphlebia 26 (13) Ala posterior muy pequeña, menos de 1/10 de la longitud del ala anterior, con una proyección costal fuerte que se extiende mas allá del ápice del ala posterior; porción posterior del ala posterior reducida (Figs. 141B-C) ..................... Hagenulus – Ala posterior variable, 1/10 a 1/4 de la longitud del ala anterior, proyección costal variable pero nunca extendiéndose mas allá del ápice del ala (Figs. 123C, F, 128B, 187C, E); porción posterior del ala posterior no reducida ..................................... 27 27 (26) Lóbulos de los penes de los machos fusionados excepto por una fisura apical, a menudo con una espina apical pequeña (Figs. 159H, M) ......................... Meridialaris – Penes de los machos totalmente divididos o fusionados en su 1/2 o 1/3 basal (Figs. 123J, 196N, 199G) ................................................................................................ 28 28 (27) Segmento basal de los fórceps del macho largo y delgado, con el 1/6 basal formando un fuerte ángulo sobre el margen interno (Figs. 199C, G); ala anterior con una vena transversa formando un ángulo de aproximadamente 45° entre las venas MA1 y R4+5 distal a la horquilla de la vena MA (Figs. 199A, E) ................. 29 – Segmento basal de los fórceps del macho no como arriba (Figs. 128D, 187J); vena transversa entre las venas MA1 y R4+5 del ala anterior variable, si está presente, ángulo usualmente mayor de 45° (Figs. 123A, D, 187B, D) ................................... 30 29 (28) Vena Sc de las alas posteriores 9/10 de la longitud total del ala (Figs. 199B, F); membrana de las alas con manchas oscuras en las venas transversas (Figs. 199A, E); ápice de los penes redondeados, cada lóbulo con una proyección digitiforme ventral (Figs. 199C, G) ........................................................................................ Ulmeritus – Vena Sc de las alas posteriores 8/10 o menos de la longitud total del ala (Fig. 196B); alas sin patrón de coloración, excepto en la membrana costal (Fig. 196A); ápice de los penes más bien rectos, con una ó más espinas (Figs. 196F, I, O, R) .......... Ulmeritoides 30 (28) Penes sin proyecciones ventrales o ventrolaterales (Figs. 128D, 181C) ............... 31 – Penes con pronunciadas proyecciones ventrales o ventrolaterales (Figs. 123H-M, 131F, 187J) ......................................................................................................................... 33

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31 (30) Penes aguzados (Fig. 126C) alas anteriores con 3 ó 4 intercalares en el área cubital (Fig. 126A) (Colombia) .....................................................................................Choroterpes – Penes largos, redondeados o algo cuadrangulares apicalmente (Figs. 128D, 181C); alas anteriores con 2 largas intercalares en el área cubital (Figs. 128A, 181A) ...... 32 32 (31) Ápice de los penes sin espinas, redondeados, con un semicírculo de setas dorsales subapicales (Fig. 128D) ................................................................................. Demoulinellus – Ápice de los penes con cortas espinas, setas apicales e indentaciones sobre el margen subapical interno (Fig. 181C) ................................................................ Secochela 33 (30) Lóbulos de los penes con una proyección apical o subapical espiniforme dirigida hacia el centro (Figs. 187J, N, P); placa estilígera del macho aguzada apicalmente, con cavidades donde articulan los fórceps unidas, formando una única cavidad (Figs. 187J, N, 188D) .......................................................................................... Thraulodes – Lóbulos de los penes con una proyección subapical dirigida ventrolateralmente (Figs. 123H-M, 131F-G) placa estilígera del macho redondeada apicalmente, cavidades donde se alojan los fórceps separadas (Figs. 123H-M, 131F) ............. 34 34 (33) Lóbulos de los penes divididos hasta la base, lóbulos con proyecciones aguzadas (Figs. 131F-G); alas con manchas y coloración mas pronunciada en la bulla (Figs. 131A, D) ............................................................................................................. Ecuaphlebia – Lóbulos de los penes fusionados en su 1/3 basal, formando una única unidad redondeada, con una proyección ventral roma (Figs. 123H-M); alas con manchas en la porción basal del área costal o sin manchas (Figs. 123A, D) ...... Atopophlebia Ninfas 1 – 2 (1)

–

3 (2) – 4 (3) – 5 (2)

Pterotecas posteriores ausentes ..................................................................................... 34 Pterotecas posteriores presentes ..................................................................................... 2 Cuerpo hidrodinámico (Fig. 186A); cabeza hipognata; filamentos caudales con setas natatorias (2/5 a 3/5 basales de los filamentos con largas setas sobre los márgenes internos de los cercos y los márgenes laterales del filamento terminal); lingua de la hipofaringe sin proyecciones laterales (Figs. 138B, 186D); glosas del labio curvada ventralmente (Fig. 138D, 186G) ............................................................ 3 Cuerpo más bien aplanado (Figs. 143A, 149A, 157A, 170A); cabeza prognata; filamentos caudales con espinas o setas, pero no formando setas natatorias; lingua de la hipofaringe con proyecciones laterales (Fig. 133E); glosas del labio variable (Fig. 133F) ............................................................................................................................ 5 Branquias laminares, afinándose gradualmente hacia el ápice y terminando en un único filamento (Fig. 186I) ................................................................................... Terpides Branquias laminares, porciones dorsales de las branquias II a VI terminadas en un filamento y 1 o 2 lóbulos laterales (Fig. 138G) ........................................................... 4 Porción ventral de las branquias II-VI sin lóbulos desarrollados (Fig. 138G) ....... ................................................................................................................................. Fittkaulus Porción ventral de las branquias II-VI con un lóbulo apical interno y otro externo marcado (Fig. 192N) ................................................................................................ Tikuna Ancho del labro igual o mayor que el ancho de la cabeza (Figs. 151I, 170A, 174H); con aparato bucal filtrador (tercer segmento del palpo maxilar y segundo

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segmento del palpo labial con setas gruesas ordenadas en hileras) (Figs. 151K, 174I); segmento basal del palpo maxilar en forma de taza, articulado lateralmente a la maxila (Figs. 146F, 195C) .......................................................................................... 6 Ancho del labro menor que el ancho de la cabeza (Figs. 149A, 157A); sin aparato bucal filtrador (setas sobre los palpos maxilares y labiales no ordenados en hileras); segmento basal del palpo maxilar sin forma de taza en la base de la maxila (Figs. 173D, 191C) ........................................................................................................... 12 Branquias con 8 o más terminaciones filamentosas (Figs. 151M, 195F) ............... 7 Branquias con 3 o menos (puede no presentar) terminaciones filamentosas (Figs. 146K, 152F) ......................................................................................................................... 8 Branquias ovaladas, orladas de filamentos (Fig. 195F); dentículos de las uñas tarsales de tamaño semejante (Fig. 195E) ....................................................... Traverella Branquias más bien cuadrangulares, truncadas apicalmente con 8 a 10 proyecciones apicales filamentosas (Fig. 151M); dentículos de las uñas tarsales de tamaño semejante, excepto el apical que es mucho mayor (Fig. 151L) ........ Hylister Segmento II del palpo labial elongado, aproximadamente 6 veces más largo que el segmento I (Fig. 152D) .................................................................................... Leentvaaria Segmento II del palpo labial no más de 3 veces la longitud del segmento I (Figs. 146H, 174I) ......................................................................................................................... 9 Branquias terminadas en lóbulos laterales y proceso mediano más largo (Fig. 146K) ....................................................................................................................Hermanella Un único proceso digitiforme presente en alguna o todas las branquias (Figs. 150N, 170F, 174L) ........................................................................................................... 10 Colmillo maxilar grande (aproximadamente 1/2 del ancho del ápice de la maxila) (Fig. 150K) ..................................................................................................... Hydrosmilodon Colmillo maxilar pequeño (aproximadamente 0,25 o menos del ancho del ápice de la maxila) (Fig. 170D) ...................................................................................................... 11 Branquias presentes en los segmentos abdominales I–VI, branquia en el segmento VII vestigial; proceso apical variable (Fig. 174L), pequeño o ausente en algunas otras branquias; labro recurvado (Fig. 174H) ............................................... Paramaka Branquias presentes y similares en forma en los segmentos abdominales I–VII, porciones dorsal y ventral con proceso digitiforme (Fig. 170G); labro no recurvado (Fig. 170B) ...................................................................................... Needhamella Cada branquia abdominal compuesta de una única lámina ovoide, terminando abruptamente en un proceso filamentoso grueso (Fig. 153G) .............. Magallanella Branquias en los segmentos abdominales II–VI compuestas de dos láminas angosta a anchas (una porción ventral y otra dorsal) ............................................... 13 Márgenes de las branquias con 6 o más filamentos (Figs. 117G, 124H, 198I) ... 14 Márgenes de las branquias con 0 a 3 filamentos (Figs. 155J, 180H) ..................... 17 Todo el margen de las branquias rodeados de procesos filamentosos (Fig. 198I) .............................................................................................................................................. 15 Procesos filamentosos solo presentes en la porción distal de la branquia (Figs. 124H, 127G) ..................................................................................................................... 16

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15 (14) Hilera de setas dorsales largas del labro dividida, sin setas medialmente (Fig. 200A); 5 dentículos aplanados en la emarginación anteromediana del labro (Fig. 200B) ..... .................................................................................................................................. Ulmeritus – Hilera de setas dorsales largas del labro entera, formando una hilera sinusoidal continua (Figs. 198A, C); 5 dentículos en la emarginación anteromediana del labro, el medio algunas veces bien desarrollado (Figs. 198B, D) ........... Ulmeritoides 16 (14) Branquias I–VI con procesos filamentosos en el margen apical (Fig. 124H) branquia VII reducida a un par de filamentos; márgenes de las mandíbulas orlado de largas setas, más densas en su 1/2 apical (Fig. 124D); emarginación anteromediana del labro ancha, poco profunda (Figs. 124B-C) ........... Atopophlebia – Branquias I–VII iguales, con procesos filamentosos en su margen apical (Fig. 127G); márgenes de las mandíbulas con un penacho mediano de setas, y algunas setas en su mitad basal (Fig. 127D); emarginación anteromediana del labro con una profunda hendidura (Figs. 127B-C) .................................................. Dactylophlebia 17 (13) Branquias presentes en los segmentos abdominales I–VI, branquias anchas con un filamento terminal (Figs. 155J–K) o sin éste (Fig. 155L) .......................... Massartella – Branquias presentes en los segmentos I–VII, branquias variables, pero no como arriba ................................................................................................................................... 18 18 (17) Branquias en el segmento abdominal I reducido a un único filamento, branquias en los segmentos II–VII con 3 proyecciones, filamento mediano de la porción dorsal largo y expandido (Fig. 126D) ............................................................Choroterpes – Branquias en el segmento abdominal I de la misma forma que las otras branquias .............................................................................................................................................. 19 19 (18) Branquias en los segmentos abdominales VII o VI–VII vestigiales, reducida casi a un filamento (Fig. 143I) .................................................................................................. 20 – Branquias en los segmentos abdominales VI y VII de la misma forma que las II a V .............................................................................................................................................. 21 20 (19) Branquias presentes en los segmentos abdominales I–VI, branquia con 3 filamentos distales (Fig. 180H), proyección mediana a veces un poco más larga, porcion ventral más pequeña, con proyecciones similares, branquia VII reducida casi a un único filamento ................................................................................... Rhigotopus – Porción dorsal de las branquias de los segmentos abdominales I–V anchas con un filamento distal delgado; porción ventral pequeña, sin filamento terminal (Fig. 143H); branquias VI y VII reducidas a delgados filamentos (Fig. 143I) ................. ............................................................................................................................. Hapsiphlebia 21 (19) Branquias en los segmentos abdominales I–VII ovales, con la porción dorsal 3/4 de la longitud y 1/2 del ancho de la parte ventral (Fig. 163E) .............. Microphlebia – Branquias en los segmentos abdominales I–VII variables, porción dorsal de la branquia de igual o mayor tamaño que la porción ventral (Figs. 167E, 191F–H) ....................... 22 22 (21) Branquias con un largo y delgado filamento terminal, 4/5 a 1/2 de la longitud de la parte basal; parte basal más ancha, usualmente con lóbulos laterales (Figs. 167E–K) .................................................................................................................. Miroculis – Filamento distal de las branquias, si está presente, igual o menos de 1/3 de la porción basal de la branquia (Figs. 177I, 191G); porción basal no como arriba ..................... 23

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23 (22) Proyecciones posterolaterales presentes en los segmentos abdominales II o III– IX; márgenes del clipeo divergentes apicalmente (Figs. 158A, 191A) .................. 24 – Proyecciones posterolaterales presentes en los segmentos abdominales VI o VII o VIII–IX; márgenes laterales del clipeo paralelos o convergentes apicalmente (Figs. 149B, 177A) ....................................................................................................................... 27 24 (23) Márgenes laterales del labro divergentes, formando un ángulo con el margen posterior (Fig. 191A) ...................................................................................... Thraulodes 26 – Bordes laterales del labro redondeados (Figs. 158A, 161A–B) .............................. 25 25 (24) Margen externo de las mandíbulas glabro, excepto por un delgado penacho submediano (Fig. 182C); dentículos basales de las uñas tarsales de tamaño desigual, resto progresivamente mayores hacia el ápice (Fig. 182F) ........... Secochela – Margen externo de las mandíbulas con setas en por lo menos la mitad de su extensión (Figs. 158C, 161F); dentículos de las uñas tarsales creciendo progresivamente desde la base hacia el ápice (Figs. 158E, 161H) ......................... 26 26 (25) Branquias con la tráquea principal y sus ramificaciones pigmentadas (Fig. 158F) .......................................................................................................................... Massartellopsis – Branquias con la tráquea principal pigmentada, sus ramificaciones mucho menos pigmentadas (Figs. 161M–O) ......................................................................... Meridialaris 27 (23) Branquias abdominales anchas y ovales (ancho aproximadamente 1/3 de la longitud total de la branquia) con filamento apical afinado apicalmente (Figs. 177I–J); labio con glossa ventral a la paraglossa (Fig. 177F) ................... Penaphlebia – Branquias abdominales variables, lanceoladas (ancho menor a 1/4 la longitud total de la branquia) (Fig. 149E) a angostas (Fig. 136D); labio con glosa en el mismo plano, o dorsal a la paraglosa (Fig. 184G) .................................................................. 28 28 (27) Proyecciones posterolaterales presentes en los segmentos abdominales VIII – IX .............................................................................................................................................. 29 – Proyecciones posterolaterales presentes en los segmentos abdominales VI o VII – IX .............................................................................................................................................. 31 29 (28) Branquias muy angostas (ancho máximo aproximadamente 1/10 del largo), como en la Fig. 136D ........................................................................................................Farrodes – Branquias más anchas (ancho máximo entre 1/5 y 1/3 del largo) como en las Figs. 149E, 184I ......................................................................................................................... 30 30 (29) Ancho máximo de las branquias aproximadamente 1/5 de la longitud (Fig. 184I) ......................................................................................................................... Simothraulopsis – Ancho máximo de las branquias aproximadamente 1/3 de la longitud (Fig. 149E) ............................................................................................................................Homothraulus 31 (28) Antenas 2,5 veces la longitud de la cabeza (Norte de Sudamérica) ................. ...............................................................................................................................Ecuaphlebia – Antenas hasta 1,5 veces la longitud de la cabeza (Andes del centro y sur de Argentina y Chile) ............................................................................................................ 32 26

Las ninfas del grupo de géneros Thraulodes, Meridialaris, Massartellopsis y Secochella son muy similares y presentan algunas variaciones dentro de cada género. Los adultos son necesarios para la confirmación de nuevos registros.

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32 (31) Maxila con un colmillo en el ápice de la galea-lacinia (Fig. 139E) .......... Gonserellus – Maxila sin colmillo en el ápice de la galea-lacinia (Figs. 130D, 173D) ................. 33 33 (32) Labro con los márgenes laterales muy redondeados, con una emarginación anteriomediana ancha y profunda (Fig. 130A), con dentículos pequeños y de igual tamaño (Fig. 130B) ........................................................................................ Demoulinellus – Labro con los márgenes laterales suavemente redondeados, emarginación anteromediana poco profunda (Fig. 173A), con un dentículo central más grande que los laterales (Fig. 173B) ....................................................................................Nousia 34 (1) Branquias anchas, mitad apical con procesos digitiformes (Fig. 122E) ....... Askola – Branquias delgadas, afinándose progresivamente hacia el ápice (Figs. 125F, 140K) .............................................................................................................................................. 35 35 (34) Dos hileras de dentículos en las uñas tarsales (Figs. 125E, 178J), mandíbulas fuertemente curvadas (Figs. 125B, 178D) ................................................................... 36 – Una hilera de dentículos en las uñas tarsales, dentículo apical mucho mas grande que el resto (Fig. 140J); mandíbulas suavemente curvadas (Fig. 140H) ............................................................................................................................. Hagenulopsis 36 (35) Uña tarsal con 3 dentículos grandes ubicados en la zona media (Fig. 178J); área media de la mandíbula con una protuberancia redondeada (Fig. 178D) ................. ........................................................................................................................ Perissophlebiodes – Uña tarsal con 1 dentículo cerca de la zona media (Fig. 125E); área media de la mandíbula sin protuberancias marcadas (Fig. 125B) ...................................... Bessierus Genus Archethraulodes PESCADOR & PETERS Genus D PESCADOR & PETERS, 1980b: 47. Archethraulodes PESCADOR & PETERS, 1982: 1; SAVAGE, 1987b: 207; DOMÍNGUEZ et al., 2001: 41. (Type-species: Archethraulodes spatulus PESCADOR & PETERS, original designation).

Taxonomy: PESCADOR & PETERS (1982) established the genus Archethraulodes based on male imagos. Its phylogenetic relationships were analyzed previously by PESCADOR & PETERS (1980b), “Genus D”. The nymph and female imago are unknown. Adult Characteristics: 1) Fork of vein MA of fore wings symmetrical (Fig. 120A); 2) hind wings lacking a costal projection (Fig. 120B); 3) length of vein Sc of hind wings at least three quarters maximum length of hind wings (Fig. 120B); 4) tarsal claws of a pair similar, each apically hooked with an opposing hook (Fig. 120D); and 5) penis lobes fused in basal half, each lobe with a long thick subapical spine (Fig. 120C). Nymphal Characteristics: Unknown. Distribution: Southwestern Argentina (Neuquen Prov.) and Chile (Aisen and Curico Prov.). Biology: The only species known from this genus appears to have a wide altitudinal range, with adults collected from sea level to 2500 m. The adults were collected in February.

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MA

A

Sc

B

E

C

D

Plate 120. Archethraulodes, imago. Figs. 120A-E, A. spatulus adult. 120A, fore wing; 120B, hind wing; 120C, male genitalia, v.v.; 120D, male tarsal claws; 120E, abdominal terga IV-VII. (MA = fork of MA). Figs. 120A-E modified from PESCADOR & PETERS, 1982.

Archethraulodes spatulus PESCADOR & PETERS Archethraulodes spatulus PESCADOR & PETERS, 1982: 2 (male); HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 210; DOMÍNGUEZ et al., 2001: 41.

Taxonomy: Archethraulodes spatulus is the only species in the genus and is presently known only from the male imago. The pattern of dark brown maculae on the abdominal terga is shown in Fig. 120E. Distribution and Biology: Distribution and biology are discussed in the generic description. Genus Askola PETERS Askola PETERS, 1969: 253. (Type-species: Askola froehlichi PETERS, original designation).

Taxonomy: The genus Askola was established by PETERS (1969) based on adults of both sexes and nymphs from Eastern Brazil. Adult Characteristics: 1) Stem of vein MA of fore wings sagged so fork sometimes slightly asymmetrical (Fig. 121A); 2) vein MP of fore wing symmetrical; 3) vein ICu1 of

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fore wings joined at base to vein CuA (Fig. 121A); 4) hind wings absent; 5) claws of a pair dissimilar, one apically hooked and the other obtuse, pad-like (Fig. 121B); 6) penes divided, tubular, slender, apically pointed, and without appendages or spines (Fig. 121C); and 7) female with egg guide which is prolonged to middle of abdominal sternum IX (Fig. 121D); 8) female sternum IX deeply cleft apically (Fig. 121E). Nymphal Characteristics (general aspect as in Fig. 122A): 1) Anteromedian emargination of labrum with two denticles of approximately same size (Fig. 122B); 2) mandibles with setae as in Fig. 122C; 3) denticles of tarsal claws progressively larger apically (Fig. 122D); 3) dorsal and ventral portions of abdominal gills I-VII equal and plate-like, with the apical half of each plate bordered with finger-like processes (Fig. 122E); 4) posterolateral spines present on abdominal segments V-IX or VI-IX; 5) hind wing pads absent. Distribution: Brazil (São Paulo, Paraná and Santa Catarina). Undescribed species recorded from Amazon region. Biology: Nymphs of this species were collected in calm water along margins of small streams or rivers of the coastal mountains of Southern Brazil. The adults have been collected from August to January. MP

MA

A

B ICu1 CuA

C

D

E

F

Plate 121. Askola, imago. Figs. 121A-E, A. froehlichi: 121A, fore wing; 121B, fore claw; 121C, male genitalia, v.v. (roman numbers = number of segments). 121D, female abdomen showing egg guide; 121E, apex of female sternum IX; 121F, male imago abdominal terga V-VII. (MA = fork of MA). Figs. 121A-F, modified from PETERS, 1969.

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Askola froehlichi PETERS Askola froehlichi PETERS, 1969: 255 (male, female, nymph); HUBBARD, 1982a: 263; DA-SILVA, 2002a: 1.

Taxonomy: Askola froehlichi is the only species described in the genus, and can be characterized by the structure of the male genitalia and the lack of coloration in the wings. The abdominal pattern in the holotype is as in Fig. 121F. DA-SILVA (2002a) described some specific variations in the nymph. Distribution and Biology: Distribution and biology are discussed in the generic description. Genus Atopophlebia FLOWERS “Ulmeritus ally” EDMUNDS et al., 1976: 239. Atopophlebia FLOWERS, 1980: 162; FLOWERS, 1987b: 203; DOMÍNGUEZ & MOLINERI, 1996: 55. (Type-species: Atopophlebia fortunensis FLOWERS, original designation).

Taxonomy: Atopophlebia was established by FLOWERS in 1980, based on two male imagos from Panama. In 1987, the same author described the female imago and the nymph of this species, and two new species from Northern South America. DOMÍNGUEZ & MOLINERI (1996) described a third species from Northern Argentina and Southern Bolivia; in the same paper they stated that no significant morphological differences could be found between the nymphs of these two species. The recent collection of A. fortunensis in Colombia has increased the number of recognized species in South America to four. There is a wide variation in the shape of the hind wing among the species, especially marked between the Northern and Southern species. Adult Characteristics: 1) Fork of vein MA symmetrical (Fig. 123A) to slightly asymmetrical (Fig. 123D), 2) vein MP2 attached at base to veins MP1 and CuA with a cross vein (Figs. 123A, D); 3) hind wings with costal projection located from about 1/2 (Figs. 123E-F) to 5/6 distance from wing base to apex (Figs. 123B-C); 4) vein Sc of hind wings from 3/5 to 4/ 5 length of wing (Figs. 123B-C, E-F); 5) tarsal claws of a pair dissimilar, one apically hooked, other obtuse, pad-like (Fig. 123G); 6) penis lobes narrow and divided on apical 2/3, each lobe twisted ventrally and laterally, each ending in a terminal blunt projection (Figs. 123H-M); and 7) female sternum IX shallowly emarginate apically (Fig. 123O). Nymphal Characteristics (Fig. 124A): 1) Labrum wide, somewhat angled laterally (Fig. 124B), with distinct broad-based denticles on anteromedian emargination (Fig. 124C); 2) outer margin of mandibles curved, fringed with long setae, denser at apical 1/2 (Fig. 124D); 3) maxilla as in Fig. 124E; 4) segment I and II of labial palpi subequal, segment III about 1/3 length of segment II; 5) submentum of labium with few setae at base (Fig. 124F); 6) tarsal claws with denticles increasing in size toward apex (Fig. 124G); 7) abdominal gills I-VI fringed on apical margin (Fig. 124H), gill VII reduced; 8) posterolateral projections on abdominal segments II-IX. Distribution: Colombia, Ecuador, Peru, Bolivia and Argentina. Biology: The nymphs of the known species of this genus were collected from different sized streams, with strong to slow current, but generally from shaded habitats. In many streams, the bottom was composed of rocks and boulders, but in Colombia the nymphs were collected from sandy-bottomed streams. Nevertheless, the nymphs are very difficult to find,

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B

C

D

D

A

E

Plate 122. Askola, nymph. Figs. 122A-C, A. froehlichi: 122A, nymphal habitus; 122B, clypeus and labrum, d. v.; 122C, left mandible, d.v.; 122D, fore claw; 122E, gill IV. (D = denticle). Fig. 122A, drawing by C. Molineri. Figs. 122B-E modified from PETERS, 1969.

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and are very scarce in collections. Subimagos in the laboratory emerged from 20:00 to 21:00, and molted to imagos after 24 hours. The largest number of adults were observed at the end of the rainy period, on clear sunny days. The adults were collected from April to June. KEYS

TO

SOUTH AMERICAN ATOPOPHLEBIA

27

Adults 1 – 2 (1) – 3 (2) –

CLAVES

Costal projection of hind wings located at approximately 1/2 distance from wing base to apex (Figs. 123E-F); a row of small spines on each penis lobe (Figs. 123J-K) ................................................................................................................ Atopophlebia flowersi Costal projection of hind wings located at approximately 5/6 distance from base to margin (Figs. 123B-C); penis lobes without spines (Figs. 123H-I, L-M) ......... 2 Tibia of fore leg dark brown; abdominal tergum I yellowish; penis lobes as in Fig. 123L ............................................................................................... Atopophlebia obrienorum Tibia of fore leg yellowish, with apex tinged with dark brown; abdominal tergum I yellowish, with posterior margin brown; penis lobes not as above (Figs. 123I, M) ........ 3 Abdominal tergum I without a pair of anterolateral spots; penis lobes as in Fig. 123I .................................................................................................. Atopophlebia fortunensis Abdominal tergum I with a pair of anterolateral black spots; penis lobes as in Fig. 123M ............................................................................................... Atopophlebia yarinacocha PARA LOS

ATOPOPHLEBIA

SUDAMERICANOS 28

Adultos 1 – 2 (1) – 3 (1) –

Proyección costal de las alas posteriores localizada a aproximadamente 1/2 de la distancia de la base del ala al ápice (Figs. 123E-F); una hilera de pequeñas espinas sobre cada lóbulo de los penes (Figs. 123J-K) ........................... Atopophlebia flowersi Proyección costal de las alas posteriores localizada a aproximadamente 5/6 de la distancia de la base del ala al ápice (Figs. 123B-C); sin hileras de espinas en cada lóbulo de los penes (Figs. 123H-I, L-M) ....................................................................... 2 Tibia de la pata anterior castaño oscura; tergo abdominal I amarillento; lóbulos de los penes como en la Fig. 123L ................................................ Atopophlebia obrienorum Tibia de la pata anterior amarillenta, con el ápice teñido de castaño oscuro; tergo abdominal I amarillento, con el borde posterior castaño; lóbulos de los penes no como en el caso anterior (Figs. 123I, M) ..................................................................... 3 Tergo abdominal I sin un par de manchas anterolaterales negruzcas; lóbulos de los penes como en la Fig. 123I ........................................................ Atopophlebia fortunensis Tergo abdominal I con un par de manchas anterolaterales negruzcas; lóbulos de los penes como en la Fig. 123M ............................................. Atopophlebia yarinacocha

The only nymphs known are those of A. flowersi and A. fortunensis, that are only separable by the abdominal coloration. For this reason a key for nymphs is not provided. 28 Las únicas ninfas conocidas son las de A. flowersi and A. fortunensis, y se separan solo por la coloración abdominal, por lo que no se incluye clave para nymphs. 27

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Atopophlebia flowersi DOMÍNGUEZ & MOLINERI Atopophlebia flowersi DOMÍNGUEZ & MOLINERI, 1996: 55 (male, female, nymph).

Taxonomy: This species was described based on imagos and subimagos of both sexes and the nymph. A. flowersi can be characterized in the adults by the following combination of characters: 1) Fore tibiae blackish brown, except basal 1/7 and apex yellowish; 2) terga MA

MA

A

D CuA MP2 MP1

CP

B

CP Sc

Sc

C

E

F

I K

L

G

M H

J

O N Plate 123. Atopophlebia, adults. Figs. 123A-F, Wings: 123A-C, A. obrienorum, 123A, fore wing; 123B, hind wing; 123C, same, enlarged; 123D-G, A. flowersi, 123D, fore wing; 123E, hind wing; 123F, same, enlarged. 123G, male tarsal claw. 123H-M, male genitalia: 123H-I, A. fortunensis: 123H, genitalia, v.v.; 123I, penes, l. v.; 123J-K, A. flowersi, 123J, genitalia, v.v.; 123K, apex of penis lobe, detail; 123L, A. obrienorum, penes, l.v.; 123M, A. yarinacocha, penes, l.v. 123N, A. flowersi, abominal terga II-IV. 123O, A. fortunensis, female sternum IX. . (CP = Costal Projection). Figs. 123A-C, I, L-M and O modified from FLOWERS, 1987b; 123G-H from FLOWERS, 1980 and 123D-F, J-K and N from DOMÍNGUEZ & MOLINERI, 1996.

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yellowish brown, with posterior margins of segments I-IX black and divergent lines from posterior and anterior margins (Fig. 123N); 3) veins C, Sc and R1 of fore wing brownish, remaining longitudinal and cross veins blackish; cross veins in costal and subcostal areas clouded with black (Fig. 123D); 4) costal projection of hind wings located about 1/2 distance from base to apex as in Figs. 123D-F; 5) ventrolateral projections of penes lobes with spines as in Figs. 123J-K. The nymph can be separated from A. fortunensis, the only other species known in the nymph, by the abdominal color pattern (Fig. 123N). Distribution and Biology: Bolivia (Cochabamba) and Argentina (Salta, Orán). The specimens were found in mountain rain forests streams between 700 and 2400 m in rockbottomed streams with strong current. The adults were collected in October and November, attracted to light as subimagos. Atopophlebia fortunensis FLOWERS “Ulmeritus ally” EDMUNDS et al., 1976: 239, Fig. 141. Atopophlebia fortunensis FLOWERS, 1980: 162 (male); FLOWERS, 1987b: 205 (female, nymph); DOMÍNGUEZ & MOLINERI, 1996: 55; ZUÑIGA et al., 1997: 264; MOSQUERA et al., 2001: 159.

Taxonomy: This species was described originally based on male imagos from Panamá. Later, FLOWERS (1987) described the female and nymph, based also in Central American specimens, stating that the wing venation he described originally from the type material was atypical. A. fortunensis can be characterized in the adults by the following combination of characters: 1) Fore tibiae dark brown at apex, remainder yellow; 2) abdominal terga IX-X brown, with a pair of anterolateral black maculae on terga VIII-IX; 3) longitudinal veins of fore and hind wings yellow, cross veins light brown, membrane of fore and hind wings hyaline, except extreme apex of fore wing, and apical 1/3 of hind wing shaded with amber; 4) penis lobes narrow, widely separated in apical 2/3, with ventrolateral projections as in Figs. 123H-I. The nymph can be separated from A. flowersi, the only other species known in the nymphs, by the abdominal color pattern (Fig. 124A). Distribution and Biology: Panamá, Costa Rica and Colombia. The nymphs were collected in a heavily shaded, forested, steep, ravine stream, with small pools and riffles. The adults were collected in May-June. Atopophlebia obrienorum FLOWERS Atopophlebia obrienorum FLOWERS, 1987b: 207.

Taxonomy: This species is known from one male subimago. FLOWERS (1987b) justified the description based on a subimago, because it was ready to molt and important characters were already visible and fully developed (wing venation and genitalia). A. obrienorum can be characterized by the following combination of characters: 1) Fore tibiae and two apical segments of abdomen entirely dark brown; 2) veins of fore and hind wings (Figs. 123A-C) yellow; membrane of fore and hind wings hyaline, membrane of hind wings washed with smoky brown at base; 3) penis lobes narrow, widely separated in apical 2/3, each lobe with a ventrolateral projection near apex (Fig. 123L). Distribution and Biology: Ecuador (Pastaza). The adult was collected in April.

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B

C D

D

D

G

A

F

H

E

Plate 124. Atopophlebia fortunensis, nymph. Figs. 124A-H: 124A, habitus; 124B, labrum d.v.; 124C, detail of anteromedian emargination showing denticles; 124D, left mandible; 124E, maxilla; 124F, labium (left dorsal, right ventral); 124G, tarsal claw; 124H, gill III. (D = Denticle). Figs. 124A-H modified from FLOWERS, 1987b.

Atopophlebia yarinacocha FLOWERS Atopophlebia yarinacocha FLOWERS, 1987b: 208.

Taxonomy: This species is known from male imago and subimagos. A. yarinacocha can be characterized by the following combination of characters: 1) Abdominal tergum I brown, with a pair of anterolateral black maculae; 2) wing veins hyaline, membrane amber; subcostal area of hind wing amber stained in basal 1/2; 3) penes fused in basal 1/3, with a curved ventrolateral projection near apex of each penis lobe (Fig. 123M). Distribution and Biology: Peru (Loreto). The adults were collected in April.

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Genus Bessierus THOMAS & ORTH Bessierus THOMAS & ORTH, in THOMAS et al., 2000: 49 (nymph). (Type-species: Bessierus doloris THOMAS & ORTH, original designation).

Taxonomy: Bessierus was established by THOMAS & ORTH (2000) based on nymphs from French Guiana. According to these authors, this genus was related to Hagenulopsis by the absence of hing wing pads and spines on the posterior margin of abdominal terga and to Perissophlebiodes by the deeply emarginated labrum, shape of the labial palpi and structure of the tarsal claws. Adult Characteristics: Unknown. Nymphal Characteristics: 1) Labrum deeply emarginated (Fig. 125A), emargination without denticles; 2) mandibles strongly curved, with a few setae as in Fig. 125B; 3) maxillae without protuberance on anterolateral margin of galea-lacinia (Fig. 125C, compare with Fig. 178E); 4) labial palpi rather long and narrow, not inflated (Fig. 125D); 5) tarsal claws with one median row of denticles, with one large apical denticle, and one subapical row of small denticles (Fig. 125E); 6) hind wing pads absent; 7) gills long and narrow, tapering apically (Fig. 125F); 8) posterolateral projections present on abdominal segments VIII-IX.

A

B C

D

D E

F

Plate 125. Bessierus doloris, nymph. Figs. 125A-F: 125A, labrum, d.v.; 125B, left mandible; 125C, maxilla; 125D, labium, v.v.; 125E, tarsal claw; 125F, gill IV. (D = denticle). Figs. 125A-F modified from THOMAS et al., 2000.

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Distribution: French Guiana (Oyapok and Maroni Basins). Biology: The nymphs were collected in two rivers in areas of low water current on the bottom gravel and in aquatic macrophytes. The rivers where nymphs were found are subjected to high anthropogenic disturbances. Bessierus doloris THOMAS & ORTH Bessierus doloris THOMAS & ORTH, in THOMAS et al., 2000: 50.

Taxonomy: This is the only species known in the genus. It can be characterized by: 1) Light yellow coloration, with a strong violaceous band across the abdominal segment I; and 2) its small size (body length= 3.2-3.3 mm). Distribution and Biology: Distribution and biology are discussed in the generic description. Genus Choroterpes EATON Choroterpes EATON, 1881: 194. (Type-species: Choroterpes picteti EATON, original designation).

Taxonomy: No species of this genus has been reported from Continental South America, and all species previously assigned to Choroterpes have been moved to other genera. Only recently an unnamed species has been reported from the region (PETERS, et al., 2005). For this reason, we included it in the generic key, but will not discuss it any further. Ilustrations for the generic key are provided in Plate 126. III II

A I

B C

D Plate 126. Choroterpes. Figs. 126A-C, C. albiannulata, imago. 126A, fore wing (arrows = cubital intercalavies); 126B, hind wing; 126C, genitalia, v.v. (Roman numbers = segment numbers). 126D, Choroterpes sp. nymph, from Colombia, gill IV. Figs. 126A-C, modified from EDMUNDS, JENSEN & BERNER, 1976.

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Distribution: Ethiophian, Oriental, Holarctic and Neotropical. In South America an unpublished record from Colombia, Antioquia, by Humberto Moreno. Genus Dactylophlebia PESCADOR & PETERS Genus A of PESCADOR & PETERS, 1980b: 47. New genus A of PESCADOR; LANDA, SOLDÁN & PETERS, 1980: 192. Dactylophlebia PESCADOR & PETERS, 1980c: 332; DOMÍNGUEZ et al., 1994: 41. (Type-species: Dactylophlebia carnulenta PESCADOR & PETERS, original designation).

Taxonomy: PESCADOR & PETERS (1980c) established the genus Dactylophlebia for the species D. carnulenta. The genus is presently known only from the nymph. Adult Characteristics: Unknown. Nymphal Characteristics (general aspect as in Fig. 127A): 1) Clypeus narrower than labrum with lateral margins divergent (Fig. 127B); 2) lateral margins of labrum abruptly curved (Fig. 127B); 3) anteromedian emargination of labrum Vshaped, with weakly developed blunt denticles (Fig. 127C); 4) outer margin of mandibles slightly angular with median setae tuft and setae along the basal half (Fig. 127D); 5) galea-lacinia of maxillae apically narrow, segments I and II of palpi equal in length, segment III a little shorter (Fig. 127E); 6) tarsal claws with double row of denticles, progressively larger apically (Fig. 127F); 7) posterolateral projections present on abdominal segments VI-IX; 8) gills on abdominal segments I-VII similar, plate-like, with dorsal and ventral portions terminating in finger-like processes (Fig. 127G). Distribution: Restricted to Southern Chile, Osorno Prov. (40º34’S; 73º09’W, X region) and Southwestern Argentina (Neuquen Prov). Biology: The nymphs were collected at altitudes above 1000 m under rocks, sometimes covered with algae. They are primarily detritivores, with gut contents consisting of detritus, mineral particles, diatoms and filamentous algae. Mature nymphs were collected in December, so the adults probably fly around this time of the year, although none were collected or reared. The streams had rocky bottoms and swift current. Nymphs are very delicate and lose the gills easily. Dactylophlebia carnulenta PESCADOR & PETERS Dactylophlebia carnulenta PESCADOR & PETERS, 1980c: 333 (nymph); HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 211; DOMÍNGUEZ et al., 1994: 42.

Taxonomy: Dactylophlebia carnulenta is the only species in the genus. Its small size (up to 7 mm), dark coloration, fleshy finger-like gills (Fig. 127G) and unique double rows of tarsal denticles make the nymph (Fig. 127F) easily distinguishable from any other species of the South American Leptophlebiidae. Distribution and Biology: Distribution and biology are discussed in the generic description.

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C

B

D

A G D

E

F

Plate 127. Dactylophlebia carnulenta, nymph. Figs. 127A-G: 127A, habitus; 127B, clypeus and labrum, d.v.; 127C, anteromedian emargination of labrum, detail; 127D, left mandible; 127E, maxilla; 127F, fore claw; 127G, gill IV. (D = denticle). Figs. 127A-G, modified from PESCADOR & PETERS, 1980c.

Genus Demoulinellus PESCADOR & PETERS Genus F of PESCADOR & PETERS, 1980b: 50. New genus F of PESCADOR; LANDA, SOLDÁN & PETERS, 1980: 171. Demoulinellus PESCADOR & PETERS, 1982: 10; SAVAGE, 1987b: 206; DOMÍNGUEZ et al., 1992: 23; DOMÍNGUEZ et al., 2001: 29, 40. (Type-species: Demoulinellus coloratus PESCADOR & PETERS, original designation).

Taxonomy: The genus Demoulinellus was established by PESCADOR & PETERS (1982) based on adults, eggs, and nymphs of D. coloratus collected from Chile. Previously (as Genus F), PESCADOR & PETERS (1980b) considered it a “phyletic enigma”. After studying

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the eggs, nymphs and adults, PESCADOR & PETERS (1982) proposed that Demoulinellus belonged to the same phyletic lineage as Zephlebia and related genera from New Zealand. Adult Characteristics: 1) Fork of vein MA of fore wings symmetrical (Fig. 128A); 2) hind wings without costal projection (Fig. 128B); 3) vein Sc of hind wings approximately 9/10 maximum length of hind wings (Fig. 128B); 4) tarsal claws of a pair dissimilar, one apically hooked and the other obtuse and pad-like (Fig. 128C); 5) penis lobes fused in basal half (Fig. 128D), each lobe with dorsal, semi-circularly arranged, subapical setae (Fig. 128E); and 6) female sternum IX entire and prominent apically (Fig. 128F). Nymphal Characteristics (general aspect as in Fig. 129A): 1) Clypeus slightly narrower than labrum, anteromedian emargination of labrum as in Fig. 130B; lateral margins of clypeus smoothly curved (Fig. 130A); 2) outer margin of mandibles curved with minute setae on basal half and tuft of median setae (Fig. 130C); 3) segment II of maxillary palpi approximately 1-1/3 times length of segment I, segment III approximately 1/2 length of segment II, inner apical margin of segment II with subapical setae (Fig. 130D); 4) segment II of labial palpi subequal in length to segment I, segment III 3/4 length of MA

A

Sc

F

B

ss

C E

D

G

Plate 128. Demoulinellus coloratus, imago. Figs. 128A-G: 128A, fore wing; 128B, hind wing; 128C, male tarsal claws; 128D, male genitalia, v.v.; 128E, penis lobe, d.v.; 128F, female sternum IX; 128G, abdominal terga IV-VII. (MA = fork of MA; ss = subapical setae). Figs. 128A-G modified from PESCADOR & PETERS, 1982.

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segment II (Fig. 130E); 5) tarsal claws with denticles progressively larger apically (Fig. 130F); 6) posterolateral projections on abdominal segments VI-IX; and 7) abdominal gills on segments I-VII similar, dorsal and ventral portions slender and apically tapered (Fig. 130G). Distribution: Chile (Bío Bío, Cautín, Chiloé, Coquimbo, Curico, Malléco, Osorno and Santiago Provinces). Biology: The nymphs of Demoulinellus occur in moderately flowing streams and rivers. They are excellent swimmers and mostly inhabit sun-exposed shallow areas among rocks, stones, and gravel. The adults have been collected from late January to late February.

A Plate 129. Demoulinellus coloratus, nymph. Fig. 129A, habitus. Figure modified from PESCADOR & PETERS, 1982.

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Demoulinellus coloratus PESCADOR & PETERS Demoulinellus coloratus PESCADOR & PETERS, 1982: 12 (male, female, nymph, egg).

Taxonomy: This is the only species in the genus. The pattern of dark brown maculae on the abdominal terga of nymphs (Fig. 129A) and adults (Fig. 128G) is characteristic. Distribution and Biology: Distribution and biology are discussed in the generic description. Genus Ecuaphlebia DOMÍNGUEZ Ecuaphlebia DOMÍNGUEZ, 1988b: 227; ZUÑIGA et al., 2004: 27. (Type-species: Ecuaphlebia rumignaui DOMÍNGUEZ, original designation).

Taxonomy: Ecuaphlebia was established in 1988, by DOMÍNGUEZ for male imagos and nymphs. Since then no other adults have been reported, nevertheless, nymphs assignable to this genus were collected from several localities in Ecuador and Colombia (ZUÑIGA et

A

C

D

B

E

F

G

Plate 130. Demoulinellus coloratus, nymph. Figs. 130A-G. 130A-E: Mouthparts. 130A, Clypeus and labrum, d.v.; 130B, anteromedian emargination of labrum, enlarged; 130C, left mandible; 130D, maxilla; 130E, labium (left dorsal, right ventral); 130F, fore claw; 130G, gill IV. Figs. 130A-G modified from PESCADOR & PETERS, 1982.

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al., 2004). For this reason, it is very likely that more species will be described when more adults will be available. Adult Characteristcs: 1) Stem of vein MA of fore wings sagged so MA fork sometimes slightly asymmetrical (Figs. 131A, D); 2) vein MP2 of fore wings attached at base to veins MP1 and CuA by cross veins (Figs. 131A, D); 3) vein ICu1 of fore wings attached to vein CuP; 4) hind wings with well developed costal projection (Figs. 131B-C, E); 5) vein Sc of hind wings ending about 7/10 distance from base to apex (Figs. 131B-C, E); 6) vein MP of hind wings forked (Fig. 131C); 7) tarsal claws of a pair dissimilar, one apically hooked and other obtuse, pad-like (Fig. 131H); and 8) penes divided, tubular; each lobe with a broad ventrolaterally oriented spine (Figs. 131F-G). MA

A

D

CuP

ICu1

CuA

B

MP2

MP1

MA2

CP

E

Sc

C MP

H

G

F I

J

Plate 131. Ecuaphlebia, adults. Figs. 131A-E: Wings. 131A-C, E. rumignaui. 131A, fore wing; 131B, hind wing; 131C, hind wing, enlarged. 131D-E, E. sp. A. 131D fore wing; 131E, hind wing. 131F-G, male genitalia, v.v.; 131F, E. rumignaui; 131G, E. sp. A. penes, v.v.; 131H, tarsal claw, E. rumignaui. 131I-J: abdominal terga II-VI: 131I, E. rumignaui, male imago; 131J, E. sp. A., male subimago. (CP = Costal Projection). Figs. 131A-J modified from DOMÍNGUEZ, 1988b.

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Nymphal Characteristics: 1) General aspect as in Fig. 132A; 2) lateral margins of labrum rounded, anteromedian emargination with 4-5 denticles (Figs. 133A-B); and labrum a little wider than clypeus; 3) mandible with a few setae on basal half of outer margin and a tuft of setae medially (Fig. 133C); 4) segment III of labial palpi as in Figs. 133F-G; 5) abdominal gills on segments I-VII lanceolate, dorsal and ventral portions similar with unbranched tracheae (Fig. 133H); 6) posterolateral projections on abdominal segments VI or VII-IX; and 7) denticles on tarsal claws progressively larger, except apical denticle much larger (Fig. 133I). Distribution: Ecuador and Colombia. Biology: Specimens of this genus have been collected from altitudes ranging from approximately 800 m to 3750 m. The nymphs were found in a variety of habitats, from slow rivers to fast running streams. KEY

ECUAPHLEBIA

TO

Imagos 1 –

CLAVE

Length of body 6.7-6.9 mm, cross veins of fore wings between veins C-Sc, Sc-R1 and R1-R2 clouded with brown (Fig. 131A) ............................. Ecuaphlebia rumignaui Length of body 8.2-8.3 mm, cross veins of fore wings between veins C-Sc, Sc-R1 and R1-R2 lightly clouded with gray (Fig. 131D) ............................. Ecuaphlebia sp. A PARA

ECUAPHLEBIA

Imagos 1 –

Longitud del cuerpo 6,7-6,9 mm; venas transversas de ala anterior entre venas C-Sc, Sc-R1 and R1-R2 sombreadas con castaño (Fig. 131A) ........... Ecuaphlebia rumignaui Longitud del cuerpo 8,2-8,3 mm; venas transversas del ala anterior entre venas C-Sc, Sc-R1 and R1-R2 sombreadas levemente con gris (Fig. 131D) ........ Ecuaphlebia sp. A

Ecuaphlebia rumignaui DOMÍNGUEZ Ecuaphlebia rumignaui DOMÍNGUEZ, 1988b: 231 (male).

Taxonomy: Ecuaphlebia rumignaui is known from a male imago and can be differentiated from the other species of the genus by the following combination of characters: 1) Length of body 6.7-6.9 mm; 2) cross veins of fore wings between veins C-Sc, Sc-R1 and R1R2 shaded with brown (Fig. 131A); and 3) abdominal color pattern as in Fig. 131I. Distribution: Ecuador (P.N. Cotopaxi). Ecuaphlebia sp. A Ecuaphlebia sp. A DOMÍNGUEZ, 1988b: 232 (male, nymph).

Taxonomy: This is the only species in the genus known from nymphs. As this species was described on the basis of a single male subimago, the author preferred not to give it a formal name. Ecuaphlebia sp. A can be distinguished from the other species by the

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following combination of characters: 1) Length of body 8.2-8.3 mm; 2) cross veins of fore wings between veins C-Sc, Sc-R1 and R1-R2 lightly shaded with gray (Fig. 131D); and 3) abdominal color pattern as in Fig. 131J. Distribution and Biology: Ecuador (Chimborazo). Nymphs of Ecuaphlebia sp. A were collected between 1800 and 2600 m in fast flowing, forested streams. The subimago was collected flying at 10:00 h.

A

Plate 132. Ecuaphlebia sp. A. Fig. 132A, nymphal habitus. Drawing by A. Dupuy.

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A

C

B

D

III

G

II

F

I

E D

H

I

Plate 133. Ecuaphlebia sp. A , nymph. Figs. 133A-G: Mouthparts. 133A, labrum, d.v.; 133B, anteromedian emargination, detail; 133C, left mandible; 133D, maxilla; 133E, hypopharynx; 133F, labium (left dorsal, right ventral); 133G, detail of labial palp segment III, dorsal. 133H, gill 2; 133I, fore claw. (D = denticle; Roman numbers = segment numbers). Figs. 133A-E modified from DOMÍNGUEZ, 1988b.

Genus Farrodes PETERS Farrodes PETERS, 1971: 5; DOMÍNGUEZ & SAVAGE, 1987: 43; SAVAGE, 1987b: 207; KLUGE, 1994: 247; DOMÍNGUEZ, MOLINERI & PETERS, 1996: 87; DOMÍNGUEZ, 1999: 156. (Type-species: Farrodes hyalinus PETERS, original designation). Homothraulus; EDMUNDS, JENSEN & BERNER, 1976: 224 (in this work, Homothraulus included Farrodes and Fig. 375 is the genitalia of Farrodes; in the key, Hermanellopsis included nymphs of Farrodes).

Taxonomy: Farrodes was established by PETERS (1971) for three species from the Antilles, and originally was restricted to these islands. Later, one species was described

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from Texas (DAVIS, 1987) and two from Argentina (DOMÍNGUEZ & SAVAGE, 1987). In 1996, DOMÍNGUEZ et al. reviewed the species of the F. bimaculatus group from Central and South America, describing 8 new species. In 1999, DOMÍNGUEZ published the cladistic analysis and biogeography of the genus, reviewing the F. caribbianus group, describing 3 new species and transferring 3 more from other genera. In the same analysis were included the other members of the Farrodes lineage: Homothraulus and Simothraulopsis raising the number of species of the genus from 6 in 1987 to more than 20 in 1999, making it one of the most widespread and species-rich genera of Leptophlebiidae in Central and South America. At the present time, it is almost impossible to separate to species the few nymphs known in the genus, and for this reason a key is not included. Adult Characteristics: 1) Male eyes slightly separated or meet medially on dorsum of head; 2) fork of vein MA of fore wings (Fig. 134A) asymmetrical; 3) vein MP of hind wings (Figs. 134B-C, L) unforked; 4) costal projection of hind wings acute and located near middle of wing (Figs. 134C, L); 5) styliger plate of male projected posterolaterally (Figs. 134D, F, 135A, E); 6) penes divided in apical 2/3, venter of each lobe with an apical appendage (Figs. 134D, F, H); 7) abdominal sternum IX of female apically blunt or slightly cleft (Fig. 135P); and 8) tarsal claws dissimilar, one apically hooked and other blunt (Fig. 135O). Nymphal Characteristics (general aspect as in Fig. 136A): 1) Labrum rounded laterally (Fig. 136B); 2) abdominal gills I-VII lanceolate, narrow, dorsal and ventral portions similar (Fig. 136D); 3) posterolateral spines on abdominal segments VIII and IX; and 4) denticles on tarsal claws equal in size, except apical denticle much larger (Fig. 136E). Distribution and Biology: Pan American from Texas to Northern Argentina, including also the Caribbean Islands. The nymphs of this genus are found in a variety of small to large streams and rivers over its wide geographical range. Nymphs have been collected from slow to fast current. Some among leaf packs, and others associated with stone and gravel substrates. This suggests a wide range of specific tolerance. In some species the adults were collected over an extended period of time, suggesting a long emergence period. The mating flight is still unknown. KEY

TO

SOUTH AMERICAN FARRODES

Male Imagos 1 – 2 (1) –

Projections of penes shaped as an inverted funnel (Fig. 135E ) or conical, curved inward (Fig. 134D) or curved apically upward (Fig. 135G) ........................................ ................................................................................... Farrodes caribbianus species group, 2 Projections of penes cylindrical, long (Figs. 134F, H, J, M) .................................................................................. Farrodes bimaculatus species group, 4 Medial projection of styliger plate present (Fig. 135E); projections of penes ventral, shaped as an inverted funnel (Fig. 135E); abdominal color pattern as in Fig. 135F ...................................................................................................... Farrodes roundsi Medial projection of styliger plate absent; projections of penes lateral and conical, curved apically inward (Fig. 134D), or upward (Fig. 135G); abdominal color pattern not as above .......................................................................................................... 3

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MP MA

CP

L

C MP

B

A

G D

E

F

I

J

H

K

M

N

Plate 134. Farrodes, imagos. Figs. 134A-E, F. caribbianus: 134A, fore wing; 134B, hind wing; 134C, hind wing, detail; 134D, male genitalia, v.v.; 134E, abdominal terga II-VII. 134F-G, F. carioca: 134F, male genitalia, v.v.; 134G, abdominal terga II-VII. 134H-I, F. iguazuanus: 134H, male genitalia, v.v.; 134I, abdominal terga II-VII. 134J-K, F. longispinus: 134J, male genitalia, v.v.; 134K, abdominal terga II-VII. 134L-N, F. maculatus: 134L, hind wing, detail; 134M, male genitalia, v.v.; 134N, abdominal segments III-VI, l.v. (CP = Costal Projection). Figs. 134A-E, L-N, modified from DOMÍNGUEZ, 1999; 134F-G, J-K, from DOMÍNGUEZ et al., 1996 and H-I, from DOMÍNGUEZ & SAVAGE, 1987.

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Posterolateral projections of styliger plate rounded, widely based (Fig. 134D); base of penes broad, angled, with small setae; apical projections of penes conical, recurved inwardly (Fig. 134D); abdominal pattern as in Fig. 134E .......................... ................................................................................................................. Farrodes caribbianus – Posterolateral projections of styliger plate with base well defined, apex rounded (Fig. 135G); base of penes roundly widened; projections of penes conical, not curved apically inward (Fig. 135G); abdominal color pattern as in Fig. 135H ....... ........................................................................................................................ Farrodes savagei 4 (1) Posterolateral projections of styliger plate short and wide (Fig. 134F); abdominal terga washed with black, color pattern as in Fig. 134G ..................... Farrodes carioca – Posterolateral projections of styliger plate long and narrow (Figs. 134J, 135A); abdominal color pattern not as above ........................................................................... 5 5 (4) Projections of penes rounded apically, with terminal flap folded (Fig. 135A); abdominal color pattern as in Fig. 135B .......................................... Farrodes ochraceous – Projections of penes not rounded apically, with terminal flap extended (Figs. 134J, 135C, K); abdominal color pattern not as above ........................................................ 6 6 (5) Lateral projections of penes with a long ventral spine basal to terminal flap (Fig. 134J); abdominal color pattern as in Fig. 134K ............................ Farrodes longispinus – Lateral projections of penes without long ventral spines (Figs. 135C, K); abdominal color pattern not as above ........................................................................... 7 7 (6) Posterolateral projections of styliger plate somewhat acute apically (Fig. 135C); abdominal color pattern as in Fig. 135D ............................................. Farrodes pakitza – Posterolateral projections of styliger plate rounded apically (Figs. 134M, 135I); abdominal color pattern not as above ........................................................................... 8 8 (7) Genital forceps segment I uniformly brown or grayish-brown, segments II and III lighter; abdominal segments with extensive whitish marks (Figs. 134N, 135L) .......... 9 – Basal 1/2 of genital forceps segment I yellow or yellowish-brown, apical 1/2 and segments II and III yellow-white; abdominal segments with small whitish marks (Figs. 134I, 135J, N) ......................................................................................................... 10 9 (8) Costal projection of hind wings large (more than 0.19 of total width) (as in Fig. 134C); apex of projections of penes forming a strong angle (Fig. 135K) .............. .......................................................................................................................... Farrodes xingu – Costal projection of hind wings small (less than 0.16 of total width), as in Fig. 134L; apex of projections of penes not forming a strong angle (Fig. 134M) ........ .................................................................................................................... Farrodes maculatus 10 (8) Costal projection of hind wings large (more than 0.19 of total width) (Fig. 134C); abdominal color pattern as in Fig. 135J .................................................... Farrodes tepui – Costal projection of hind wings small (less than 0.16 of total width), as in Fig. 134L; abdominal color pattern not as above ............................................................. 11 11 (10) Basal 1/2 of forceps segment I yellow, remainder of forceps yellowish-white; styliger plate yellow ..............................................................................Farrodes iguazuanus – Basal 1/2 of forceps segment I brown, washed with brownish-black; remainder of forceps yellowish-white; styliger plate brown, washed with brownish-black .... ................................................................................................................... Farrodes yungaensis

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CLAVE

PARA LOS

FARRODES

SUDAMERICANOS

Imagos Machos 1. – 2 (1) – 3 (2)

–

4 (1) – 5 (4) – 6 (5) – 7 (6) – 8 (7)

Proyecciones de los penes con forma de un embudo invertido (Fig. 135E ) o cónicas, curvadas hacia adentro (Fig. 134D) o curvadas apicalmente hacia arriba (Fig. 135G) .................................................. grupo de especies Farrodes caribbianus......2 Proyecciones de los penes cilíndricas, largas (Figs. 134F, H, J, M) ........................... ...................................................................... grupo de especies Farrodes bimaculatus......4 Proyección medial de la placa estilígera presente (Fig. 135E); proyección de los penes ventral, con forma de un embudo invertido (Fig. 135E); patrón de coloración abdominal como en la Fig. 135F ........................................ Farrodes roundsi Proyección medial de la placa estilígera ausente; proyecciones de los penes laterales y cónicas, curvadas apicalmente hacia adentro (Fig. 134D), o hacia arriba (Fig. 135G); patrón de coloración abdominal no como arriba ................................ 3 Proyecciones posterolaterales de la placa estilígera redondeadas, con base amplia (Fig. 134D); base de los lóbulos de los penes ensanchadas en ángulo recto, con pequeñas espinas; proyecciones de los penes cónicas, curvadas hacia dentro (Fig. 134D); patrón de coloración abdominal como en la Fig. 134E ............................... Farrodes caribbianus Proyecciones de la placa estilígera con una base bien definida, ápice redondeado (Fig. 135G); base de los lóbulos de los penes ensanchados y redondeados; proyecciones de los penes cónicas, no curvadas apicalmente hacia adentro (Fig. 135G); patrón de coloración abdominal como en la Fig. 135H ........................................... Farrodes savagei Proyecciones posterolaterales de la placa estilígera corta y ancha (Fig. 134F); tergos abdominales teñidos de negruzco, patrón de coloración abdominal como en la Fig. 134G ............................................................................................ Farrodes carioca Proyecciones posterolaterales de la placa estilígera larga y delgada (Figs. 134J, 135A); patrón de coloración abdominal no como arriba .......................................... 5 Proyecciones laterales de los penes redondeadas apicalmente, con una corta aleta terminal plegada (Fig. 135A ); patrón de coloración abdominal como en la Fig. 135B ................................................................................................................... Farrodes ochraceous Proyecciones laterales de los penes no redondeadas apicalmente, con una corta aleta terminal (Figs. 134J, 135C, K); patrón de coloración abdominal no como arriba ...... 6 Proyecciones laterales de los penes con una larga espina ventral, basal a la aleta terminal (Fig. 134J); patrón de coloración abdominal como en la Fig. 134K ........ .................................................................................................................. Farrodes longispinus Proyecciones laterales de los penes sin largas espinas ventrales (Figs. 135C, K); patrón de coloración abdominal no como arriba ....................................................... 7 Proyecciones posterolaterales de la placa estilígera algo aguzadas apicalmente (Fig. 135C); patrón de coloración abdominal como en la Fig. 135D ...... Farrodes pakitza Proyecciones posterolaterales de la placa estilígera redondeada apicalmente (Figs. 134M, 135I); patrón de coloración abdominal no como arriba ............................... 8 Segmento I de los fórceps castaños o castaño-grisáceo uniformes, segmentos II y III más claros; segmentos abdominales con marcas blancas extensas (Figs. 134N, 135L) ..................................................................................................................................... 9

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371

C

B

D

G

E

F

H

J

K

L

I

M O

P

N Plate 135. Farrodes, imagos. Figs. 135A-B, F. ochraceous: 135A, male genitalia, v.v.; 135B, abdominal terga II-VII. 135C-D, F. pakitza: 135C, male genitalia, v.v.; 135D, abdominal terga II-VII. 135E-F, F. roundsi: 135E, male genitalia, v.v. (forceps removed); 135F, abdominal terga II-VII. 135G-H, F. savagei: 135G, male genitalia, v.v.; 135H, abdominal terga II-VII. 135I-J, F. tepui: 135I, male genitalia, v.v.; 135J, abdominal terga II-VII. 135K-L, F. xingu: 135K, male genitalia, v.v.; 135L, abdominal terga IIVII. 135M-P, F. yungaensis: 135M, male genitalia, v.v.; 135N, abdominal terga II-VII; 135O, male fore tarsal claw; 135P, apex of female sternum IX. Figs. 135A-D, I-L, modified from DOMÍNGUEZ et al., 1996; 135E-H from DOMÍNGUEZ, 1999; 135M-N, P from DOMÍNGUEZ & SAVAGE, 1987.

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Mitad basal del segmento I de los fórceps amarilla o castaño-amarillenta, 1/2 apical y segmentos II y III amarillo-blanquecinos; segmentos abdominales con pequeñas marcas blanquecinas (Figs. 134I, 135J, N) ................................................ 10 9 (8) Proyección costal de las alas posteriores grande (más de 0,19 del ancho total) como en la Fig. 134C; ápice de las proyecciones peneanas formando un ángulo marcado (Fig. 135K) .................................................................................... Farrodes xingu – Proyección costal de las alas posteriores pequeña (menos de 0,16 del ancho total) como en la Fig.134L; ápice de las proyecciones peneanas no formando un ángulo marcado (Fig. 134M) ............................................................................. Farrodes maculatus 10 (8) Proyección costal de las alas posteriores grande (más de 0,19 del ancho total) (Fig. 134C); patrón de coloración abdominal como en la Fig. 135J ............. Farrodes tepui – Proyección costal de las alas posteriores pequeña (menos de 0,16 del ancho total) como en la Fig. 134L; patrón de coloración abdominal no como arriba ............ 11 11 (10) Mitad basal del segmento I de los fórceps amarillento, resto de los fórceps amarillo-blanquecinos; placa estilígera amarillenta ....................... Farrodes iguazuanus – Mitad basal del segmento I de los fórceps castaño, teñido de castaño-negruzo; resto de los fórceps amarillo-blanquecino; placa estilígera castaña, teñida de castaño-negruzco ................................................................................... Farrodes yungaensis Farrodes caribbianus (TRAVER) Thraulus caribbianus TRAVER, 1943: 79 (male, female); TRAVER, 1960b: 73; HUBBARD, 1982a: 267. Farrodes caribbianus; DOMÍNGUEZ, 1999: 159 (nymph).

Taxonomy: Thraulus caribbianus was described by TRAVER (1943), based only on adults. According to her, this species was allied to T. maculatus (now Farrodes maculatus), due to the type of hind wing and the uniqueness of the lateral appendages of the penes. Based on a cladistic analysis DOMÍNGUEZ (1999), transferred this species to the genus Farrodes. The imagos of this species can be separated from the other species of the genus by the following combination of characters: 1) Costal projection of hind wing large (more than 0.19 of total width, Fig. 134C); 2) posterolateral projections of styliger plate short, blunt and widely based (Fig. 134D); 3) apex of penis lobes produced in an obtuse angle (Fig. 134D); 4) base of penes widened at right angle, with few setae (Fig. 134D); 5) projections of penes lateral, conical, curved inward; 6) abdominal color pattern as in Fig. 134E. There is a wide variation in coloration among the specimens known, even from the same locality. In the nymph: 1) Spines on outer margin of hind tarsi long (Fig. 136C); 2) spines on dorsum of hind femora acute (Fig. 136C); 3) abdominal color pattern as in Fig. 134E. Distribution and Biology: Colombia, Venezuela, Costa Rica and Panama. The specimens were collected from sea level to around 1300 m. Based on collection data, the adults appear to emerge during most of the year. Farrodes carioca DOMÍNGUEZ, MOLINERI & PETERS Farrodes carioca DOMÍNGUEZ, MOLINERI & PETERS, 1996: 92 (male, female); DA-SILVA, 2002b: 2 (nymph).

Taxonomy: Farrodes carioca was described based on males and a female imago from Brazil. Later, DA-SILVA (2002b) described the possible nymph of this species, based on

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material from the same area where the adults were collected. Farrodes carioca can be separated from the other species of the genus by the following combination of characters: In the adult: 1) Costal projection of hind wing large (more than 0.19 of total width, Fig. 134C); 2) posterolateral projections of styliger plate short, wide, with apex blunt (Fig. 134F); 3) basal 1/4 of penis lobes fused; 4) apex of penis lobes rounded; 5) lobes of penes cylindrical, long, ending in a rounded short flap (Fig. 134F); and 6) abdominal color pattern as in Fig. 134G. In the nymphs (according to DA-SILVA, 2002): 1) One row of pectinate setae close to “broom” setae row at apex of maxilla, measuring about 2/3 of length (in other species row of pectinate setae more subapical and smaller); and 2) leg and abdominal pattern. Nevertheless, according to this author, this coloration is more similar to the one of F. hyalinus PETERS, from Jamaica. Distribution and Biology: Brazil (Rio de Janeiro). The adults were collected in April, at 950 m altitude. The nymphs came from different habitats (DA-SILVA, 2002b), one with rithral characteristics and the other with potamal ones. The nymphs were also found in different microhabitats, suggesting a generalist character. Farrodes iguazuanus DOMÍNGUEZ & SAVAGE Farrodes iguazuanus DOMÍNGUEZ & SAVAGE, 1987: 49 (male, female); HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 211; DOMÍNGUEZ, MOLINERI & PETERS, 1996: 89; DOMÍNGUEZ, 1999: 175.

Taxonomy: Farrodes iguazuanus is known only from imagos, which can be separated from the other species of this genus by the following combination of characters: 1) Costal projection of hind wing small (less than 0.16 of total width, Fig. 134L); 2) apical 2/3 of penes divided (Fig. 134H); 3) apex of penis lobes rounded; 4) lobes of penes long, cylindrical, ending in a short flap; 5) abdominal terga translucent, tinted variably with blackish brown, and terga II-VII with anteromedian area pale and divided by a much darker wedge which extends anteriorly from dark medial area (Fig. 134I); 6) basal 1/2 of segment I of forceps yellowish, remainder of segment I and segments II-III whitish yellow. Distribution and Biology: Argentina (Misiones). The adults are known to emerge between November and December. Farrodes longispinus DOMÍNGUEZ, MOLINERI & PETERS Farrodes longispinus DOMÍNGUEZ, MOLINERI & PETERS, 1996: 95 (male imago).

Taxonomy: Farrodes longispinus is known from a single male imago from Cerro de la Neblina Tepui, Venezuela. It can be separated from the other species of the genus by the following combination of characters: 1) Costal projection of hind wing large (more than 0.19 of total width) (Fig. 134C); 2) posterolateral projections of styliger plate long and narrow with apex blunt (Fig. 134J); 3) basal 1/2 of penes lobes fused; 4) apex of penis lobes rounded; 5) lateral projections of penes cylindrical, long, ending in a short flap, with a long ventral spine basal to terminal flap (Fig. 134J); 6) abdominal color pattern as in Fig. 134K. Distribution and Biology: Venezuela (T. F. Amazonas). The adults were collected in February, at an altitude of 140 m.

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F

C

D

A

E

B

D

Plate 136. Farrodes, nymphs. Figs. 136A-F. Fig. 136A, F. yungaensis, habitus. 136B-C, F. caribbianus: 136B, labrum, d.v.; 136C, leg III. 136D-E, F. yungaensis: 136D, gill IV; 136E, fore tarsal claw. 136F, F. savagei: leg III, d.v. (D = denticle). Fig. 136A, drawing by C. Molineri. Figs. 136B-C, F, modified from DOMÍNGUEZ, 1999; 136D-E from DOMÍNGUEZ & SAVAGE, 1987.

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Farrodes maculatus (NEEDHAM & MURPHY) Thraulus maculatus NEEDHAM & MURPHY, 1924: 45 male imago); ULMER, 1943: 28; TRAVER, 1947a: 149. Homothraulus maculatus; TRAVER, 1960b: 73; PETERS & TSUI, 1972: 565; HUBBARD, 1982a: 265. Farrodes maculatus; DOMÍNGUEZ, 1999: 162.

Taxonomy: This species was described from males imagos, stating that it was “near [Thraulus] misionensis” NEEDHAM & MURPHY (1924). Later, it was transferred to Homothraulus, following DEMOULIN’S (1955) transfer of T. misionensis. Finally, in his cladistic revision of the generic complex, DOMÍNGUEZ (1999) transferred it to Farrodes. Farrodes maculatus adults can be separated from the other species of the genus by the following combination of characters: 1) Costal projection of hind wings small (less than 0.16 of total width, Fig. 134L); 2) posterolateral projections of styliger plate long and narrow, base wide; 3) apical 1/2 of penes divided (Fig. 134M); 4) lateral projections of penes long, rather pointed apically with a small flap; and 5) abdominal color pattern as in Fig. 134N. Distribution and Biology: Argentina. Adults collected in March. Farrodes ochraceous DOMÍNGUEZ, MOLINERI & PETERS Farrodes ochraceous DOMÍNGUEZ, MOLINERI & PETERS, 1996: 94 (male imago); DOMÍNGUEZ, 1999: 175.

Taxonomy: Farrodes ochraceous was described from a single imago, from Reserva Ducke, near Manaus. The adults of this species can be separated from the other species of the genus by the following combination of characters: 1) Costal projection of hind wings large (more than 0.19 of total width, Fig. 134C); 2) posterolateral projections of styliger plate long and narrow, with apex acute (Fig. 135A); 3) basal 2/5 of penes fused; 4) apex of penis lobes rounded; 5) lateral projections of penes cylindrical, long and wide, with a short flap folded ventrally; and 6) abdominal segments washed with orange-brown (Fig. 135B). Distribution and Biology: Brazil (Amazonas). The adult was collected in February. Farrodes pakitza DOMÍNGUEZ, MOLINERI & PETERS Farrodes pakitza DOMÍNGUEZ, MOLINERI & PETERS, 1996: 97 (male imago); DOMÍNGUEZ, 1999: 175.

Taxonomy: Farrodes pakitza was described only from male imagos, collected at light and malaise traps. The adults of this species can be separated from the other species of the genus by the following combination of characters: 1) Costal projection of hind wings large (more than 0.19 of total width, Fig. 134C); 2) posterolateral projections of styliger plate long and narrow, with somewhat acute apex (Fig. 135C); 3) basal 1/2 of penes fused; 4) apex of penis lobes rounded; 5) lateral projections of penes cylindrical, long, ending in a small narrow flap (Fig. 135C); and 6) abdominal segments washed with black (Fig. 135D). Distribution and Biology: Peru (Madre de Dios). The adult was collected in September.

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Farrodes roundsi (TRAVER) Thraulus roundsi TRAVER, 1947a: 153 (male, female); TRAVER, 1960b: 73. Farrodes roundsi; DOMÍNGUEZ, 1999: 161.

Taxonomy: The male and female imagos of this species were described in Thraulus by TRAVER (1947), although she noted that the genitalia was so different that it might not belong to this genus. In 1999 DOMÍNGUEZ transferred it to Farrodes, although he stated that until the nymphs were known, the position would be uncertain. Farrodes roundsi adults can be separated from the other species of the genus by the following combination of characters: 1) Costal projection of hind wing large (more than 0.19 of total width, Fig. 134C); 2) posterolateral projections of styliger plate medium-sized, cylindrical, and curved inwardly (Fig. 135E); 3) medial projection of styliger plate present, shaped as in Fig. 135E; 4) ventral projection of penes shaped as an inverted funnel; 5) abdominal segments translucent yellowish to olive-brown, coloration pattern as in Fig. 135F. Distribution and Biology: Panama, Costa Rica and Colombia. Adults were collected from November to February. Farrodes savagei DOMÍNGUEZ Farrodes savagei DOMÍNGUEZ, 1999: 164 (male, female, nymph).

Taxonomy: This species was described based on adults and nymphs from the Perija mountains, associated by rearing. Farrodes savagei can be separated from the other species of the genus by the following combination of characters. In the imagos: 1) Costal projection of hind wing large (more than 0.19 of total width, Fig. 134C); 2) posterolateral projections of styliger plate medium sized, cylindrical, straight and rounded apically (Fig. 135G); 2) basal 1/4 of penes fused; 3) ventral projections of penes conical, curved apically upward (Fig. 135G); and 4) abdominal terga II-VI hyaline, with black markings as in Fig. 135H. In the nymph: 1) Spines on outer margin of tarsi short (Fig. 136F); 2) thick setae on basal half of outer margin of labial palpi II; 3) abdominal pattern as in male imago (Fig. 135H). Distribution and Biology: Venezuela (Zulia). Adults were collected in October and November. Farrodes tepui DOMÍNGUEZ, MOLINERI & PETERS Farrodes tepui DOMÍNGUEZ, MOLINERI & PETERS, 1996: 98 (male, female); DOMÍNGUEZ, 1999: 175.

Taxonomy: This species was described based on adults of both sexes from Cerro de La Neblina Tepui. Farrodes tepui can be separated from the other species of the genus by the following combination of characters: 1) Costal projection of hind wing large (more than 0.19 of total width, Fig. 134C); 2) posterolateral projections of styliger plate long and narrow, with apex blunt (Fig. 135I); 3) basal 3/5 of penes fused; 4) apex of penis lobes rounded; 5) lateral projections of penes cylindrical, long, ending in a short flap (Fig. 135I) and 6) abdominal segments yellowish translucent, washed with black as in Fig. 135J. Distribution and Biology: Venezuela (Amazonas). Adults were collected in March.

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Farrodes xingu DOMÍNGUEZ, MOLINERI & PETERS Farrodes xingu DOMÍNGUEZ, MOLINERI & PETERS, 1996: 97 (male, female); DOMÍNGUEZ, 1999: 175.

Taxonomy: This species was described based on adults of both sexes from Rio Xingu, Brazil. Farrodes xingu can be separated from the other species of the genus by the following combination of characters: 1) Costal projection of hind wing large (more than 0.19 of total width, Fig. 134C); 2) projections of styliger plate long and narrow, with apex rounded (Fig. 135K); 3) approximately basal 1/3 of penes fused; 4) apex of penis lobes rounded; 5) lateral projections of penes cylindrical, long, ending in a short flap (Fig. 135K); 6) forceps segment I greyish-brown, segments II-III yellowish-white; and 7) abdominal segments with extensive whitish marks (Fig. 135L). Distribution and Biology: Brazil (Pará). Adults were collected in October. Farrodes yungaensis DOMÍNGUEZ & SAVAGE Farrodes yungaensis DOMÍNGUEZ & SAVAGE, 1987: 45 (male, female, nymph); HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 211; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 43; DOMÍNGUEZ, MOLINERI & PETERS, 1996: 88; DOMÍNGUEZ, 1999: 175.

Taxonomy: This species was described based on imagos of both sexes and nymphs from Northwestern Argentina. Farrodes yungaensis can be distinguished from the other species of the genus by the following combination of characters. In the imago: 1) Costal projection of hind wing small (less than 0.16 of total width, Fig. 134L); 2) projections of styliger plate long and narrow, with apex rounded (Fig. 135M); 3) approximately basal 1/2 of penes fused; 4) apex of penis lobes rounded; 5) lateral projections of penes cylindrical, long, ending in a short flap (Fig. 135M); 6) segment I of genital forceps brown with the remainder yellowish white and 7) the abdominal terga dark blackish-brown, with the anterolateral angles, and the anterosubmedian and mediolateral areas pale (Fig. 135N). In the nymph: 1) Spines on outer margin of tarsi short, as in Fig. 136F; 2) thick setae along outer margin of labial palpi segment II; 3) the abdominal color pattern similar to that of the male imago (Fig. 135N). Distribution and Biology: Mountain rain forest (Yungas) of Northwestern Argentina (Tucumán and Salta Provs) and Southwestern Bolivia (up to Cochabamba), between 500 and 1200 m altitude. The nymphs were collected in a rocky bottomed stream with a width of 10 to 15 m and some 45-70 cm depth. The nymphs were found on the underside of the rocks or in the leafpacks from small shaded stream 3 to 4 m wide to rivers with strong currents. Mature nymphs were collected in April. Imagos were collected between November and April, and adult emergence appears to last at least until May. The subimagos emerge at dusk and molt at dawn. Genus Fittkaulus SAVAGE & PETERS Fittkaulus SAVAGE & PETERS, 1978: 293; SAVAGE, 1986: 257; ORTH et al., 2000: 30. (Type-species: Fittkaulus maculatus SAVAGE & PETERS, original designation).

Taxonomy: Fittkaulus was established by SAVAGE & P ETERS (1978) for the species F. maculatus from northern Brazil, based on male and female imagos and

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nymphs. In 1986, S AVAGE revised the genus and described two additional species based solely on females. Adult Characteristics: 1) Male upper portion of eyes on tall stalk (Fig. 137A), with 10-13 large facets in longest row; 2) vein ICu1 of fore wings joined at base to vein CuA (Figs. 137B, E); 3) hind wings (Figs. 137C-D, F-G) small (about 1/7 the length of fore wings); 4) tarsal claws of a pair similar, with apical hook and opposing blunt lobe (Fig. 137H); 5) genital forceps two-segmented (Fig. 137I); 6) penes short and tubular, fused in basal 2/3 and without projections (Fig. 137I); and 7) female abdominal sternum IX deeply cleft apically (Fig. 137J). Nymphal Characteristics: 1) Head hypognathous; 2) labrum (Fig. 138A) with amteromedian emargination well developed; 3) linguae of hypopharynx without lateral processes (Fig. 138B); 4) large apically curved unpectinate seta near a large pectinate seta on anterior internal angle of maxillae (Fig. 138C); 5) glossae curved ventrally (Fig. 138D); 6) tarsal claws with 5-6 short basal denticles and 4-5 larger, sharper subapical denticles (Fig. 138E); 7) gills on abdominal segments I-VII with ventral portion smaller (3/5-4/5 length dorsal portion); gill I (Fig. 138F) and gill VII lanceolate, gills II-VI with large inner and small outer lobe on dorsal portion, lobes not developed on ventral portion (Fig. 138G); 8) posterolateral spines on abdominal segments III-VI and VIII-IX; and 9) caudal filaments with swimming setae (basal 2/5-3/5 of filaments with long setae on inner margin of cerci and lateral margins of terminal filament). Distribution: Surinam, French Guiana and Brazil. Unpublished records from Peru and Ecuador. Biology: The nymphs of Fittkaulus were collected in small streams, 3-5 m wide, at a depth of 20-50 cm, with sandy and rocky bottoms. The nymphs were on leaves and large detritus generally, but some were in fine detritus or sand in calm areas. The adults were collected in March and April. KEY

SOUTH AMERICAN FITTKAULUS

TO

Imagos 1 – 2(1)

–

Fore wings with a dark macula on the fork of vein MA (Figs. 137B, E); abdominal terga as in Figs. 137K-L .................................................................................................... 2 Fore wings without dark marks on the fork of vein MA; abdominal terga as in Fig. 137M .................................................................................................... Fittkaulus cururuensis Apex of hind wings obtuse, broadly rounded, costal projection of hind wings well developed, with apex located 7/10 or more distance from base of wing to apex (Figs. 137C-D); color pattern of abdominal terga as in Fig. 137K. .......................... ................................................................................................................ Fittkaulus maculatus Apex of hind wings acute, rounded, costal projection developed, with apex located less than 7/10 of the distance from the base of wing to apex (Figs. 137FG); color pattern of abdominal terga as in Fig. 137L .....................Fittkaulus cuiabae

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MP

379

MA

B

E

C

ICu1 CuA

F CP

Sc

A D

G

I

J

H

K

L

M

Plate 137. Fittkaulus, imagos. Figs. 137A-D, F. maculatus, male: 137A, eyes, l. v.; 137B, fore wing; 137C, hind wing; 137D, hind wing, detail. Figs. 137E-G, F. cuiabae, female: 137E, fore wing; 137F, hind wing; 137G, hind wing, detail. Figs. 137H-K, F. maculatus: 137H, fore tarsal claw, male; 137I, male genitalia, v.v.; 137J, female sternum IX; 137K, abdominal terga II-VII. Fig. 137L, F. cuiabae, female: abdominal terga IV-VII. Fig. 137M, F. cururensis female: thorax and abdominal terga I-IV. (CP = Costal Projection). Figs. 137A-M modified from SAVAGE, 1986.

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CLAVE

PARA LOS

FITTKAULUS

SUDAMERICANOS

Imagos 1 – 2(1)

–

Alas anteriores con una mancha oscura en la bifurcación de la vena MA (Figs. 137B, E); tergos abdominales como en las Figs. 137K-L ......................................... 2 Alas anteriores sin marcas oscuras en la bifurcación de la vena MA; tergos abdominales como en la Fig. 137M .............................................. Fittkaulus cururuensis Ápice de las alas posteriores obtuso, ampliamente redondeado, proyección costal bien desarrollada, conel ápice ubicado a 7/10 ó más allá de la distancia de la base al ápice (Figs. 137C-D); patrón de coloración abdominal como en la Fig. 137K .. ................................................................................................................ Fittkaulus maculatus Ápice de las alas posteriores redondeado, con base amplia, proyección costal desarrollada y con el ápice localizado a menos de 7/10 de la distancia de la base al ápice alar (Figs. 137F-G); patrón de coloración abdominal como en la Fig. 137L .....................................................................................................................Fittkaulus cuiabae

Fittkaulus cuiabae SAVAGE Fittkaulus cuiabae SAVAGE, 1986: 265 (female).

Taxonomy: The imagos of F. cuiabae can be separated from the other species of the genus by the following combination of characters: 1) Fore wings with dark macula at fork of vein MA (Fig. 137E); 2) apex of hind wings rounded and widely based (Figs. 137F-G); 3) hind coxae with two blackish brown maculae, one large and other small; and 4) abdominal sterna I-II without small brown posteromedian marks, and abdominal terga as in Fig. 137L. Distribution and Biology: Northern and central Brazil. From the dates of collection, the adults emerge in April. Fittkaulus cururuensis SAVAGE Fittkaulus cururuensis SAVAGE, 1986: 268 (female); DOMÍNGUEZ, ZUÑIGA & MOLINERI, 2002: 462.

Taxonomy: The imagos of F. cururuensis can be separated from the other species of the genus by the following combination of characters: 1) Fore wings without dark spot at fork of vein MA; 2) apex of hind wings acute and rounded; 3) hind coxae with two large blackish brown maculae; and 4) abdominal sterna I-II with small brown posteromedian marks, and abdominal terga as in Fig. 137M. Distribution and Biology: Northern Brazil. The adults were collected in February. Fittkaulus maculatus SAVAGE & PETERS Fittkaulus maculatus SAVAGE & PETERS, 1978: 294 (male, female, nymph); HUBBARD, 1982a: 264; SAVAGE, 1986: 262 (male, female, nymph); DOMÍNGUEZ, ZUÑIGA & MOLINERI, 2002: 462.

Taxonomy: The imagos of F. maculatus can be separated from those of the other species of the genus by the following combination of characters: 1) Fore wings with dark macula at fork of vein MA (Fig. 137B); 2) apex of hind wings blunt and widely rounded

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US

B A D

C

E

D

F G

Plate 138. Fittkaulus, nymphs. Figs. 138A-G, F. maculatus: 138A, clypeus and labrum, d.v.; 138B, hypopharynx, v.v.; 138C, maxilla; 138D, labium: left dorsal, right ventral; 138E, fore claw; 138F, gill I; 138G, gill IV. (US = unpectinate setae; D = denticle). Figs. 138A-G modified from SAVAGE, 1986.

(Figs. 137C-D); 3) hind coxae with two small blackish brown maculae; and 4) abdominal sterna I-II without small posteromedian marks, and abdominal terga as in Fig. 137K. This is the only species for which nymphs are known. They can be characterized by the following combination of characters: 1) Anteromedian emargination of labrum with six small denticles, apically flattened (Fig. 138A); and 2) legs without blackish brown maculae. Distribution and Biology: Surinam and Northern Brazil. The nymphs were collected in small sandy streams in parts with exposed rocks. The majority of the nymphs were collected from detritus in areas of slow current. The adults were collected in March. Genus Gonserellus PESCADOR Gonserellus PESCADOR, 1997: 237. (Type-species: Gonserellus atopus PESCADOR, original designation).

Taxonomy: The genus Gonserellus was established by PESCADOR (1997), from unique nymphs from Southern Chile.

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Adult Characteristics: Unknown. Nymphal Characteristics (general aspect as in Fig. 139A): 1) Clypeus slightly narrower than labrum, with lateral margins parallel (Fig. 139B); 2) lateral margins of labrum smoothly curved (Fig. 139B); 3) anteromedian emargination of labrum broad, U-shaped with 5 denticles (Fig. 139C); 4) outer margins of mandibles smoothly curved, with fine setae and a submedian seta tuft (Fig. 139D); 5) galea-lacinia of maxillae with a long, sharp subapical tusk (Fig. 139E); 6) bipectinate setae on dorsal surface of tibia; 7) tarsal claws with small equal-sized basal denticles, remainder of denticles progressively larger apically, with the apical one enlarged (Fig. 139F); 8) gills on abdominal segments I-VII similar, narrow and tapered apically (Fig. 139G); and 9) posterolateral spines present on abdominal segments VI-IX.

A

B

C D

D

T

D

E

G

F Plate 139. Gonserellus, nymph. Figs. 139A-G, G. atopus: 139A, habitus; 139B, clypeus and labrum, d. v.; 139C, anteromedian emargination of labrum, detail; 139D, left mandible, d.v.; 139E, left maxila, d.v.; 139F, fore claw; 139G, gill 4. (D = denticle, T = tusk). Figs. 139A-G, modified from PESCADOR, 1997.

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Distribution: Chile (Comuna Los Lagos, Provincia de Valdivia, X Region., Rio Quinchilca 39º51´S). Biology: Mature nymphs of this species were collected in a cold water river from Southern Chile. Gonserellus atopus PESCADOR Gonserellus atopus PESCADOR, 1997: 241 (nymph).

Taxonomy: Gonserellus atopus is the only species known from the genus. It has a dark brown color pattern on the abdomen (Fig. 139A), but the range of variation for the species is unknown. Distribution and Biology: Discussed in the generic description. Genus Hagenulopsis ULMER Hagenulopsis ULMER, 1920a: 34; NEEDHAM & MURPHY, 1924: 43; TRAVER, 1946: 427; PETERS, 1969: 258; EDMUNDS, JENSEN & BERNER, 1976: 221; PETERS & DOMÍNGUEZ, 2001: 353; ORTH et al., 2000: 30. (Type-species: Hagenulopsis diptera ULMER, original designation).

Taxonomy: Hagenulopsis was described by ULMER (1920a) based on male and female adults from Santa Catarina, Brazil. NEEDHAM & MURPHY (1924) later gave a superficial description of a nymph from Peru, which they associated with this genus because of a lack of hind wing pads. In 1943 SPIETH described another species of this genus (H. minutus) based on a dipterous female with an ovipositor, differentiating it from the species described by ULMER on the basis of its small size. TRAVER (1944) fully described the nymph of Hagenulopsis as “Hagenulopsis ally”. Later, in 1946, TRAVER indicated that the female description of Choroterpes emersoni NEEDHAM & MURPHY, was actually that of the female of H. minuta. From the same original material of NEEDHAM & MURPHY, TRAVER (1946) found and described 2 male imagos of H. minuta. PETERS & DOMÍNGUEZ (2001) reviewed the genus, describing the nymph of H. minuta for the first time. Adult Characteristics: 1) Fork of vein MA of fore wings symmetrical or asymmetrically sagged (Fig. 140A); 2) vein ICu1 of fore wings attached at base to vein CuP; 3) hind wings absent; 4) tarsal claws of a pair dissimilar, one apically pointed, other obtuse, pad-like; 5) penes (Fig. 140B) divided, tubular, apically acute, with two small ventral spines located about 1/4 distance from apex; 6) inner angle of forceps segment I located about 1/2 distance from base (Fig. 140B); 7) female with well developed and apically acute egg guide (Fig. 140D) and 8) female sternum IX deeply cleft apically (Fig. 140E). Nymphal Characteristics: 1) Labrum with margins rounded (Fig. 140F), and anteromedian emargination (Fig. 140G) with 5 denticles (usually three small and two large, although relative size and apparent number can vary); 2) mandible with few or no setae on apical third of outer margin (Fig. 140H); 3) apical 1/2 of segment III of labial palpi constricted (Fig. 140I); 4) denticles of tarsal claws progressively larger toward apex, except apical denticle much larger (Fig. 140J); 5) gills on abdominal segments I-VII lanceolate, narrow, dorsal and ventral portions similar (Fig. 140K); 4) posterolateral spines on

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abdominal segments VI-IX, larger posteriorly; and 5) acute projections at apex of sternum IX (similar to female, Fig. 140E). Distribution: Brazil, Colombia, French Guiana, Peru, Surinam, Guyana, Venezuela, north to Honduras. Also West Indies. Biology: The nymphs of this genus were collected in mountain streams, with rocky bottoms, under cobbles or submerged logs. The adults were collected from December to March. The swarm, with the typical up and down flight pattern, was observed in late morning, under a tree canopy, about 2-3 m above the stream. KEY

HAGENULOPSIS

TO

Imagos 1

–

CLAVE

29

Body length 3.0 to 3.5 mm; fore wings 3.0 to 4.0 mm; cross veins of fore wings pale to hyaline, lighter than longitudinal veins, not surrounded with dark clouds (Fig. 140A); well developed bridge between stalks of upper portion of male eyes (Fig. 140C); caudal filaments with darker annulations at articulations only at base ................................................................................................................. Hagenulopsis minuta Body length 6.0 to 7.0 mm; fore wings 6.0 to 6.5 mm; cross veins of fore wings blackish brown with dark clouds surrounding veins heaviest in costal area, cross veins darker than longitudinal veins; upper portion of male eyes without stalk or bridge between eyes; caudal filaments with darker annulations along most of length ..................................................................................................... Hagenulopsis diptera PARA

HAGENULOPSIS

30

Imagos 1

–

Cuerpo 3,0 a 3,5 mm; alas anteriores 3,0 a 4,0 mm; venas transversas de las alas anteriores pálidas a hialinas, más claras que las venas longitudinales, no sombreadas de castaño (Fig. 140A); un puente bien desarrollado entre la porción dorsal de los ojos de los machos (Fig. 140C); anillos más oscuros en la articulación de los segmentos, solo en la base de los filamentos caudales .................. Hagenulopsis minuta Cuerpo 6,0 a 7,0 mm; alas anteriores 6,0 a 6,5 mm; venas transversas de las alas anteriores castaño-negruzcas, con sombreado oscuro alrededor de las venas, más fuerte en el área costal, venas transversas más oscuras que las venas longitudinales; sin un puente entre la porción dorsal de los ojos de los machos; anillos más oscuros en las articulaciones de los segmentos, en casi toda la longitud de los filamentos caudales ............................................................................. Hagenulopsis diptera

Although the nymph of H. diptera is not known with certainty, the presence of wing markings in wing pads and the markings on the caudal filaments should serve to distinguish the nymphs of these two species. 30 Aunque la ninfa de H. diptera no es conocida con seguridad, la presencia de manchas en las venas en desarrollo, y las manchas en los filamentos caudales debería servir para separar las ninfas de estas especies. 29

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MA

A

385

D CuP

Eg

ICu1

B

C

E

F I

III II

H

G

I D

D

J

K

Plate 140. Hagenulopsis. Figs. 140A-K, H. minuta. 140A-E, imago: 140A, male fore wing; 140B, male genitalia, v.v.; 140C, male head, d. v.; 140D, female abdominal segments VII-X, l.v.; 140E, female sternum IX. Figs. 140F-K, nymph: 140F, labrum, d. v; 140G, anteromedian emargination of labrum; 140H, left mandible; 140I, labium (left dorsal, right ventral); 140J, fore claw; 140K, gill IV. (Eg = Egg guide; D = denticle; Roman numbers = segment numbers). Figs. 140A-K modified from PETERS & DOMÍNGUEZ, 2001.

Hagenulopsis diptera ULMER Hagenulopsis diptera ULMER, 1920a: 34 (male, female); LESTAGE, 1922a: 33; EDMUNDS, JENSEN & BERNER, 1976: 221; HUBBARD, 1982a: 264; PETERS & DOMÍNGUEZ, 2001: 356.

Taxonomy: The imagos of Hagenulopsis diptera can be separated from the other species of the genus by the following combination of characters: 1) Length of body 6.07.0 mm; 2) length of fore wings 6.0-6.5 mm; 3) fore wings of males and females with blackish marks around cross veins, cross veins darker than longitudinal veins; 4) cross veins in fore wing numerous (17-18 in costal area and 12-13 in subcostal area); 5) no bridge present between upper portions of male eyes; 6) dark annulations at articulations along most of the length of caudal filaments. Distribution: Brazil.

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Hagenulopsis minuta SPIETH Hagenulopsis minutus SPIETH, 1943: 10 (female); TRAVER, 1946: 427 (male); HUBBARD, 1982a: 264. Hagenulopsis minuta; PETERS & DOMÍNGUEZ, 2001: 354.

Taxonomy: Hagenulopsis minuta was described by SPIETH (1943) from a female imago. Later, TRAVER (1946) found that the allotype and two male specimens of the type-series of Choroterpes emersoni were actually H. minuta and subsequently described the males of this species. Additional information was given by PETERS & DOMÍNGUEZ (2001). The imagos of Hagenulopsis minuta can be distinguished from the other species of the genus by the following combination of characters: 1) Length of body 3.0-3.5 mm; 2) few cross veins in fore wings, approximately 4-8 cross veins in costal area and 4-7 in subcostal area (Fig. 140A); 3) well developed bridge between stalks of upper portion of male eyes (Fig. 140C); 4) fore wings of males and females without blackish clouds around cross veins (Fig. 140B); and 5) dark annulations at articulations of only basal part of caudal filaments of imagos of both sexes. The nymphs of this species have unicolorous caudal filaments, and there are no clouds around the cross veins in the developing wing pads (as in the adults). Distribution and Biology: Guyana, Surinam and Venezuela. Biology as in generic discussion. Genus Hagenulus EATON Hagenulus EATON, 1882: 207. (Type-species: Hagenulus caligatus EATON, original designation).

Taxonomy: The genus was known only from the Caribbean. Recently, PETERS et al. (2005) described a new species, based on adults, from Ecuador. The described nymphs of this genus may be very different, and as the nymphs of the South American species are not known, we prefer neither discuss the nymphs here nor include this genus in the nymphal generic key. Adult Characteristics: 1) Eyes of male separated on meson of head; 2) fork of vein MP of fore wings symmetrical to slightly asymmetrical (Fig. 141A); 3) vein ICu1 of fore wings attached to CuP (Fig. 141A); 4) costal projection of hind wing very long, located at apex of wing (Figs. 141B-C); 5) genital forceps segments II and III short, segment I long, base of forceps broad, inner margin forming an angular bend (Fig. 141D); 6) penes divided, tubular (Fig. 141D). Nymphal Characteristics: Unknown. Distribution: Ecuador. Also Cuba, Haiti, Jamaica and Puerto Rico. Hagenulus marshali PETERS, FLOWERS, HUBBARD, DOMÍNGUEZ & SAVAGE Hagenulus marshali PETERS et al., 2005: 56.

Taxonomy: This species was described from two male imagos. Although the fork of vein MP of the fore wings in this species is not symmetrical as in the other species of Hagenulus, the authors placed this species in Hagenulus because of the unique condition of the hind wings, the attachment of ICu1 to vein CuP, and genital forceps typical for the

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III II

A D

CuP CP

ICu1

MP2

B

I

MP1

C E III

IV

V

Plate 141. Hagenulus, imago. Figs. 141A-C, Hagenulus sp., 141A, fore wing; 141B, hind wing; 141C, hind wing, detail; 141D, male genitalia, v.v.; 141E, abdominal segments III-V, l.v. (CP = Costal Projection; Roman numbers = segment numbers).

genus. The slightly asymmetrical fork of vein MP may eventually lead to placement of the species in another genus or subgenus. The male imago of H. marshali can be distinguished from the other species of the genus by the following combination of characters: 1) Fork of vein MP of fore wings asymmetrical (MP2 attached to MP1 at a cross vein) (Fig. 141A); 2) hind wings as in Figs. 141B-C; 3) penes without spines and slightly twisted (Fig. 141D); and 5) abdominal color pattern as in Fig. 141E. Distribution: Ecuador (30 Km W of Puyo). Genus Hapsiphlebia PETERS & EDMUNDS Atalophlebia EATON, 1881: 91 [partim]; DEMOULIN, 1955b: 10 [partim]; DEMOULIN, 1955e: 6 [partim]. Hapsiphlebia PETERS & EDMUNDS, 1972: 1401; TSUI & PETERS, 1975: 548; PESCADOR & PETERS, 1980b: 43. (Type-species: Atalophlebia anastomosis DEMOULIN, original designation).

Taxonomy: The type species was described originally by DEMOULIN (1955b) in Atalophlebia. Later, PETERS and EDMUNDS (1972) established Hapsiphlebia and designated H. anastomosis (DEMOULIN) as the type species. PESCADOR & PETERS (1980b) analyzed the phylogenetic relationships of this genus, as well as the other Southern South America leptophlebiids. Presently, this genus is known only from the type species. Adult Characteristics: 1) Fork of vein MA of fore wings symmetrical (Fig. 142A); 2) cross veins anastomosed in stigmatic area (Fig. 142A); 3) vein Sc of hind wings approximately 9/10 length of hind wings (Fig. 142B); 4) claws of a pair similar, both apically hooked, each with an opposing hook (Fig. 142C); 5) penis lobes divided and tubular (Fig. 142D), and each lobe with one subapical, ventral spine; and 6) female with

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small egg guide extended slightly past anterior margin of segment VIII; and 7) female sternum IX deeply cleft apically (Fig. 142E). Nymphal Characteristics (Fig. 143A): 1) Clypeus and labrum subequal in width; (Fig. 143B), length of labrum approximately 1/2 maximum width; 2) anteromedian emargination of labrum (Fig. 143C) shallow, U-shaped, with 4-6 subequal sized denticles; 3) outer margin of mandibles straight to slightly curved with long thick setae on basal 1/ 2 (Fig. 143D); 4) segment II of maxillary palpi 4/5 length of segment I, segment III equal to or little longer than segment II, inner margin of segment II with long setae (Fig. 143E); 5) submentum of labium with long lateral setae and glossae strongly curved ventrally (Fig. 143F); 6) tarsal claws with small, subequal denticles (Fig. 143G); 7) posterolateral spines on segments II-IX; and 8) abdominal gills I-VII dissimilar, gills I-V plate-like with well developed dorsal and ventral portions (Fig. 143H), gills VI and VII reduced to slender filaments (Fig. 143I). Distribution and Biology: Chile (Magallanes province (ca. 53ºS, XII region) north to Coquimbo Province (ca. 30ºS IV region). The nymphs live in a wide variety of lotic environments from old irrigation canals and small streams to large rivers. The nymphs are most abundant in small shallow streams partially buried in crevices or on surfaces of stones, rocks or any solid objects heavily coated with silt or mud. Partially decomposed leaves trapped in quiet shallow sections of streams are also densely populated by the nymphs. Collection records indicate that nymphs were found from November to January, and adults from December to January.

MA

A

Sc

D

C

B

E

F

Plate 142. Hapsiphlebia, imago. Figs. 142A-F, H. anastomosis: 142A, fore wing; 142B, hind wing; 142C, male tarsal claw; 142D, male genitalia, v.v.; 142E, female sternum IX; 142F, abdominal terga III-VI. Figs. 142A-F modified from PETERS & EDMUNDS, 1972.

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Hapsiphlebia anastomosis (DEMOULIN) Atalophlebia anastomosis DEMOULIN, 1955b: 10 (male, female); DEMOULIN, 1955e: 6 (female). Hapsiphlebia anastomosis; PETERS & EDMUNDS, 1972: 1401 (male, female, nymph); TSUI & PETERS, 1975: 548 (male, nymph); HUBBARD, 1982a: 26.

Taxonomy: Hapsiphlebia anastomosis was originally described in Atalophlebia by DEMOULIN (1955). PETERS & EDMUNDS (1972) subsequently transferred and designated it as the type species of Hapsiphlebia. H. anastomosis is the only species described in the genus, but can probably be characterized by the structure of the male genitalia and the distinctive

B C

D

A

E

F

G

H

I

Plate 143. Hapsiphlebia, nymph. Figs. 143A-I, H. anastomosis nymph: 143A, habitus; 143B, clypeus and labrum, d. v.; 143C, anteromedian emargination of labrum, detail; 143D, left mandible; 143E, maxilla; 143F, labium, left dorsal, right ventral; 143G, fore claw; 143H gill III; 143I, gill VII. Figs. 143A-I modified from PETERS & EDMUNDS, 1972.

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color pattern of nymphs and imagos (Fig. 142F); body lengths of male and female imagos are 13.0-14.0 mm and 13.0-15.0 mm respectively. Distribution and Biology: Distribution and biology are discussed in the generic description. Genus Hermanella NEEDHAM & MURPHY Hermanella NEEDHAM & MURPHY, 1924: 39; ULMER, 1939: 490; SPIETH, 1943: 8; TRAVER, 1947a: 158; DEMOULIN, 1955a: 7; TRAVER, 1960a: 19; EDMUNDS, JENSEN & BERNER, 1976: 221; SAVAGE, 1987b: 207; DOMÍNGUEZ & FLOWERS, 1989: 555; DOMÍNGUEZ & CUEZZO, 2002: 149; ORTH et al., 2000: 30. (Type-species: Hermanella thelma NEEDHAM & MURPHY, original designation).

Taxonomy: The genus Hermanella was established by NEEDHAM & MURPHY (1924) for the species H. thelma from two nymphs from “Cataratas del Iguazú”, Argentina. A good part of the description referred to the wings based on the wing pads. The illustrations apparently represented two nymphs, although they differed somewhat from each other. SPIETH (1943) noted NEEDHAM & MURPHY’s (1924) error with respect to the geographic locality (but cited H. thelma as H. velma) and described a new species, Hermanella incertans, from Surinam. TRAVER (1947) considered it improbable that the adult described by SPIETH as Hermanella and the nymphs illustrated by NEEDHAM & MURPHY belonged to the same genus. Then DEMOULIN (1955) separated the species into two subgenera, Hermanella (s.s.) thelma and Hermanella (Hermanellopsis) incertans, and illustrated a nymph he believed belonged to H. (Hermanellopsis). TRAVER (1960) described, but did not name, a nymph in the “Hermanella” complex, noting that the wing pads did not totally cover either the abdomen nor the gills, in contradiction to the original description by NEEDHAM & MURPHY. In discussing the Hermanella complex she recognized four types of nymphs, compared their variability, and noted that wings extracted from the wing pads and those of adults were not comparable. EDMUNDS et al. (1976), using data of WL PETERS, who reared nymphs of the Hermanella complex, elevated Hermanellopsis to generic status. EDMUNDS et al. (1976) cited as a generic character the two lateral lobes and digitiform median projection on the apex of the gills (as DEMOULIN had illustrated for Hermanellopsis), but they gave illustrations of Needhamella. Until DOMÍNGUEZ & FLOWERS (1989) studied the type of H. thelma, all adults of the Hermanella complex of genera had been placed in Thraulus or Traverella. DOMÍNGUEZ & FLOWERS (1989) analyzed the adult characters of Hermanella, and studied reared material of Traverella ehrhardti for which they established the genus Needhamella and at the same time established two subgenera in Hermanella. Based on egg chorion characters of the Hermanella-Traverella complex, DOMÍNGUEZ & CUEZZO (2002) concluded that the subgenera of Hermanella were paraphyletic. We follow this conclusion and do not recognize subgenera within Hermanella. Adult Characters: 1) Fork of veins MA and MP in fore wings asymmetrical, with slanted cross vein above fork of vein MA (Figs. 144A, D, G); 2) costal projection of hind wings (Figs. 144B-C, E-F, H-I) acute; 3) vein Sc of hind wings ending in cross vein near base of costal projection; 4) tarsal claws of a pair dissimilar, one apically hooked, other obtuse (as in Fig. 151E); 5) penes (Figs. 144J-M) divided in apical 1/2, with ventral projections variable, laterally broader in basal 1/3 of each lobe; 6) styliger plate with a pair

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MP MA

A

D Sc

CP

B

E

C

F Pp

sCV

MP MA

SPp

J G Sc

CP

H

I Pp

SPp

M

K

L

R

N

O

P

Q

Plate 144. Hermanella, adults. Figs. 144A-P. Figs. 144A-C, H. maculipennis: 144A, fore wing; 144B, hind wing; 144C, hind wing, detail. 144D-F, H. guttata: 144D, fore wing; 144E, hind wing; 144F, hind wing, detail. 144G-I, H. grandis: 144G, fore wing; 144H, hind wing; 144I, hind wing, detail. 144J-M, male genitalia, v.v.: 144J, H. maculipennis; 144K, H. thelma; 144L, H. guttata; 144M, H. froehlichi. 144NQ, abdominal terga V-VII: 144N, H.maculipennis; 144O, H. guttata; 144P, H. grandis; 144Q, H. froehlichi. 144R, H. maculipennis, prosternum. (sCV = slanted cross-vein; CP = Costal Projection; Pp = Penes projections; SPp = Styliger Plate projection). Figs. 144A-L, N-P and R, modified from DOMÍNGUEZ & FLOWERS, 1989; 144M, Q modified from FERREIRA & DOMÍNGUEZ, 1992.

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of dorsal projections; 7) prosternum with straight median carina (Fig. 144R); and 8) female sternum IX cleft apically (as in Fig. 151H). Nymphal Characteristics (Fig. 145A): 1) Labrum as wide as head with long dorsal setae (Figs. 146A, D), anteromedian emargination as in Figs. 146B, E; 2) outer margin of mandibles angular (Fig. 146F); 3) small tusk on anteromedian margin of maxillae (Fig. 146G); 4) submentum with thick lateral setae (Fig. 146H); 5) apical denticle of tarsal claws much larger than the other denticles (Fig. 146J); 6) posterolateral projections on abdominal segments VIII-IX; and 7) plate-like abdominal gills with two lateral lobes and a digitiform apical projection on segments I-VII (Fig. 146K). Distribution: Argentina, Brazil, French Guiana, Uruguay. Biology: The nymphs of this genus were collected under rocks in small to medium sized streams and rivers, with sandy or rocky bottoms. They were not very abundant. The adults of one species were observed swarming at twilight. Adults of other species (mostly females) were attracted to light. In many cases, the females arrived at the light with a sphere of eggs extruded from the abdomen. KEYS

TO

HERMANELLA

Male Imagos 1 – 2(1) – 3(1) – 4(3) –

Styliger plate (Figs. 144KL) with broad projections at base of penes; penes with broad, lobe-like ventral projections ................................................................................ 2 Styliger plate (Figs. 144J, M) with slender projections at base of penes; penes with curved, spine-like ventral projections ............................................................................ 3 Wings without maculae around cross veins; abdomen blackish brown ................... .................................................................................................................... Hermanella thelma Wings with maculae around cross veins (Figs. 144D-F); abdomen orange-brown (Fig. 144N) .............................................................................................. Hermanella guttata Wing membrane hyaline, costal and subcostal area orange-brown (Fig. 144G) ...... 4 Wing membrane with brown maculae around cross veins (Figs. 144A-C) ............. .......................................................................................................... Hermanella maculipennis Length of fore wings 9.5 to 10.0 mm; color pattern of abdomen as in Fig. 144P ................................................................................................................... Hermanella grandis Length of fore wings 6.8 to 8.7 mm; color pattern of abdomen as in Fig. 144Q ................................................................................................................ Hermanella froehlichi

Nymphs 1 – 2(1) –

Outer margins of middle and hind tibiae with numerous long setae ....................... ................................................................................................................ Hermanella froehlichi Outer margins of middle and hind tibiae glabrous or with small scattered setae (Figs. 146C, I) ...................................................................................................................... 2 Tibiae with relatively short dorsal setae and outer margins glabrous (Fig. 146C); body length > 9 mm ......................................................................................................... 4 Tibiae with longer dorsal setae and outer margin with scattered setae (Fig. 146I); body length < 8 mm ......................................................................................................... 3

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4(2) –

Abdominal terga almost completely blackish, with no distinct pattern ................... .................................................................................................................... Hermanella thelma Abdominal terga orange-brown, with pattern as in male imago (Fig. 144O) ......... ................................................................................................................... Hermanella guttata Wing pads showing traces of spotted imaginal wings ......... Hermanella maculipennis Wing pads without traces of spots, outer margins dark ............... Hermanella grandis

CLAVES

PARA

–

HERMANELLA

Imagos Machos 1 – 2(1) – 3(1) – 4(3) –

Placa estilígera (Figs. 144K-L) con proyecciones ventrales anchas en la base de los penes; penes con proyecciones ventrales anchas, en forma de lóbulos ................. 2 Placa estilígera (Figs. 144J, M) con proyecciones ventrales delgadas, como espinas, en la base de los penes; penes con proyecciones ventrales curvadas, como espinas ................................................................................................................................................ 3 Alas sin manchas alrededor de las venas transversas; coloración general del abdomen castaño-negruzca .................................................................. Hermanella thelma Alas con manchas alrededor de las venas transversas (Figs. 144D-F); coloración general del abdomen castaño-anaranjada (Fig. 144N) .................. Hermanella guttata Alas con la membrana hialina, areas costal y subcostal castaño-anaranjada (Fig. 144G) .................................................................................................................................... 4 Alas con manchas castañas alrededor de las venas transversas (Figs. 144A-C) ...... .......................................................................................................... Hermanella maculipennis Longitud del ala I 9,5 a 10,0 mm; patrón de coloración del abdomen como en la Fig. 144P ................................................................................................. Hermanella grandis Longitud del ala I 6,8 a 8,7 mm; patrón de coloración del abdomen como en la Fig. 144Q ............................................................................................. Hermanella froehlichi

Ninfas 1 – 2(1)

3(2) – 4(2) –

Márgenes externos de tibias II y III con numerosas setas largas ........................... ................................................................................................................ Hermanella froehlichi Márgenes externos de tibias II y III glabros o con pocas setas dispersas (Figs. 146C, I) ................................................................................................................................. 2 Tibias con setas dorsales relativamente cortas y margenes externos glabros (Fig. 146C); largo del cuerpo > 9 mm .................................................................................... 4 Tibias con setas dorsales más largas y margenes externos con setas dispersas (Fig. 146I); largo del cuerpo < 8 mm ..................................................................................... 3 Tergos abdominales casi completemente negruzcos, sin patrón aparente .............. .................................................................................................................... Hermanella thelma Tergos abdominales castaño-anaranjados, con patrón como en el imago macho (Fig. 144O) .............................................................................................. Hermanella guttata Pterotecas con indicios de las alas imaginales manchadas ....... Hermanella maculipennis Pterotecas sin indicios de manchas, margenes externos oscuros ..... Hermanella grandis

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A

Plate 145. Hermanella froehlichi. Fig. 145A, nymphal habitus. Modified from DOMÍNGUEZ et al., 1994.

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Hermanella costalis (NAVÁS) Thraulus costalis NAVÁS, 1934a: 26; TRAVER, 1960b: 73; PETERSEN & GAEDIKE, 1968: 960. Hermanella costalis; PETERS et al., 2005: 55.

Taxonomy: This species was described from Brazil, based on a male imago in poor condition. The specimen had costal coloration on fore and hind wings. TRAVER (1960) stated that it was not Thraulus, but did not reassign the species to other genus. PETERS et al. (2005), observed that the specimen belonged to the Hermanella complex. Nevertheless, the unusual condition of the styliger plate (blunt median protuberance and no paired dorsal projections) do not allow definitive placement of this species to any described genus in the complex and it is excluded from the key. Distribution: Brazil (Rio de Janeiro). Hermanella froehlichi FERREIRA & DOMÍNGUEZ Hermanella (Guayakia) froehlichi FERREIRA & DOMÍNGUEZ, 1992: 179 (male, female, nymph). Hermanella froehlichi; DOMÍNGUEZ & CUEZZO, 2002: 159.

Taxonomy: Hermanella froehlichi can be separated from the other species of the genus by the following combination of characters. In the imagos: 1) Fore wings with costal and subcostal areas tinged with orange-brown; 2) abdominal color pattern as in Fig. 144Q; 3) fore femora with black spot on basal 1/3; and 4) ventral projections on penes narrow and strongly curved (Fig. 144M). In the nymph (Fig. 145A): 1) Abdominal gills blackish with base and apex whitish; 2) outer margin of tibiae of middle and hind legs with numerous long setae. Distribution and Biology: Southern Brazil. The nymphs were collected on and under medium-sized rocks. The imagos and subimagos were attracted to a light trap at dusk. Hermanella grandis DOMÍNGUEZ & FLOWERS Hermanella (Guayakia) grandis DOMÍNGUEZ & FLOWERS, 1989: 565 (female, nymph); HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 212. Hermanella grandis; DOMÍNGUEZ & CUEZZO, 2002: 160.

Taxonomy: Hermanella grandis, known from female imagos and nymphs, can be separated from the other species of the genus by the following combination of characters. In the female imago: 1) Wings 12.0-12.5 mm in length, without macualae (Figs. 144G-I); 2) abdominal color patterns as in Fig. 144P; and 3) black spots present on the basal 1/2 and apex of femora. In the nymph: 1) Abdominal gills blackish; 2) dorsal spines on fore tibiae short (as in Fig. 146C); 3) black spots on basal 1/2 of femora; and 4) wing pads with dark outer margins but without maculae. Distribution and Biology: Northwestern Argentina and Southern Brazil. The females were attracted to light in December. Nymphs of this species were found under rocks in small to medium sized streams.

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Hermanella guttata DOMÍNGUEZ & FLOWERS Hermanella (Hermanella) guttata DOMÍNGUEZ & FLOWERS, 1989: 561 (male, female, nymph); HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 212. Hermanella guttata; DOMÍNGUEZ & CUEZZO, 2002: 160.

Taxonomy: Hermanella guttata can be separated from the other species of the genus by the following combination of characters. In the imago: 1) Wings with brown maculae (Figs. 144D-F; 2) legs without spots on base of femora; 3) abdominal color pattern as in Fig. 144O; and 4) ventral projections of penes broad (Fig. 144L). In the nymph: 1) Legs B

E

F A

D

I G

C

H

K J

Plate 146. Hermanella, nymphs. Figs. 146A-K. Figs. 146A-C, H. maculipennis: 146A, labrum, d.v.; 146B, detail of anteromedian emargination; 146C, foreleg. 146D-K, H. thelma: 146D, labrum, d.v.; 146E, detail of anteromedian emargination; 146F, left mandible; 146G, maxilla; 146H, labium, right ventral, left dorsal; 146I, foreleg; 146J, tarsal claw; 146K, gill I. Figs. 146A-K, modified from DOMÍNGUEZ & FLOWERS, 1989.

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without spots on femora; 2) abdominal color pattern as in male imago (Fig. 144O); and 3) abdominal gills whitish with median area washed with grayish black. Distribution and Biology: Southeastern Argentina, Uruguay, Paraguay and Southern Brazil. The nymphs were collected under rocks in medium sized streams. Hermanella maculipennis (ULMER) Thraulus maculipennis ULMER, 1920a: 30 (male); NEEDHAM & MURPHY, 1924: 44; ULMER, 1943: 28. Traverella maculipennis; EDMUNDS, 1950b: 551; ALLEN, 1973a: 1287; HUBBARD, 1982a: 267. Hermanella (Guayakia) maculipennis; DOMÍNGUEZ & FLOWERS, 1989: 563 (male, female, nymph); HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 212. Hermanella maculipennis; DOMÍNGUEZ & CUEZZO, 2002: 160.

Taxonomy: Hermanella maculipennis was described with H. ehrhardti in the genus Thraulus by ULMER (1920) and later transferred to Traverella by EDMUNDS (1950). DOMÍNGUEZ & FLOWERS (1989) finally transferred this species to Hermanella in their revision of the genus. This species can be separated from the other species of the genus by the following combination of characters. In the imago: 1) Wings with brown maculae (Figs. 144AC); 2) legs with black spot on basal 1/3 of femora; 3) abdominal color pattern as in Fig. 144N; and 4) ventral projections of penes narrow and curved (Fig. 144J). In the nymph: 1)Abdominal gills blackish with grayish apex; 2) dorsal spines on fore tibiae short (Fig. 146C); 3) femora with prominent spot on basal 1/3; and 4) wing pads with spots. Distribution and Biology: Southern Brazil, Northeastern Argentina. The adults were attracted to light immediately after dark from November to April. Adults have been observed flying above the river in large swarms during twilight. Some nymphs were collected in small streams with sandy bottoms. Hermanella thelma NEEDHAM & MURPHY Hermanella thelma NEEDHAM & MURPHY, 1924: 40 (nymph); LESTAGE, 1931b: 57 (nymph); TRAVER, 1947a: 159 [Fig. 129 of NEEDHAM & MURPHY prob. not this genus]; DEMOULIN, 1955a: 7; EDMUNDS, JENSEN & BERNER, 1976: 222; HUBBARD, 1982a: 264; DOMÍNGUEZ & CUEZZO, 2002: 160. Hermanella (Hermanella) thelma; DOMÍNGUEZ & FLOWERS, 1989: 561 (male, nymph); HUBBARD, DOMÍNGUEZ & PESCADOR, 1990: 213. Hermanella velma; SPIETH, 1943: 8.

Taxonomy: Hermanella thelma was originally described from nymphs. DOMÍNGUEZ & FLOWERS (1989), based on new material from the type-locality, redescribed the nymph and described for the first time the male subimago. H. thelma can be differentiated from the other species of this genus by the following combination of characters. In the subimagos: 1) Abdominal terga almost completely blackish, with no apparent pattern; and 2) genitalia as in Fig. 144K, penes with broad ventral projections. In the nymph: 1) Legs without spots; 2) dorsal spines on fore tibiae relatively long (Fig. 146I); 3) abdominal terga blackish, with no apparent pattern; and 4) gills grayish at base and whitish on apical projections. Distribution and Biology: Argentina (Misiones).The nymphs have been collected under rocks in marginal zones of the central river bed of the river Iguazú, running below

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the cataracts. Although numerous females which could belong to this species have been collected at light, male imagos have never been collected. Genus Hermanellopsis DEMOULIN Hermanella [partim] SPIETH, 1943: 9; TRAVER, 1947a: 159. Hermanella (Hermanellopsis) DEMOULIN, 1955a: 8. Hermanellopsis; EDMUNDS, JENSEN & BERNER, 1976: 222; SAVAGE & PETERS, 1983: 576. (Type-species: Hermanella incertans SPIETH, monotypy).

Taxonomy: The type species of Hermanellopsis was described by SPIETH (1943) from three male imagos in the genus Hermanella, a genus known only from nymphs. TRAVER (1947) pointed out some differences between the wings of H. incertans and H. thelma, and, based on her commentary, DEMOULIN (1955a) established a new subgenus for SPIETH’s species. EDMUNDS et al. (1976) raised the subgenus Hermanellopsis to generic level and provided the generic characters for the supposed nymph of this genus. SAVAGE & PETERS (1983) revised the genus, as well as other related genera, and placed in doubt the identity of the proposed nymph; the true nymph of this genus remains unknown. They also described a second species of the genus. Adult Characteristics: 1) Fork of vein MA of fore wings asymmetrical (Figs. 147A, D); 2) vein MP2 joined at base to MP1 and CuA by a cross vein (Fig. 147A) or fork of MP asymmetrical (Fig. 147D); 3) hind wing narrow and acutely pointed (Figs. 147B-C); 4) costal projection of hind wings acute, well developed, apex of projection 2/5 to less than1/2 distance from base (Figs. 147B-C, E); 5) claws of a pair dissimilar, one acute, apically hooked, and other blunt (Fig. 147F), similar in structure on all legs; 6) penes divided, tubular, long and slender, without appendages (Figs. 147G-H); and 7) abdominal sternum IX of female cleft apically (Fig. 147I). Nymphal Characteristics: Unknown. Distribution: Surinam, Guyana, French Guiana and northern Brazil. Biology: The adults were collected in April and June-July. KEY

HERMANELLOPSIS

TO

Imagos 1 –

CLAVE

Vein MP2 of fore wings joined at base to MP1 and CuA by a cross vein (Fig. 147A); penes as in Fig. 147G ..................................................... Hermanellopsis incertans Vein MP2 of fore wings joined to MP1 forming an asymmetrical fork (Fig. 147D); penes as in Fig. 147H ......................................................................... Hermanellopsis arsia PARA

HERMANELLOPSIS

Imagos 1 –

Vena MP2 de las alas anteriores unidas en la base a MP1 y CuA por un vena transversa (Fig. 147A); penes como en la Fig. 147G ............. Hermanellopsis incertans Vena MP2 de las alas anteriores unidas a MP1 formando una horquilla asimétrica (Fig. 147D); penes como en la Fig. 147H ...................................... Hermanellopsis arsia

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MA

399

A G

CP

CuA MP2 MP1

B

C

D

H E

F

I

Plate 147. Hermanellopsis, adults. Figs. 147A-C, H. incertans male imago: 147A, fore wing; 147B, hind wing; 147C, hind wing, detail. 147D-F, H. arsia male subimago: 147D, fore wing; 147E, hind wing; 147F, fore tarsal claw. 147G-H. Male genitalia, v.v., 147G, H. incertans, male imago; 147H, H. arsia, male subimago. 147I, female sternum IX. (CP = Costal Projection). Figs. 147A-I modified from SAVAGE & PETERS, 1983.

Hermanellopsis arsia SAVAGE & PETERS Hermanellopsis arsia SAVAGE & PETERS, 1983: 580 (male).

Taxonomy: Hermanellopsis arsia was described from male subimagos, therefore some of the characters included in the description are not comparable with the other species of the genus, which is known only from imagos. In the adults: 1) Upper portion of male eyes with 30-40 facets in longest row; and 2) vein MP2 of fore wings appears to be asymmetrically joined at base to MP1 (Fig. 147D). Distribution and Biology: Northern Brazil. From the dates of collection, the adults fly in April.

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Hermanellopsis incertans (SPIETH) Hermanella incertans SPIETH, 1943: 9 (male); TRAVER, 1947a: 159 (male, female); DEMOULIN, 1966a: 11. Hermanella (Hermanellopsis) incertans; DEMOULIN, 1955a: 8. Hermanellopsis incertans; EDMUNDS, JENSEN & BERNER, 1976: 223 (male, female, nymph); HUBBARD, 1982a: 264; SAVAGE, 1983b: 128; SAVAGE & PETERS, 1983: 579 (male, female).

Taxonomy: SPIETH (1943) described this species from three male imagos, and included as paratypes two female subimagos, which he did not include in the description. SAVAGE & PETERS (1983) described these females. The imagos of this species can be distinguished from the other species of the genus by the following combination of characters: 1) Upper portion of male eyes with 25-31 facets in longest row; and 2) vein MP2 of fore wings joined at base to veins MP1 and CuA by cross veins (Fig. 147A). Distribution and Biology: Guyana and Surinam. According to the dates of collection, the adults of this species emerge in June and July. Genus Homothraulus DEMOULIN Homothraulus DEMOULIN, 1955a: 11; TRAVER, 1960b: 74; EDMUNDS, JENSEN & BERNER, 1976: 224, [non] Fig. 375 (see discussion under Farrodes); SAVAGE, 1987b: 207; DOMÍNGUEZ et al., 1994: 48; DOMÍNGUEZ et al., 1997: 141. (Type-species: Thraulus misionensis ESBEN-PETERSEN, original designation).

Taxonomy: The genus Homothraulus was established by DEMOULIN (1955a) to include the American species formerly referred to Thraulus which EDMUNDS (1948) had not transferred to Traverella. Thraulus misionensis ESBEN-PETERSEN, from Uruguay and Argentina, was designated as type-species. DEMOULIN (1955a) also described a nymph as Homothraulus sp., on which he principally based this genus. TRAVER (1960) pointed out various errors which were acknowledged by DEMOULIN (in TRAVER 1960b) among which were the fact that the nymph described by DEMOULIN (1955a) did not belong to this genus but was very close to Ulmeritus and that not all the American species of Thraulus must belong to Traverella or Homothraulus. TRAVER (1960b) described the presumed nymph of Homothraulus and a new species, H. lucretiae from Uruguay. In addition to these species, the genus includes tentatively H. larensis (NAVÁS) from Argentina (HUBBARD et al., 1990). This genus is closely related to Simothraulopsis and Farrodes. In this group, Simothraulopsis is restricted to the Guyana Shield and Amazonian basin, while Homothraulus is found only in Southern Brasil and the Paraná Basin. Of the species described from this genus, only H. misionensis has been described from nymphs and adults; H. lucretiae from male and female subimagos and H. larensis only from male imagos. Adult Characteristics: 1) Eyes of male separated dorsally by a distance approximately equal to width of median ocellus; 2) fork of vein MA of fore wings asymmetrical (Fig. 148A); 3) fork of vein MP of fore wings symmetrical (Fig. 148A); 4) vein MP of hind wings (Figs. 148B-C) unforked; 5) costal projection of hind wings acute and located near middle of anterior border; 6) penes of male (Fig. 148D) divided in apical half with apex rounded, each penis lobe with a ventral spine-like projection (in

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subimagos, spine more apically directed as in Fig. 148E); 7) apex of female sternum IX blunt or truncated (Fig. 148F); and 8) tarsal claws of a pair dissimilar, one hooked and the other blunt (Fig. 148G). Nymphal Characteristics (general aspect as in Fig. 149A): 1) Labrum a little broader than clypeus, lateral margins rounded (Fig. 149B); 2) mandible with rounded lateral margin and with a small tuft of setae near middle (Fig. 149C); 3) apical denticle of claws much larger than others (Fig. 149D); 4) abdominal gills on segment I-VII lanceolate, dorsal and ventral portions similar (Fig. 149E); and 5) posterolateral spines on abdominal segments VIII-IX. Distribution: Argentina, Uruguay, Brazil. Biology: The nymphs have been collected from small to medium-sized rivers, generally in areas with swifter current. The nymphs were found mainly in the rocky bottoms, but some were in debris and vegetation. Adults emerge between December and March. Imagos were mainly captured with aerial nets, but the subimagos were attracted by light. MP

MA

A

CP

B

F

G

C MP

D

Pp

E

H

I

Plate 148. Homothraulus, adults. Figs. 148A-D, H. misionensis: 148A, fore wing; 148B, hind wing; 148C, hind wing, detail; 148D, genitalia, v.v. 148E, H. lucretiae, genitalia, v.v. 148F-148H, H. misionensis: 148F, apex of female sternum IX; 148G, fore claw; 148H, abdominal terga II-VII. 148I, H. lucretiae, abdominal terga II-VII. (CP = Costal Projection; Pp = Penes projections).

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HOMOTHRAULUS

TO

Imagos 1 –

CLAVE

Longitudinal veins whitish yellow; femora yellowish brown with a darker black macula on the dorsal side .............................................................. Homothraulus lucretiae Longitudinal veins yellowish brown; femora whitish yellow with a blackish longitudinal line and a brown incomplete dorsal ring near the apex ........................ ......................................................................................................... Homothraulus misionensis PARA

HOMOTHRAULUS

Imagos 1 Venas longitudinales amarillo-blanquecinas; fémures castaño-amarillentos con una mancha más oscura en el lado dorsal ................................. Homothraulus lucretiae – Venas longitudinales castaño-amarillentas; fémures amarillo-blanquecinos con una linea longitudinal negruzca y un anillo castaño, incompleto dorsalmente, cerca del ápice ........................................................................................ Homothraulus misionensis Homothraulus larensis (NAVÁS) Thraulus larensis NAVÁS, 1926b: 33 (male); NAVÁS, 1930e: 130; ULMER, 1943: 28; PETERSEN & GAEDIKE, 1968: 961; HUBBARD, 1982a: 267; ALBA-TERCEDOR & PETERS, 1985: 223; DOMÍNGUEZ, 1989b: 273. Thraulus lorensis [sic]; BERTHÉLEMY, 1965: 2. Homothraulus larensis; HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 213; DOMÍNGUEZ et al. 1994: 49.

Taxonomy: This species was placed in Homothraulus by HUBBARD et al., 1992 based on examination of syntypes by P. GRANT (Ph.D. dissertation, in Ms.,1985: 217). Due to limited data, it is presently impossible to characterize this species. Types are located at the Museo del Ayuntamiento de Zaragoza, Barcelona, España (ALBA-TERCEDOR & PETERS, 1985); at the Deutches Entomologisches Institut, Berlin, Germany (PETERSEN & GAEDIKE, 1968) and at the Museo Argentino de Ciencias Naturales, Buenos Aires, Argentina (DOMÍNGUEZ, 1989b). Distribution: Argentina (Buenos Aires). Homothraulus lucretiae TRAVER Homothraulus lucretiae TRAVER, 1960b: 81; HUBBARD, 1982a: 265.

Taxonomy: TRAVER (1960b) justified the desgination of a new species based only on subimagos because of its characteristic color pattern. One of us (ED) had the opportunity to study the material determined by TRAVER (1960b) as belonging to this species and now we consider it a valid species. Homothraulus lucretiae can be differentiated from the other species of the genus by the following combination of characters. In the subimago: 1) Abdominal terga I-III and VI-X grayish brown, IV-V yellowish, with 2 submedian blackish lines on VI-X (Fig. 148I); 2) dark brown macula on middorsal portion of femora; and 3) longitudinal veins of fore wings whitish yellow. Distribution: Uruguay.

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Homothraulus misionensis (ESBEN-PETERSEN) Thraulus misionensis ESBEN-PETERSEN, 1912: 339 (male); ULMER, 1920c: 117; ULMER, 1921: 263; NEEDHAM & MURPHY, 1924: 44; ULMER, 1943: 29 (male). Homothraulus misionensis; DEMOULIN, 1955a: 11; TRAVER, 1956a: 12 (nymph); TRAVER, 1960b: 78 (male, female, nymph); THEW, 1960b: 122; EDMUNDS, JENSEN & BERNER, 1976: 224; HUBBARD, 1982a: 265; HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 212; DOMÍNGUEZ et al., 1994: 48.

Taxonomy: This is the only species of the genus for which adults and nymphs are known. The imagos of H. misionensis can be distinguished from the other species of the genus by the

B

S

A

C

D

D

E

Plate 149. Homothraulus, nymph. Figs. 149A-E, H. misionensis: 149A, habitus; 149B, clypeus and labrum, d. v.; 149C, left mandible; 149D, tarsal claw; 149E, gill IV. (D = denticle; S = Setae). Fig. 149A, drawing by A. Dupuy.

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following combination of characters: 1) Abdominal terga II-VII translucent with blackish wash over posterior 2/3 of each segment, and VIII-IX almost totally black (Fig. 148H); 2) legs without middorsal brown maculae; and 3) longitudinal veins of fore wings yellowish brown. Distribution and Biology: Argentina, Brazil, and Uruguay. According to the collection data, the nymphs are found in rapids of streams and rivers; adults emerge between December and March. It appears only the subimagos are attracted to light because the few adults present in collection were collected with a net. Genus Hydrosmilodon FLOWERS & DOMÍNGUEZ Hydrosmilodon FLOWERS & DOMÍNGUEZ, 1992: 655; DOMÍNGUEZ & CUEZZO, 2002: 160; ORTH et al., 2000: 30. (Type-species: Thraulus primanus EATON, original designation).

Taxonomy: The genus Hydrosmilodon was established by FLOWERS & DOMÍNGUEZ (1992) for the species Thraulus primanus EATON from Mexico and for a new species from Argentina (H. saltensis). In a cladistic analysis, FLOWERS & DOMÍNGUEZ (1991) found that T. primana was more related to the Hermanella complex (Hermanella NEEDHAM & MURPHY, Hylister DOMÍNGUEZ & FLOWERS and Needhamella DOMÍNGUEZ & FLOWERS) than to the genus Traverella where it was transferred by TRAVER (1960). This was confirmed by a separate analysis, based on the egg chorion morphology of the whole complex (DOMÍNGUEZ & CUEZZO, 2002). Besides these two species, material corresponding to other undescribed South American species was examined, but the authors of the genus did not describe them due to the lack of adults. Most recently, THOMAS et al. (2004) added two new species, H. gilliesae and H. mikei, from nymphs from French Guiana. Adult Characteristics: 1) Forks of veins MA and MP of fore wings asymmetrical (Fig. 150A), with slanted cross vein close to MA fork; 2) vein Sc of hind wings ending in transverse vein near base of costal projection; 3) vein MP of hind wings unforked (Figs. 150B-C); 4) costal projection of hind wings (Figs. 150B-C) acute, rounded at apex; 5) tarsal claws of a pair dissimilar, one apically hooked, other blunt (Fig. 150D); 6) penes (Fig. 150E) divided in apical 1/2, each lobe with short median spine-like projection; basal 1/2 of penes with swollen lateral margins; 7) styliger plate with spines close to base of forceps (Fig. 150E); 8) prosternum with short median carina (Fig. 150F); and 9) female sternum IX apically cleft (Fig. 150G). Nymphal Characteristics: 1) Labrum as wide as head, with long setae (Fig. 150I); 2) outer margin of mandible angular (Fig. 150J); 3) maxillae with large blade-like tusk (tusk length approximately 1/2 width of the apex of maxilla) (Fig. 150K), rows of long setae on maxillary palpi; 4) apical denticle of tarsal claws same size as other denticles or much larger (Figs. 150M, O, Q), small apical denticles sometimes present; 6) posterolateral projections on abdominal segments VII or VIII-IX; and 7) gills (Figs. 150N, P, R) on abdominal segments I-VII, narrow and plate-like, with a single finger-like process, to narrow and tapered apically; gills on VII sometimes vestigial. Distribution: Northwestern Argentina (Salta Prov.), Southern Bolivia and French Guiana. Also Central and Tropical South America. Biology: The nymphs are found in rocky streams, from low to middle elevations. Adults were attracted to light. The adults flight during most of the year in Central America, but apparently this period is much more restricted in the Subtropics.

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TO

405

SOUTH AMERICAN HYDROSMILODON

Nymphs 1

Denticles of tarsal claws subequal in size (Fig. 150Q); posterolateral projections on abdominal segments VII-IX ...................................................... Hydrosmilodon mikei Subapical denticle of tarsal claws much larger than other denticles (Fig. 150M, O); posterolateral projections on abdominal segments VIII-IX, sometimes reduced on VIII ................................................................................................................................. 2 Gills elongate-oval on abdominal segments I-VII, gills with apical filament and tracheal trunk with many branches (Fig. 150N); claws without small apical denticles (Fig. 150M); posterolateral projections on abdominal segments VIII-IX ............................................................................................................ Hydrosmilodon saltensis Gills narrow on abdominal segments I-VII, tapered to terminal filament on segments I-VI, and tracheal trunk without branches (Fig. 150P) ; gills vestigial on segment VII; claws with one larger apical denticle (Fig. 150O); posterolateral projections of abdominal segment reduced, very small on segment VIII .............. ............................................................................................................. Hydrosmilodon gilliesae

– 2(1)

–

CLAVE

PARA LOS

HYDROSMILODON

SUDAMERICANOS

Ninfas 1 – 2(1)

–

Dentículos de las uñas tarsales subiguales en tamaño (Fig. 150Q), proyecciones posterolaterales en segmentos abdominales VII-IX ................... Hydrosmilodon mikei Dentículo subapical de las uñas tarsales mucho mayor que los otros dentículos (Fig. 150M, O); proyecciones posterolaterales en segmentos abdominales VIII-IX, las del segmento VIII a veces reducidas ....................................................................... 2 Branquias ovales, elongadas, en segmentos abdominales I-VII, branquias con un filamento apical y traquea con muchas ramificaciones (Fig. 150N); uñas sin dentículos apicales pequeños (Fig. 150M); proyecciones posterolaterales en segmentos abdominales VIII-IX ................................................ Hydrosmilodon saltensis Branquias delgadas en segmentos abdominales I-VII, afinadas hasta un filamento terminal en segmentos I-VI, y traquea sin ramificaciones (Fig. 150P); branquias vestigiales en segmento VII; uñas con el dentículo apical más grande (Fig. 150O); proyecciones posterolaterales de los segmentos abdominales reducidas, muy pequeñas en el segmento VIII ..................................................... Hydrosmilodon gilliesae

Hydrosmilodon gilliesae THOMAS & PÉRU Hydrosmilodon gilliesae THOMAS & PÉRU in THOMAS et al., 2004: 66 (nymph).

Taxonomy: H. gilliesae can be distinguished from the other species of the genus by the following combination of characters in the nymph: 1) Outer margin of mandibles without setae; 2) clypeus without median projection; 3) lateral margins of pronotum with 2-3 long, sharp setae; 4) tarsal claws with large subapical denticle and a smaller apical denticle (Fig. 150O); 5) gills slender, tapered to apical filament, and without tracheal

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MP

MA

sCV Pp

SPp

A Sc

B

CP

C

E

F V

VI

D

G

VII

H

I

J N D

K L

M

O

P

Q

R

Plate 150. Hydrosmilodon. Figs. 150A-N: H. saltensis. 150A-H, male imago: 150A, fore wing; 150B, hind wing; 150C, hind wing, detail; 150D, fore tarsal claw; 150E, genitalia, v.v.; 150F, prosternum; 150G, female sternum IX; 150H, abdominal terga V-VII. 150I-N, nymph. 150I, labrum, d.v.; 150J, left mandible; 150K, maxilla; 150L, labium, right ventral, left dorsal; 150M, fore tarsal claw; 150N, gill IV. H. gilliesae: 150O-P. 150O, claw III; 150P, gill V. H. mikei: 150Q, claw III; 150R, gill V. (sCV = slanted cross-vein; CP = Costal Projection; Pp = Penes projections; SPp = Styliger Plate projection; D = denticle; Roman numbers = segment numbers). Figs. 150A-N modified from FLOWERS & DOMÍNGUEZ, 1992; 150O-R modified from THOMAS et al., 2004.

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branches (Fig. 150P) on abdominal terga I-VI , vestigial on VII; and 6) posterolateral projections on abdominal terga VIII-IX, that on VIII very small. Distribution: French Guiana. Hydrosmilodon mikei THOMAS & BOUTONNET Hydrosmilodon mikei THOMAS & BOUTONNET in THOMAS et al., 2004: 72 (nymph).

Taxonomy: This unique species is tentatively placed in this genus until the adult will be described. It can be distinguished from the other known species by the following combination of character in the nymph: 1) Outer margin of mandibles without setae; 2) clypeus without median projection; 3) lateral margins of pronotum with 2 short setae; 4) denticles on tarsal claws subequal in size, median denticles a little larger, and with a row of small apical denticles (Fig. 150Q); 5) gills narrowly oval without tracheal branches and with median apical filament on abdominal terga I-VI (Fig. 150R), vestigial on VII; and 6) posterolateral projections on abdominal segment VII-IX. Distribution: French Guiana. Hydrosmilodon saltensis FLOWERS & DOMÍNGUEZ Hydrosmilodon saltensis FLOWERS & DOMÍNGUEZ, 1992: 659 (male, female, nymph).

Taxonomy: H. saltensis, can be distinguished from the other species of the genus by the following combination of characters: In the imago: 1) Styliger plate with short and straight spines close to base of forceps (Fig. 150E); 2) general coloration blackish-brown, abdominal color pattern as in Fig. 150H; 3) forceps orange brown. In the nymph: 1) Outer margin of mandibles with basal setae (Fig. 150J); 2) clypeus with small median projection; 3) lateral margins of pronotum with 3 long, pointed setae; 4) tarsal claws with large subapical denticle and without small apical denticles (Fig. 150M); 5) gills elongate-oval with tracheal branches and with distinct apical filament on both portions on abdominal segments I-VII (Fig. 150N); and 6) posterolateral projections on abdominal segments VIII-IX. Distribution and Biology: Northwestern Argentina (Salta Prov.) and Southern Bolivia. The nymphs were collected in mountain rain forest streams, 10-17 m wide and 3570 cm deep, with exposed bedrock bottoms. Abundant leaves and debris were deposited between the rocks. The nymphs were found under stones in water 40 - 50 cm deep. This species appears to be highly seasonal, occurring in abundance during February, but are rare or absent at other times of the year. Subimagos emerge at dusk. Imagos were attracted to light traps. Genus Hylister DOMÍNGUEZ & FLOWERS Hylister DOMÍNGUEZ & FLOWERS, 1989: 565; FLOWERS & DOMÍNGUEZ, 1991: 51; DOMÍNGUEZ, FERREIRA & NIETO, 2001: 316; DOMÍNGUEZ & CUEZZO, 2002: 158. (Type-species: Hylister plaumanni DOMÍNGUEZ & FLOWERS, original designation).

Taxonomy: Hylister was established based on male and female imagos and nymphs. According to FLOWERS & DOMÍNGUEZ (1991) and DOMÍNGUEZ et al. (2001), it belongs in the

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MA

A B D

Sc

CP

F

C

E

H

V

G

VI

VII

I

J III

D

II

L

I

M

K

Plate 151. Hylister plaumanni. Figs. 151A-M. 151A-G, male imago: 151A, fore wing; 151B, hind wing; 151C, hind wing, detail; 151D, genitalia, v.v.; 151E, fore claw; 151F, prosternum; 151G, abominal terga V-VII. 151H, apex of female sternum IX. 151I-M, nymph: 151I, labrum, d. v.; 151J, left mandible; 151K, labium, right ventral, left dorsal; 151L, fore claw; 151M, gill I. (CP = Costal Projection; D = denticle; Roman numbers = segment numbers). Figs. 151A-M modified from DOMÍNGUEZ & FLOWERS, 1989.

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Hermanella complex and is a sister group of the genus Hermanella. This was confirmed by a separate study, based on egg chorion morphology of the group (DOMÍNGUEZ & CUEZZO, 2002). Adult Characteristics: 1) Forks of veins MA and MP of fore wings asymmetrical (Fig. 151A), with slanted cross vein above fork of vein MA; 2) costal projection of hind wings (Figs. 151B-C) sharply pointed; 3) vein Sc of hind wings ending in transverse vein near base of costal projection (Figs. 151B-151C); 4) tarsal claws dissimilar, one apically pointed, other blunt (Fig. 151E); 5) penes (Fig. 151D) divided in apical 1/4, ventral projections of penes distal to ventral groove, with lateral swellings on basal half of each penis lobe; 6) styliger plate without ventral projections; 7) prosternum with straight median carina (Fig. 151F); and 8) female sternum IX apically cleft (Fig. 151H). Nymphal Characteristics: 1) Labrum as wide as head, with setae as in Fig. 151I; 2) outer margin of mandible (Fig. 151J) angular, with angle rounded; 3) submentum with thick lateral setae and short ventral setae (Fig. 151K); 4) apical denticle of tarsal claws much larger than other denticles (Fig. 151L); 5) posterolateral projections on abdominal segments VII-IX; and 6) abdominal gills on segments I-VI (Fig. 151M) plate-like, apically truncated, with 8-10 finger-like projections. Distribution: Southeastern Brazil. Biology: The nymphs were found under rocks of medium to large size in a totally shaded mountain stream. The adults were attracted to light near dusk in February. Hylister plaumanni DOMÍNGUEZ & FLOWERS Hylister plaumanni DOMÍNGUEZ & FLOWERS, 1989: 567 (male, female, nymph, egg); DOMÍNGUEZ & CUEZZO, 2002: 158.

Taxonomy: H. plaumanni, the only species known in the genus, can be characterized by the following characters. In the imago: 1) Membrane of fore wings (Fig. 151A) hyaline, costal and subcostal areas translucent brown, darker near wing base; 2) genitalia with spines strongly curved as in Fig. 151D; and 3) abdominal color pattern as in Fig. 151G. In the nymph: 1) Gills grayish, with whitish projections (Fig. 151M); and 2) abdominal terga and sterna brownish orange, and 3) color pattern as in male imago (Fig. 151G). Distribution and Biology: As in generic discussion. Genus Leentvaaria DEMOULIN Leentvaaria DEMOULIN, 1966A: 13; DOMÍNGUEZ, FERREIRA & NIETO, 2001: 313; ORTH et al., 2000: 31. (Type-species: Leentvaaria palpalis DEMOULIN, original designation).

Taxonomy: Leentvaaria was established by DEMOULIN (1966a) for a single species, known only from nymphs. In 2001, DOMÍNGUEZ et al. redescribed the nymphs, based on new material, and studied the phylogenetic relationships of this genus. Adult Characteristics: Unknown. Nymphal Characteristics: 1) Labrum as wide as head, with shape and dorsal setae as in Fig. 152A; 2) outer margin of mandible angular (Fig. 152B); 3) long setae on maxillary palpi in even rows, prominent tusk on inner apical margin of maxillae (Fig. 152C); 4) labial palpi very long, segment I with basal prominence; segment II with long

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row of dorsal setae (Fig. 152D); 5) subapical denticle of tarsal claw very enlarged, also with a few small apical denticles (Fig. 152E); 6) gills on abdominal segments I-VII lanceolate, narrow, tapering gradually toward apex, smaller posteriorly (Fig. 152F); 7) posterolateral spines on abdominal segments VIII-IX. Distribution: Brazil, French Guiana and Surinam. Biology: Unknown.

B

A

III T

D II

C

E

I

D

F

Plate 152. Leentvaaria, nymph. Figs. 152A-F, Leentvaaria sp.: 152A, labrum, d.v.; 152B, left mandible; 152C, maxilla; 152D, labium, right ventral, left dorsal; 152E, tarsal claw; 152F, gill IV. (D = denticle; T = tusk; Roman numbers = segment numbers). Figs. 152A-F, modified from DOMÍNGUEZ et al., 2001.

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Leentvaaria palpalis DEMOULIN Leentvaaria palpalis DEMOULIN, 1966a: 14 (nymph); HUBBARD, 1982a: 265; DOMÍNGUEZ, FERREIRA & NIETO, 2001: 314.

Taxonomy: Leentvaaria palpalis is the only species known in this genus. This species was based on three young nymphs. For this reason, DOMÍNGUEZ et al. (2001), were not able to decide if the specimens they had were of the same species. According to the original description, L. palpalis can be characterized by the general brown coloration, with the abdominal terga darker laterally, and whitish legs and cerci. Distribution: Surinam. Genus Magallanella PESCADOR & PETERS Genus B PESCADOR & PETERS, 1980b: 47. Magallanella PESCADOR & PETERS, 1980c: 336. (Type-species: Magallanella flinti PESCADOR & PETERS, original designation).

Taxonomy: The genus Magallanella was described by PESCADOR & PETERS (1980c) based only on nymphs collected from Magallanes Province, Chile. Adult Characteristics: Unknown. Nymphal Characteristics (general aspect as in Fig. 153A): 1) Clypeus narrower than labrum, and lateral margins of labrum divergent and abruptly curved (Fig. 153B); 2) anteromedian emargination of labrum V-shaped, with weakly developed blunt denticles (Fig. 153C); 3) outer margin of mandibles slightly angular with a sparse median seta tuft (Fig. 153D); 4) galea-lacinia of maxillae with apex narrow, segments I and II of palpi equal in length, segment III a little shorter (Fig. 153E), segment II with tuft of setae; 5) denticles of tarsal claws progressively larger apically except apical denticle much larger (Fig. 153F); 6) posterolateral projections on abdominal segments VI-IX; and 7) abdominal gills I-VII consisting of single plate-like lamella terminating apically in a thick filamentous process (Fig. 153G). Distribution: Southern Chile (Magallanes Prov. ca. 53ºS , XII region). Biology: The nymphs were collected in small streams, 30-90 cm wide by 5-10 cm deep, with gravel and rubble bottoms and with a thick growth of moss. The streams were located in open forest and steppe. The nymphs are detritivores. Magallanella flinti PESCADOR & PETERS Magallanella flinti PESCADOR & PETERS, 1980c: 337 (nymph).

Taxonomy: Magallanella flinti, known only from the nymph, is the only described species in the genus (Fig. 153A). The shape and structure of the abdominal gills (Fig. 153G) and tarsal claws (Fig. 153F) easily differentiate the species from the superficially similar Dactylophlebia carnulenta. Distribution and Biology: Distribution and biology are discussed in the generic description.

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C

B

D

A

D

E F

G

Plate 153. Magallanella flinti, nymph. Figs. 153A-G: 153A, habitus; 153B, clypeus and labrum, d.v.; 153C, anteromedian emargination of labrum, detail; 153D, left mandible; 153E, maxilla; 153F, fore leg claw; 153G, gill IV. (D = denticle). Figs. 153A-G, modified from PESCADOR & PETERS, 1980C.

Genus Massartella LESTAGE Massartella LESTAGE, 1924a: 21; LESTAGE, 1930a: 251; ULMER, 1943: 14; TRAVER, 1946: 422; DEMOULIN, 1955a: 14; PETERS & EDMUNDS, 1972: 1404; PESCADOR & PETERS, 1980b: 47; PESCADOR & PETERS, 1990: 145; DERKA, 2002: 309. (Type-species: Atalophlebia brieni LESTAGE, original designation).

Taxonomy: LESTAGE (1930a) established Massartella for M. brieni which he had earlier described from adults (LESTAGE, 1924a) as Atalophlebia brieni. ULMER (1943)

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described two new species based also on adults. DEMOULIN (1955a) described the nymph. Subsequent taxonomic accounts of the genus were given by PETERS & EDMUNDS (1972) and PESCADOR & PETERS (1990). Adult Characteristics: Massartella can be distinguished from the other genera of the family by the following combination of characters. In the imagos: 1) Fork of vein MA of fore wings symmetrical (Fig. 154A); 2) vein ICu1 of fore wings attached at base to cross vein between veins CuA and CuP, attachment close to CuP; 3) hind wings without costal projection and vein Sc 9/10 length of the wing (Fig. 154B); 4) claws of a pair similar, both apically hooked, each with an opposing hook (Fig. 154F); 5) lobes of penes divided to base, each lobe with 1 or 2 subapical lateral spines (Figs. 154D-E); and 6) female sternum IX deeply cleft apically (Fig. 154G). Nymphal Characteristics: 1) Clypeus as wide as labrum (Fig. 155A); 2) length of labrum approximately 1/2 maximum width (Fig. 155A); 3) mandibles with slightly curved outer margin and a median seta tuft (Figs. 155B-C); 4) hypopharynx as in Fig. 155E; 5) glossae of labium ventrally curved (Fig. 155F) and submentum with long lateral spines (Fig. 155F); 6) denticles on tarsal claws progressively larger apically (Fig. 155I); 7) abdominal terga with short spines laterally, and posterior margin with broad spines and long feather-like setae; 8) posterolateral projections on abdominal segments II-IX; and 9) abdominal gills I-VI similar, plate-like, well developed, inner basal lobe of ventral portion enlarged (Figs. 155J-L), gills absent on abdominal segment VII. Distribution: Argentina, Brazil and Venezuela. Biology: The nymphs of Massartella are found from cold mountain streams and rivers to small lowland forest streams. Nymphs were collected from leaf-packs in slow flowing streams with muddy bottoms, and from rocks in the faster portions of mountain streams. Last instar nymphs climb out of water onto dry rocks to emerge. The swarm is composed of few individuals and takes place in midmorning. The adults fly from mid-September to mid-July. KEYS

TO

MASSARTELLA

Male Imagos 1

–

Wing membrane faintly toned with brownish-yellow, cross veins clouded with dark brown; Cu area of fore wings encloses a network of cross veins (Fig. 154C); dark brown maculae on abdominal terga as in Fig. 154I; penis lobes apically pointed (Fig. 154E) ............................................................................ Massartella alegrettae Wing membrane hyaline except costal and subcostal membranes of fore wings toned with brownish-yellow, cross veins not clouded; Cu area of fore wings encloses little or no network of cross veins (Fig. 154A); maculae on abdominal terga not as above (Fig. 154H); penis lobes apically blunt (Fig. 154D) ................. Massartella brieni

Nymphs 1 –

Posterolateral projections of abdominal segments long, 3/4 to equal maximum median length of segment (Fig. 155N) .................................... Massartella venezuelensis Posterolateral projections of abdominal segments short, less than 1/2 maximum median length of segment (Figs. 155M, O) ................................................................. 2

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C A Sc CuP ICu1 CuA

MP2

B

F

D

E G

H

I

J

Plate 154. Massartella, imagos. Figs. 154A-C: Wings. 154A-B, M. brieni. 154A, fore wing; 154B, hind wing. 154C, M. alegrettae cubital area. 154D, male genitalia, v.v.; 154E, M. alegrettae, penes, v.v. 154F-G, M. brieni: 154F, fore claw; 154G, apex of female sternum IX. 154H-J, abdominal terga IV-VIII: 154H, M. brieni, male imago; 154I, M. alegrettae, male subimago; 154J, M. venezuelensis, female subimago. (MA = fork of MA). Figs. 154A-J modified from PESCADOR & PETERS, 1990.

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Dorsal portion of gills with well developed apical process (Fig. 155J); abdominal sterna with prominent thick setae ........................................................ Massartella brieni Dorsal portion of gills without apical process (Fig. 155L); abdominal sterna glabrous .....................................................................................................Massartella devani PARA

MASSARTELLA

Imagos Machos 1

–

Membrana de las alas teñidas de castaño amarillento, venas transversas sombreadas de castaño obscuro; área Cubital de las alas anteriores con una malla de venas transversas (Fig. 154C); manchas castaño obscuras en tergos abdominales como en Fig. 154I; lóbulos de los penes aguzados apicalmente (Fig. 154E) ...................... ................................................................................................................ Massartella alegrettae Membrana de las alas hialina excepto membranas costal y subcostal de las alas anteriores teñidas de castaño amarillento, venas transversas no teñidas; área Cubital de las alas anteriores con pocas o sin malla de venas transversas (Fig. 154A); manchas en tergos abdominales no como las precedentes (Fig. 154H); lóbulos de los penes obtusos (Fig. 154D) ........................................... Massartella brieni

Ninfas 1. – 2. –

Espinas posterolaterales de los segmentos abdominales largos, de iguales a 3/4 de la longitud media máxima del segmento (Fig. 155N) ............ Massartella venezuelensis Espinas posterolaterales de los segmentos abdominales cortas, menos de 1/2 de la longitud media máxima del segmento (Figs. 155M, O) ............................................. 2 Porción dorsal de las branquias con un proceso apical bien desarrollado (Fig. 155J); esternos abdominales con setas gruesas y prominentes ...... Massartella brieni Porción dorsal de las branquias sin proceso apical (Fig. 155L); esternos abdominales glabros ........................................................................................................Massartella devani

Massartella alegrettae ULMER Massartella alegrettae ULMER, 1943: 21 (male); PETERS & EDMUNDS, 1972: 1404; HUBBARD, 1982a: 265; PESCADOR & PETERS, 1990: 148 (male).

Taxonomy: ULMER (1943) originally described M. alegrettae from a male imago collected from Alegrete, Rio Grande do Sul State, Brazil. The nymph and female imagos of the species are still unknown. PESCADOR & PETERS (1990) redescribed the species based on the holotype and additional subimagos. Massartella alegrettae can be distinguished from M. brieni by the following characters of the male imago: 1) Wing membranes faintly toned with brown and cross veins clouded with dark brown; 2) cubital area of fore wings enclosing a network of cross veins (Fig. 154C); 3) pattern of dark brown maculae on abdominal terga as illustrated in Fig. 154I; and 4) penis lobes apically pointed (Fig. 154E). Distribution: Paraná State and Rio Grande do Sul State, Brazil.

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C

B

D G

H

E

F

J

M

K

L

N

I

O

Plate 155. Massartella, nymphs. Figs. 155A-H: Mouthparts. M. brieni (Figs. 155A-B, D-G); M. venezuelensis (Figs. 155C, H). 155A, labrum, d.v. ; 155B-C, left mandibles; 155D, maxilla; 155E, hypopharynx; 155F, labium (left dorsal, right ventral); 155G-H, detail of segment III of labial palpi, dorsal. 155I, fore claw, M. brieni. 155J-L, gill IV: 155J, M. brieni; 155K, M. venezuelensis; 155L, M. devani. 155M-O, abdominal terga: 155M, M. brieni; 155N, M. venezuelensis; 155O, M. devani. Figs. 155B-K, M-N modified from PESCADOR & PETERS, 1990; 155L, O from DERKA, 2002.

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Massartella brieni (LESTAGE) Atalophlebia brieni LESTAGE, 1924a: 22 (male, female, egg); LESTAGE, 1930b: 439. Massartella brieni; LESTAGE, 1930a: 251 (male, female, egg); ULMER, 1943: 19 (male, female); DEMOULIN, 1955a: 14 (female, nymph); DEMOULIN, 1955f: 59; PETERS & EDMUNDS, 1972: 1404; HUBBARD, 1982a: 265; PESCADOR & PETERS, 1990: 150 (male, female, nymph, egg). Atalophlebia axillata NAVÁS 1934b: 163 (male); PETERSEN & GAEDIKE, 1968: 960. Massartella fruhstorferri ULMER, 1943: 20 (male, female); PETERS & EDMUNDS, 1972: 1404 (female, nymph); HUBBARD, 1982a: 265. Massartella spp. PETERS & EDMUNDS, 1972: 1404 (male, nymph).

Taxonomy: PESCADOR & PETERS (1990) have recently reviewed the taxonomy of the species including a brief discussion on nymphal and adult variations and placed the species as a senior synonym of Massartella fruhstorferri ULMER. The imagos of Massartella brieni can be distinguished from M. alegrettae by the following combination of characters: 1) Cubital area of fore wings lacking a network of cross veins (Fig. 154A); 2) maculae on abdominal terga as in Fig. 154H; and 3) penis lobes slightly blunt apically, each lobe with short subapical projection (Fig. 154D ). The nymphs can be distinguished from M. venezuelensis and M. devani by the following combination of characters: 1) Mandibles with thin median seta tuft (Fig. 155B); 2) segment II of maxillary palpi with pectinate setae; 3) segment III of labial palpi with two rows of short peg-like setae (Fig. 155G); 4) abdominal sterna with thick setae; 5) dorsal portion of gills with well developed apical process (Fig. 155J); 6) posterolateral projections on abdominal segments less than 1/2 maximum median length of segment (Fig. 155M). Distribution and Biology: Brazil. The nymphs were collected from rocks in faster portion of large rivers to small mountain streams, at altitudes between 800 and 1600 m. Collection records indicate that the adults have been found from September to March. The imagos mate in mid-morning in small companies, at approximately 10 m above the river. Massartella devani DERKA Massartella devani DERKA, 2002: 309.

Taxonomy: Massartella devani was described based only on nymphs, that can be distinguished from M. brieni and M. venezuelensis by the following combination of characters: 1) Mandibles with thin median setae tuft (similar to Fig. 155B); 2) segment II of maxillary palpi with spine-like setae; 3) segment III of labial palpi with one row of short peg-like setae (similar to Fig. 155H); 4) abdominal sterna glabrous; 5) dorsal portion of gills without apical process (Fig. 155L); and 6) posterolateral projections of abdominal segments short (1/5-1/6 maximum median length of segment) (Fig. 155O). Distribution and Biology: Venezuela (Bolivar). The nymphs of M. devani were found in a small stream originating in the Mt. Roraima. The stream was approximately 1.5 m wide, and the substratum composed of a mixture of gravel and stones.

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Massartella venezuelensis PESCADOR & PETERS Massartella venezuelensis PESCADOR & PETERS 1990: 155 (female, nymph); DERKA, 2002: 315.

Taxonomy: Massartella venezuelensis is presently known only from the nymph and female subimago. The abdominal color pattern of the female subimago is as in Fig. 154J. The nymph of this species can be distinguished from M. brieni and M. devani by the following combination of characters: 1) Mandibles with thick median seta tuft (Fig. 155C); 2) segment II of maxillary palpi with spine-like setae; 3) segment III of labial palpi with one row of short peg-like setae (Fig. 155H); 4) abdominal sterna glabrous; 5) dorsal portion of gills with weakly developed apical process (Fig. 155K); and 6) posterolateral projections of abdominal segments long, about 3/4 to equal to maximum length of segment (Fig. 155N). Distribution and Biology: Venezuela (Bolivar, Escalera, 108 km. S Río Cuyuni). The nymphs of M. venezuelensis were found in a tumbling, fast flowing cold stream with a mixture of gravel, rubble and bedrock substratum. The stream was approximately 10 m wide and 10-30 cm deep. Massartella sp. PESCADOR & PETERS Massartella sp. PESCADOR & PETERS, 1990: 158.

Taxonomy: PESCADOR & PETERS (1990) mentioned the presence of immature nymphs of this genus in Northeastern Argentina. Later, more nymphs (unpublished) were collected from small streams in the same area. This material was found in streams with muddy or clay bottom, and the nymphs were hiding among leaf packs. Distribution: Northeastern Argentina (Misiones Prov.). Genus Massartellopsis DEMOULIN Massartellopsis DEMOULIN, 1955e: 9; PETERS & EDMUNDS, 1972: 1409; PESCADOR & PETERS, 1980b: 43; PESCADOR & PETERS, 1987: 181; SAVAGE, 1987b: 207; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 51; DOMÍNGUEZ et al. 2001: 29. (Type-species: Massartellopsis irarrazavali DEMOULIN, original designation).

Taxonomy: DEMOULIN (1955e) first described Massartellopsis which was redescribed by PETERS & EDMUNDS (1972) based on reared adults. A more comprehensive taxonomic account of the genus was given by PESCADOR & PETERS in1987. Adult Characteristics: 1) Fork of vein MA of fore wings symmetrical (Fig. 156A); 2) hind wings without costal projection (Fig. 156B); 3) vein Sc of hind wings 9/10 maximum length of wing (Fig. 156B); 4) claws of a pair similar, both apically hooked with opposing hook (Fig. 156C); 5) penis lobes divided, tubular, with tongue-like apical projection on each lobe (Figs. 156D-E); and 6) female sternum IX entire apically (Fig. 156F). Nymphal Characteristics (Fig. 157A): 1) Clypeus narrower than labrum and lateral margins of clypeus divergent apically (Fig. 158A); 2) anteromedian emargination of labrum deeply V-shaped (Fig. 158B); 3) outer margin of mandibles angular with series of short setae on apical half and a median setae tuft (Fig. 158C); 4)

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MA

A

Sc CuP

F ICu1 CuA

D

B

C ap

G

H

E Plate 156. Massartellopsis irarrazavali, imago. Figs. 156A-H: 156A, fore wing; 156B, hind wing; 156C, fore claw; 156D, male genitalia, v.v.; 156E, penis lobe, d.v.; 156F, apex of female sternum IX; 156GH, abdominal terga IV-VII. (MA = fork of MA; ap = apical projection). Figs. 156A-H modified from PESCADOR & PETERS, 1987.

submentum of labium with thick ventrolateral spine-like setae on surface (Fig. 158D); 5) tarsal claws with denticles a little larger apically (Fig. 158E); 6) posterolateral projections on abdominal segments II-IX; and 7) gills on abdominal segments I-VII similar, gradually tapered apically, with many tracheal branches (Figs. 157A, 158F). The nymphs of this genus (Fig. 157A) are similar to those of Meridialaris, but usually can be separated by the dark, heavily branched tracheation of the gills and the presence of setae on the sides of submentum. Distribution: Mountain ranges of Bolivia southward to the Andean and Patagonic region of Chile and Argentina. Biology: The nymphs occur under stones or small rocks in cold and moderately swift flowing streams and rivers. The nymphs also inhabit the gravelly shorelines of lakes. Some nymphs collected from Southern Argentina and Chile were parasitized by the chironomid Symbiocladius wygodzinski ROBACK.

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Plate 157. Massartellopsis irarrazavali. Fig. 157A, nymphal habitus. Figure modified from PESCADOR & PETERS, 1987.

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Massartellopsis irarrazavali DEMOULIN Massartellopsis irarrazavali DEMOULIN, 1955e: 9 (male, female); PETERS & EDMUNDS, 1972: 1409 (male, female, nymph); HUBBARD, 1982a: 265; DOMÍNGUEZ & SAVAGE, 1987: 44; PESCADOR & PETERS, 1987: 183 (male, female, nymph, egg); HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 214; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 52 (male, female, nymph).

Taxonomy: Massartellopsis irarrazavali is the only known species in the genus, and can be recognized by the characters listed in the generic diagnosis. PESCADOR & PETERS (1987) presented a brief discussion on intraspecific color variations (Figs. 156G-H). Distribution and Biology: As in generic description. Genus Meridialaris PETERS & EDMUNDS Deleatidium [partim] EATON, 1899: 288; ULMER, 1920a: 23; 1938: 85; LESTAGE, 1931b: 52; DEMOULIN, 1955b: 13; 1955e: 23. Meridialaris PETERS & EDMUNDS, 1972: 1405; DOMÍNGUEZ & SAVAGE, 1987: 44; PESCADOR & PETERS, 1987: 148; SAVAGE, 1987b: 207; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 52. (Type-species: Deleatidium laminatum ULMER, original designation).

A

C B

D

E

S

F

D

Plate 158. Massartellopsis irarrazavali, nymph. Figs. 158A-F: 158A-D: Mouthparts. 158A, clypeus and labrum, d.v.; 158B, anteromedian emargination of labrum, enlarged; 158C, left mandible; 158D, labium (left dorsal, right ventral); 158E, fore claw; 158F, gill IV. (D = denticles; S = spine-like setae). Figs. 158A-F modified from PESCADOR & PETERS, 1987.

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Taxonomy: Meridialaris was established by PETERS & EDMUNDS (1972) who designated Deleatidium laminatum ULMER as the type-species. PETERS & EDMUNDS (1972) transferred all species in South America previously identified as Deleatidium to Meridialaris. A taxonomic review of Meridialaris was given by PESCADOR & PETERS (1987), who discussed the questionable taxonomic status of M. patagonica and M. lestagei (previously in Atalophlebioides). These species are not included in the keys. Adult Characteristics: 1) Fork of vein MA of fore wings symmetrical (Fig. 159A); 2) hind wings without costal projection and vein Sc 9/10 maximum length of wing (Fig. 159C); 3) tarsal claws dissimilar, one apically hooked and with small opposing hook, and other pad-like (Fig. 159D); 4) penis lobes of male genitalia fused to near apex with apical projection on each lobe (Figs. 159F-N); and 5) female sternum IX pointed apically (Fig. 159E). Nymphal Characteristics (Fig. 160A): 1) Clypeus narrower than labrum and lateral margins of clypeus divergent apically (Figs. 161A-B ); 2) anteromedian emargination of labrum shallow U-shaped to deeper V-shaped (Fig. 161A-D); 3) outer margin of mandibles slightly angular (Fig. 161F) to smoothly curved (Fig. 161E), and with a series of short setae and median seta tuft (Fig. 161F); 4) submentum of labium glabrous (Fig. 161G); 5) tarsal claws with denticles progressively larger apically (Fig. 161H); 6) posterolateral spines on abdominal segments IIIX; and 7) abdominal gills I-VII similar, slender, tapered apically (Figs. 161M-N). The nymphs of this genus (Fig. 160A) are difficult to separate from nymphs of Massartellopsis, but can be distinguished by the lack of setae on the submentum and the lighter coloration of the tracheal gills of the species of Meridialaris. Distribution: Magallanes Province (ca. 53°S) XII Region, north to Isluga (19º20´S), I Region, Chile, and in Argentina from Chubut Province to Southern Bolivia highlands, with easternmost record in Tucumán Province (ca. 65° W). Biology: The nymphs of Meridialaris occur in streams of various sizes, rivers, and lakes. The nymphs are found generally under rocks, although their location in the river varies between species. The gut content is composed of detritus, diatoms and filamentous algae. Subimagos generally emerge in the afternoon, and adults have been reported flying from November to March. Mating flights have been observed on the river banks in the afternoon (between 16:00 and 18:00 h.). Specimens of this genus have been collected from sea level in Patagonia, up to 4300 m in Northern Argentina. Some species of Meridialaris are parasitized by Symbiocladius (Chironomidae). KEYS

TO

MERIDIALARIS

Male Imagos 1 –

Penes square-shaped (Fig. 159M); well developed angular bend on inner margin of segment I of genital forceps located approximately 1/2 distance from base (Fig. 159L); maculae on abdominal terga as in Fig. 159T ........ Meridialaris laminata Penes not as above; angular bend on inner margin of segment I of genital forceps located approximately 1/3 distance from base (Fig. 159H, N); maculae on abdominal terga not as above .......................................................................................... 2

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Penes bell-shaped with blunt finger-like apical projections (Fig. 159N); angular bend on inner margin of segment I of genital forceps weakly developed (Fig. 159N); maculae on abdominal terga as in Fig. 159U ........... Meridialaris tintinnabula Penes cylindrical (Figs. 159F-G) or swollen (Figs. 159I-K); angular bend on inner margin of segment I of genital forceps well developed (Fig. 159H); maculae on abdominal terga not as above .......................................................................................... 3 Penes with prominent setae between apical projections (Figs. 159F-G); maculae on abdominal terga pronounced (as in Figs. 159O-P) ............................................... 4 Penes glabrous between apical projections (Figs. 159I-K); maculae on abdominal terga not as pronounced (Fig. 159S), or weakly developed to almost absent (Figs. 159Q-R) ................................................................................................................................ 5 Vein ICu1 of fore wings parallel to slightly divergent distally from vein CuA (Fig. 159B); maculae on abdominal terga pronounced as in Fig. 159P ............................. .............................................................................................................. Meridialaris chiloeensis Vein ICu1 of fore wings strongly divergent distally from vein CuA (Fig. 159A); maculae on abdominal terga as in Fig. 159O ............................. Meridialaris biobionica Femora lack median band; basal 3/5 of penes swollen (Figs. 159J-K ) and apical projections weakly developed (Fig. 159K); maculae on abdominal terga as in Fig. 159S .......................................................................................................... Meridialaris inflata Femora with dark brown median band; penes broad with well developed subapical projections (Fig. 159I); maculae on abdominal terga greatly reduced to almost absent as in Figs. 159Q-R ................................................................ Meridialaris diguillina

Nymphs 1 – 2(1) – 3(2) – 4(1) –

Labrum with deep, V-shaped anteromedian emargination (Figs. 161B, D) .......... 2 Labrum with broad and shallow U-shaped anteromedian emargination (Fig. 161C) ..................................................................................................................................... 4 Tarsi with long, needle-like spines (Figs. 161K-L); lateral margins of labrum angularly curved (Fig. 161A) .......................................................... Meridialaris chiloeensis Tarsi without long needle-like spines (Figs. 161I-J); lateral margins of labrum smoothly curved (Fig. 161B) ........................................................................................... 3 Outer margin of cardo with less than 25 long spines; joints of antennal flagella with reddish brown annulations; femora with 2 dorsal pale yellow spots ............... ..................................................................................................................... Meridialaris spina Outer margin of cardo with more than 25 long spines; joints of antennal flagella lack annulations; femora with 3 dorsal pale yellow spots .......................... .......................................................................................................... Meridialaris tintinnabula Maculae on abdominal terga as in Fig. 159T; abdominal gills moderately broad, main tracheal trunk with several well developed branches (Fig. 161M) .................. ............................................................................................................... Meridialaris laminata Maculae on abdominal terga as in Figs. 159Q-R; abdominal gills slender, main tracheal trunks without or with few weakly developed branches (Fig. 161O) ....... ............................................................................................................... Meridialaris diguillina

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Imagos Machos 1

– 2(1)

– 3(2) – 4(3) – 5(3) –

Penes cuadrangulares (Fig. 159M); recodo angular bien desarrollado en el margen interno de segmento I de los fórceps genitales ubicado cerca de 1/2 de la distancia de la base (Fig. 159L); manchas en tergos abdominales como en Fig. 159T .......... ............................................................................................................... Meridialaris laminata Penes no como en el caso anterior; recodo angular en el margen interno de segmento I de los fórceps genitales ubicado cerca de 1/3 de la distancia de la base al ápice (Figs. 159H, N); manchas en tergos abdominales no como en el caso anterior ................... 2 Penes en forma de campana con proyecciones apicales digitiformes obtusas (Fig. 159N); recodo angular en el margen interno de segmento I de los fórceps genitales poco desarrollado (Fig. 159N); manchas en los tergos abdominales como en Fig. 159U .................................................................................. Meridialaris tintinnabula Penes cilíndricos (Figs. 159F-G) o ensanchados (Figs. 159I-K); recodo angular en el margen interno de segmento I de los fórceps genitales bien desarrollado (Fig. 159H); manchas en los tergos abdominales no como en el caso anterior ............. 3 Penes con setas prominentes entre las proyecciones apicales (Figs. 159F-G); manchas en los tergos abdominales marcadas (como en las Figs. 159O-P) ......... 4 Penes glabros entre las proyecciones apicales (Figs. 159I-K); manchas en los tergos abdominales no pronunciadas (Fig. 159S), o débilmente desarrolladas hasta casi ausentes (Figs. 159Q-R) ............................................................................................ 5 Vena ICu1 de las alas anteriores paralela a levemente divergentes distalmente de la vena CuA (Fig. 159B); manchas en los tergos abdominales marcadas, como en Fig. 159P .................................................................................................... Meridialaris chiloeensis Vena ICu1 de las alas anteriores fuertemente divergente distalmente de la vena CuA (Fig. 159A); manchas en los tergos abdominales como en Fig. 159O ............ .............................................................................................................. Meridialaris biobionica Fémures sin banda medial; 3/5 basales de los penes ensanchados (Figs. 159J-K) y proyecciones apicales poco desarrolladas (Fig. 159K); manchas en los tergos abdominales como en Fig. 159S ........................................................ Meridialaris inflata Fémures con banda medial castaño obscura; penes anchos con proyecciones subapicales bien desarrolladas (Fig. 159I); manchas en los tergos abdominales reducidas a casi ausentes, como en las Figs. 159Q-R ................ Meridialaris diguillina

Ninfas 1 – 2(1) –

Labro con profunda emarginación anteromediana en forma de V (Figs. 161B, D) ................................................................................................................................................ 2 Labro con emarginación anteromediana ancha y poco profunda en forma de U (Fig. 161C) ........................................................................................................................... 4 Tarsos con espinas largas en forma de aguja (Figs. 161K-L); márgenes laterales del labro curvados angularmente (Fig. 161A) ................................... Meridialaris chiloeensis Tarsos sin espinas largas en forma de aguja (Figs. 161I-J); márgenes laterales del labro suavemente curvados (Fig. 161B) ........................................................................ 3

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MA Sc

A ICu1

MP1 CuA MP2

E

D

C

CuA ICu1

B

H

I

K J

F

G

N

L M O

P

Q

R

S

T

U

Plate 159. Meridialaris, imagos. Figs. 159A-U. 159A, fore wing; 159B, detail of CuA area of fore wing; 159C, hind wing; 159D, male tarsal claw; 159E, apex of female sternum IX; 159A, 159C-E: Meridialaris laminata; 159B, M. chiloeensis. 159F-N, male genitalia. 159F, M. biobionica, penes, v.v.; 159G, M. chiloeensis penes, v.v.; 159H-I, M. diguillina, 159H, genitalia, v.v., 159I, penes, d.v.; 159J-K, M. inflata, penes, 159J, v.v, 159K, d.v.; 159L-M, M. laminata, 159L, genitalia, v.v, 159M, penes, d.v.; 159N, M. tintinnabula, genitalia, v.v. 159O-U, abdominal terga. 159O, M. biobionica (IV-V); 159P, M. chiloeensis (III-VII); 159Q-R, M. diguillina (III-VII); 159S, M. inflata (III-VII); 159T, M. laminata (III-VII); 159U, M. tintinnabula (III-VII). Figs. 159A-U, modified from PESCADOR & PETERS, 1987.

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Margen externo del cardo con menos de 25 espinas largas; articulaciones del flagelo de las antenas con anillos castaño-rojizos; fémures con 2 manchas dorsales amarillo pálidas ........................................................................................ Meridialaris spina Margen externo del cardo con más de 25 espinas largas; articulaciones del flagelo de las antenas sin anillos; fémures con 3 manchas amarillo pálidas ......................... .......................................................................................................... Meridialaris tintinnabula Manchas de los tergos abdominales como en Fig. 159T; branquias abdominales moderadamente anchas, tronco traqueal principal con varias ramas bien desarrolladas (Fig. 161M) ................................................................ Meridialaris laminata Manchas de los tergos abdominales como en Figs. 159Q-R; branquias abdominales angostas, tronco traqueal principal sin o con pocas ramas débilmente desarrolladas (Fig. 161O) ................................................................. Meridialaris diguillina

Meridialaris biobionica (ULMER) Deleatidium biobionicum ULMER, 1938: 85 (male, female); ULMER, 1943: 15; TRAVER, 1946: 423; DEMOULIN, 1955b: 15; PETERSEN & GAEDIKE, 1968: 960. Meridialaris biobionica; PETERS & EDMUNDS, 1972: 1405; HUBBARD, 1982a: 265; PESCADOR & PETERS, 1987: 151 (male).

Taxonomy: ULMER (1938) described Meridialaris biobionica from one male and three female imagos from Biobío Prov., Chile, as Deleatidium biobionicum. The species was redescribed by PESCADOR & PETERS (1987), and can be distinguished from other species of Meridialaris by the following combination of characters. In the imagos: 1) Vein ICu1 of fore wings strongly divergent distally from vein CuA (as in Fig. 159A), 2) femora with dark brown median band; 3) maculae of abdominal terga covering most of each tergum (Fig. 159O); 4) angular bend on inner margin of segment I of genital forceps approximately 1/3 distance from base; 5) penes with setae between blade-like apical processes (Fig. 159F) and 6) caudal filaments without markings at annulations. The nymphs are unknown. Distribution and Biology: Southern central Chile. Except for collection records for adults in December and January, the biology of the species is unknown. Meridialaris chiloeensis (DEMOULIN) Deleatidium chiloeense DEMOULIN, 1955b: 13 (male); DEMOULIN, 1955e: 23 (male). Meridialaris chiloeense; PETERS & EDMUNDS, 1972: 1405; PESCADOR & PETERS, 1987: 154 (male, female, nymph). Meridialaris chiloeensis; HUBBARD, 1982a: 265; HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 214; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 54 (male, female). Deleatidium penai DEMOULIN, 1955e: 27 (female). Meridialaris penai; PETERS & EDMUNDS, 1972: 1405; HUBBARD, 1982a: 266.

Taxonomy: Meridialaris chiloeensis was described from male imagos by DEMOULIN (1955b as Deleatidium chiloeense) and D. penai was described later the same year from female imagos and subimagos. The species was redescribed by PESCADOR & PETERS (1987), who included the first nymphal descriptions and synonymized the two species.

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A

Plate 160. Meridialaris laminata. Fig. 160A, nymphal habitus. Figure modified from PESCADOR & PETERS, 1987.

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M. chiloeensis can be distinguished from the other species of the genus by the following combination of characters. In the imagos: 1) Vein ICu1 of fore wings parallel to slightly divergent from vein CuA (Fig. 159B); 2) femora without median band; 3) pattern of dark brown maculae on abdominal terga as in Fig. 159P; 4) well developed angular bend on inner margin of segment I of genital forceps approximately 1/3 distance from base (as in Fig. 159H); and 5) penes cylindrical with minute setae between blade-like apical processes (Fig. 159G). In the nymph: 1) Labrum with narrow, deep, V-shaped anteromedian emargination and angular lateral margins (Fig. 161D); 2) pro- and mesonota with pale yellow spots; 3) tarsi with needle-like spines (Fig. 161K-L); and 4) moderately broad abdominal gills with well developed tracheal branches (Fig. 161N). Distribution and Biology: Confined to the eastern and western slopes of the southern South American Andes. The nymphs are found under stones or small rocks in swift areas of streams and rivers, and lake shorelines. The nymphs have been collected from September to March and the adults from November to March. The adults emerged in the field between 16:00-18:00 h while reared adults emerged anytime during the day. A small group of adults was observed swarming at 16:30 h above an isolated shrub on the bank of the river, and collected specimens were mostly female imagos. Nymphs reportedly feed on detritus, mineral particles, diatoms and filamentous algae. The nymphs of the M. chiloeensis are sometimes parasitized by the chironomid, Symbiocladius wygodzinski ROBACK. Meridialaris diguillina (DEMOULIN) Deleatidium diguillinum DEMOULIN, 1955e: 23 (male, female). Meridialaris diguillina; PETERS & EDMUNDS, 1972: 1405; HUBBARD, 1982a: 265; PESCADOR & PETERS, 1987: 159 (male, female, nymph); HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 215. Meridialaris diguilina [sic]; WAIS, 1985: 2167 (nymph); WAIS & BONETTO, 1988: 1457. Meridialaris diquillina [sic]; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 55 (male, nymph).

Taxonomy: DEMOULIN (1955e) described Meridialaris diguillina (as Deleatidium diguillinum) from male and female adults. The species was subsequently redescribed by PESCADOR & PETERS (1987) who included the description of the nymph and a brief discussion on intraspecific variations. Meridialaris diguillina can be distinguished from the other species of the genus by the following combination of characters. In the imagos: 1) Vein ICu1 of fore wings strongly divergent distally from vein CuA (as in Fig. 159A); 2) femora with dark brown median band; 3) maculae on abdominal terga as in Figs. 159Q-R; 4) well developed angular bend on inner margin of segment I of genital forceps approximately 1/3 distance from base (Fig. 159H); 5) penes with slender apical projections (Fig. 159I); and 6) caudal filaments uniformly pale yellow. In the nymph: 1) Labrum with broad, shallow anteromedian emargination (as in Fig. 161C); 2) antennal flagella uniformly pale yellow; 3) tarsi with short blunt spines (Fig. 161I); and 4) slender abdominal gills with few weakly developed branches (Fig. 161O), or unbranched. Distribution and Biology: Western and eastern slopes of the southern Andes, from Rio Diguillin (37ºS), Prov. Ñuble, southward to Rio Elqui (29º54´S), IV Region, Chile. According to PESCADOR & PETERS (1987) the nymphs of M. diguillina occur in streams of

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various sizes, as well as in rivers and lakes. Compared to M. laminata and M. chiloeensis, the nymphs of M. diguillina extend to near the mid-stream where the current is swift and almost inaccessible for collecting. Collection records indicate that the nymph and adults of M. diguillina occur in September through March, and November through February respectively. The adults have been collected in abundance by light traps. Meridialaris inflata PESCADOR & PETERS Meridialaris inflata PESCADOR & PETERS, 1987: 165 (male).

Taxonomy: Meridialaris inflata was described by PESCADOR & PETERS (1987) from male imagos in Chile. The female imago, and nymphs are still unknown. Meridialaris inflata differs from the other species of the genus by: 1) Absence of median banding on femora; 2) absence of banding on annulations of caudal filaments; 3) distinctive abdominal tergal maculae (Fig. 159S); and 4) basal 3/5 of penes swollen and apical projections short (Figs. 159J-K). According to PESCADOR & PETERS (1987) the species appears closely related to M. diguillina but can be distinguished from it by the distinctive abdominal maculae, unbanded femora, and swollen penes. Distribution: Southern Chile, Malleco Prov. 38º20’S. Meridialaris laminata (ULMER) Deleatidium laminatum ULMER, 1920a: 23 (male, female); ULMER, 1920c: 115; LESTAGE, 1931b: 56; ULMER, 1938: 105; ULMER, 1943: 15; TRAVER, 1946: 423; DEMOULIN, 1955b: 15. Meridialaris laminata; PETERS & EDMUNDS, 1972: 1405; HUBBARD, 1982a: 265; WAIS, 1985: 2167; PESCADOR & PETERS, 1987: 168 (male, female, nymph, egg); WAIS & BONETTO, 1988: 1457; HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 215; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 56 (male, female, nymph). Deleatidium illapeli DEMOULIN, 1955e: 26 (female). Meridialaris illapeli; PETERS & EDMUNDS, 1972: 1405; HUBBARD, 1982a: 265.

Taxonomy: Meridialaris laminata was first described from adults by ULMER (1920a) in the genus Deleatidium. Deleatidium illapeli was described from a female imago by DEMOULIN (1955e). The species was later redescribed by PESCADOR & PETERS (1987) who included the first nymphal description and synonymized M. illapeli with M. laminata. Meridialaris laminata can be distinguished from all the other species of the genus by the following combination of characters. In the imago: 1) Vein ICu1 of fore wings strongly divergent distally from vein CuA (Fig. 159A); 2) femora with a dark brown band; 3) dark brown maculae on abdominal terga as in Fig. 159T; 4) well developed angular bend on inner margin of segment I of genital forceps approximately 1/2 distance from base (Fig. 159L); 5) penes square-shaped with blade-like apical projections (Fig. 159M) and 6) caudal filaments with reddish brown annulations at articulations. In the nymph: 1) Labrum with broad, U-shaped anteromedian emargination (Fig. 161C) and angular lateral margins (Fig. 161A); 2) pro- and mesonota with pale yellow spots; 3) tarsi with short spines (as in Fig. 161I); and 4) abdominal gills comparatively broad with many branches on both sides of main tracheal trunk (Fig. 161M).

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B

A

C

D

E

F

H

G

SM

D

I

M

N

J

K

L

O

Plate 161. Meridialaris, nymphs. Figs. 161A-O. 161A-G, nymphal mouthparts. 161A-B, Clypeus and labrum, d.v.: 161A, M. laminata; 161B, M. tintinnabula (outline); 161C-D, anteromedian emargination of labrum: 161C, M. laminata; 161D, M. chiloeensis; 161E-F, left mandible: 161E, M. tintinnabula; 161F, M. laminata; 161G, labium (left dorsal, right ventral). 161H, Fore claw, M. laminata. 161I-L, nymphal metatarsal spines: 161I, M. diguillina; 161J, M. spina; 161K-L, M. chiloeensis. 161M-O, Gill IV: 161M, M. laminata; 161N, M. chiloeensis; 161O, M. diguillina. (SM = Submentum; D = Denticle). Figs. 161A-O modified from PESCADOR & PETERS, 1987.

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Distribution and Biology: Widely distributed in Chile, from Valdivia (37º40´S) to Coquimbo (29º57´S) IV Region, in elevations from 200 to 2000 m; in Argentina, in Neuquen and Rio Negro Provinces. The nymphs occur in streams and rivers, generally in crevices of stones or under small rocks, although they also have been found under submerged logs or roots. The emergence of adults generally takes place between 16:00 and 19:00. Swarms can be composed of a few (about 10) to many (100) individuals, generally many more males than females. The location of the flights observed varied from 1.5 to 10 m above the water, and lasted between 15 to 60 minutes. Some nymphs are parasitized by the chironomid Symbiocladius wygodzinski ROBACK. Meridialaris lestagei (ULMER) “Nymphen des Leptophlebia typus” ULMER, 1904: 4. Atalophlebia fulvipes; sensu LESTAGE, 1931b: 52. Atalophlebioides lestagei ULMER, 1938: 102.

Taxonomy: This nymph was described by ULMER (1904) and eventually given the name Atalophlebioides lestagei by ULMER (1938). It is clearly not Atalophlebiodes but fits within the definition of Meridialaris (M. lestagei n. comb.). The nymph is from the same type locality as M. patagonica and may represent the same species but type material has not been examined. The species is not included in keys. Distribution: Southern Chile. Meridialaris patagonica (LESTAGE) Atalophlebia chilensis; sensu ULMER, 1904: 7 (female). Atalophlebia patagonica LESTAGE, 1931b: 52. Deleatidium patagonicum; ULMER, 1938: 105 (female); ULMER, 1943: 15; TRAVER, 1946: 419. Meridialaris patagonica; PETERS & EDMUNDS, 1972: 1405; HUBBARD , 1982a: 265; PESCADOR & PETERS, 1987: 149.

Taxonomy: ULMER (1904) identified one female imago from South Patagonia, Punta Arenas, as Atalophlebia chilensis. LESTAGE (1931b) recognized that it was not A. chilensis and established a new species A. patagonica for the specimen; ULMER (1938) transferred it to Deleatidium. Although included in Meridialaris by PETERS & EDMUNDS (1972), the inadequate description and illustration make it impossible to identify the species until types are examined. Distribution: Southern Chile. Meridialaris spina PESCADOR & PETERS Meridialaris spina PESCADOR & PETERS, 1987: 174 (nymph); HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 215; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 56 (nymph).

Taxonomy: Meridialaris spina was described from nymphs, and adults are unknown. Nymphs are distinguished from the other species of the genus by the following combination of characters: 1) Labrum with narrow, deep, V-shaped anteromedian emargination (as in Fig. 161D) and with rounded lateral margins (as in

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Fig. 161B); 2) moderately long tarsal spines (Fig. 161J); 3) femora with two dorsal pale spots; 4) moderately broad abdominal gills with several branches (as in Fig. 161N); and 5) abdominal terga yellowish brown to dark brown, with black maculae near center and near base of gills. Distribution and Biology: Andean slopes of Chile and Argentina. In Chile the species has been collected from Santiago (33º27´S) south to Rio Simpson (45º34´S), XI Region. In Argentina, it is confined in the Andean streams of Neuquén and Rio Negro Provinces. The nymphs inhabit small rocks or stones in swift flowing streams, and were collected from November to February. Meridialaris tintinnabula PESCADOR & PETERS Meridialaris tintinnabula PESCADOR & PETERS, 1987: 171 (male, nymph); HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 216; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 57 (male, nymph).

Taxonomy: PESCADOR & PETERS (1987) described M. tintinnabula from male imagos, subimagos, and nymphs collected from high mountain streams in Tucumán Province, Argentina. Meridialaris tintinnabula can be distinguished from the other species of the genus by the following combination of characters. In the imagos: 1) Vein ICu1 of fore wings parallel to slightly divergent distally from vein CuA (Fig. 159B); 2) pattern of dark brown maculae on abdominal terga as in Fig. 159U; 3) genital forceps with weakly developed angular bend on inner margin approximately 1/3 distance from base (Fig. 159N); and 4) bell-shaped penes with finger-like apical projections (Fig. 159N). In the nymphs: 1) Labrum with narrow, deep V-shaped anteromedian emargination (as in Fig. 161D) and rounded lateral margins (Fig. 161B); 2) moderately long tarsal spines (as in Fig. 161J); and 3) moderately broad abdominal gills with few to several well developed branches (as in Fig. 161N). Distribution and Biology: Andean region of Northwestern Argentina. The nymphs have been collected under rocks in mountain streams, between 800 and 4300 m in Tucumán Province, Argentina. The adults were found hidden among rocks nearby. The species can be found in rivers occupied by Thraulodes species, although Meridialaris is dominant at higher altitudes and Thraulodes at lower altitudes (DOMÍNGUEZ & BALLESTEROS VALDEZ, 1992). Genus Microphlebia SAVAGE & PETERS Microphlebia SAVAGE & PETERS, 1983: 569; ORTH et al., 2000: 31. (Type-species: Microphlebia surinamensis SAVAGE & PETERS, original designation).

Taxonomy: Microphlebia was established by SAVAGE and PETERS, and since then no other systematic reference to the genus was made, except for its inclusion in generic keys (DOMÍNGUEZ et al., 2001). This genus is known only from male subimagos and nymphs, and for this reason the key to males must be used with care. Segment II of the forceps of males is clearly long in relation to segment I, but exact ratios cannot be determined. Adult Characteristics: 1) Penes tubular, divided to base, without appendages (Figs. 162A-B); 2) fork of veins MA and MP of fore wings asymmetrical (Fig. 162C); 3) vein MP

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of hind wings not forked (Figs. 162D-E); 4) costal projection of hind wings well developed, acute and rounded (Figs. 162D-E); 5) vein Sc of hind wings terminating at base of costal projection (Figs. 162D-E); and 6) tarsal claws of a pair dissimilar, one apically hooked, other blunt (Fig. 162F), similar in structure on all legs. Nymphal Characteristics (Fig. 163A): 1) Width of labrum equal to width of clypeus (Fig. 163B); 2) outer margin of mandibles smoothly curved with few setae medially (Fig. 163C); 3) glossae of labium ventrally curved (Fig. 163D); 4) gills on abdominal segments IVII oval with margins entire, dorsal portion smaller than ventral portion (Fig. 163E); 5) posterolateral spines on abdominal segments V-VI to IX; and 6) denticles on tarsal claws progressively larger apically, except apical denticle much larger (Fig. 163F). Distribution: Surinam, French Guiana, Venezuela and northern Brazil. Biology: The nymphs live in slow rivers, in the detritus. The subimagos were collected at light in December.

MP

MA

A

C

D

CP

Sc

F B

E MP

G

H

Plate 162. Microphlebia, adults. Figs. 162A-B: male genitalia, v.v.: 162A, M. surinamensis; 162B, M. pallida. 162C-E, wings M. surinamensis: 162C, fore wing; 162D, hind wing; 162E, hind wing, detail. 162F, male fore tarsal claw of M. surinamensis. 162G-H, Subimaginal male eyes, d. v.: 162G, M. pallida; 162H, M. surinamensis. (CP = Costal Projection). Figs. 162A-H, modified from SAVAGE & PETERS, 1983.

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KEYS

TO

MICROPHLEBIA

Subimagos 1 –

Upper portion of male eyes with 23 to 30 facets in longest row (Fig. 162G); femora uniformly yellowish brown ................................................. Microphlebia pallida Upper portion of male eyes with 17 to 24 facets in longest row (Fig. 162H); femora yellowish brown, with apical 1/3 tinted blackish brown ............................... ........................................................................................................ Microphlebia surinamensis E

A

D

F

B

C

D Plate 163. Microphlebia, nymph. Figs. 163A-F, M. surinamensis. 163A, habitus. 163B-D, mouthparts: 163B, clypeus and labrum, d.v.; 163C, left mandible; 163D, labium, left dorsal, right ventral; 163E, gill IV; 163F, tarsal claw. (D = denticle). Figs. 163A-F, modified from SAVAGE & PETERS, 1983.

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Nymphs 1 –

CLAVES

Femora uniformly yellowish brown; membrane of abdominal gills light grayish ..... ................................................................................................................. Microphlebia pallida Femora yellowish brown, with apex shaded blackish (Fig. 163A); membrane of abdominal gills blackish brown ............................................... Microphlebia surinamensis PARA

MICROPHLEBIA

Subimagos 1 –

Porción superior de los ojos del macho con 23 a 30 facetas en la hilera más larga (Fig. 162G); fémures uniformemente castaño-amarillentos ....... Microphlebia pallida Porción superior de los ojos del macho con 17 a 24 facetas en la hilera más larga (Fig. 162H); fémures castaño-amarillentos, con el 1/3 apical teñido de castañonegruzco ....................................................................................... Microphlebia surinamensis

Ninfas 1 –

Fémures uniformemente castaño-amarillentos; membrana de las branquias abdominales grisáceo-claras .............................................................. Microphlebia pallida Fémures castaño-amarillentos, con el ápice sombreado de negruzco (Fig. 163A); membrana de las branquias abdominales castaño-negruzcas ..................................... ........................................................................................................ Microphlebia surinamensis

Microphlebia pallida SAVAGE & PETERS Microphlebia pallida SAVAGE & PETERS, 1983: 574 (male, nymph).

Taxonomy: Microphlebia pallida can be distinguished from the other species of the genus by the following combination of characters. In the male subimago: 1) Body length 4.4-5.0 mm; 2) upper portion of male eyes with 23-30 facets in longest row (Fig. 162G); and 3) femora uniformly yellowish brown. In the nymph: 1) Head and abdominal terga lightly washed with brownish black; 2) femora uniformly yellowish brown; and 3) membrane of abdominal gills light gray. Distribution and Biology: Northern Brazil. The nymphs were collected in detritus, in the slow current zone. Subimagos were collected at light. Microphlebia surinamensis SAVAGE & PETERS Microphlebia surinamensis SAVAGE & PETERS, 1983: 572 (male, nymph); SAVAGE, 1983b: 125; SAVAGE, 1987b: 107 (male).

Taxonomy: Microphlebia surinamensis can be differentiated from the other species of the genus by the following combination of characters. In the male subimago: 1) Body length 3.6-4.4 mm; 2) upper portion of male eyes with 17-24 facets in longest row (Fig. 162H); and 3) femora yellowish brown with apical 1/3 blackish brown. In the nymph: 1) Head and abdominal terga heavily washed with brownish black; 2) femora yellowish brown, with apical 1/5-1/3 washed with brownish black (Fig. 163A); and 3) membrane of abdominal gills blackish gray.

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Distribution and Biology: Surinam and Venezuela. The subimagos were collected at light at the end of December. Genus Miroculis EDMUNDS Miroculis EDMUNDS, 1963: 34; SAVAGE & PETERS, 1983: 496; SAVAGE, 1983b: 124; SAVAGE, 1987a: 98. (Type-species: Miroculis rossi EDMUNDS, original designation).

Taxonomy: Miroculis was described by EDMUNDS in 1963, based on three male imagos. In 1983, SAVAGE & PETERS described the nymph of the genus and nine new species, additionally dividing the genus into four subgenera. In 1987, SAVAGE added another two new species, raising the total to twelve known species. Because of the fact that neither females nor nymphs are separable at the subgeneric level, we give only characters of the male imago in the discussion of the subgenera. A revision of subgeneric characters of male eyes (PETERS et al., in press) is followed here. Adult Characteristics: 1) Fork of vein MA of fore wings asymmetrical (Fig. 164A); 2) vein MP2 of fore wings joined at base to MP1 and CuA by cross veins (Fig. 164A) or vein MP2 reduced and joined to IMP by a cross vein (Fig. 164D); 3) marginal intercalaries present along posterior margin of fore wings (Figs. 164D, F); 4) costal projection of hind wings well developed, acute, apex tent-shaped and projecting from costal margin basal to point where Sc meets C (Figs. 164B-C, E, I); 5) prothoracic tarsal claws of male differing in shape from other claws; 6) male prothoracic claws of a pair unique, one ending in acute finger-like projection, other blunt (Fig. 164L); 7) penes tubular, divided, without appendages (Figs. 165A-G); 8) male forceps segment II long, more than 1/4 length of segment I (Figs. 165A-K) and 9) female sternum IX apically cleft (Fig. 164M). Nymphal Characteristics (general aspect as in Fig. 166A): 1) Labrum somewhat wider than clypeus (Figs. 167A-C), lateral margins from rounded to forming an angle (Figs. 167A-C); 2) tuft of setae medially on lateral margin of mandibles (Fig. 167D); 3) gills on abdominal segments I-VII, with long median apical projection and two shorter posterolateral projections, shape of dorsal and ventral projections sometimes differing (Figs. 167E-K); 4) glossae of labium curved over ventrally; 5) posterolateral spines present on abdominal segments II or III-IX, more developed on V or VI-IX; and 6) denticles on tarsal claws progressively larger apically, apical denticle often much larger (Fig. 167L). Distribution: Northern South America to Southern Brazil and Argentina. Biology: The nymphs of Miroculis have been found in streams and rivers, with slow to rapid current, and in a variety of substrata. According to unpublished data, the emergence period can be rather long (at least from November to April). KEYS

TO

MIROCULIS

Male Imagos 1

Upper portion of eyes on narrow, dorsally-directed stalks, 5 to 20 facets in longest row of dorsal portion, eyes separated on meson of head by a distance 0.5 to 1.5 width of an upper portion (Figs. 165L-M, O-P) ................. Miroculis (Miroculis).....4

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Upper portion of eyes not on stalks (Figs. 165N, Q, 218B), or upper portion on short, wide stalks separated on meson of head by a distance 0.1 to 0.2 maximum width of an upper portion (Figs. 165R, 218C), with 27 to 40 facets of medium size in the longest row ...................................................................................................... 2 MA

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M J K Plate 164. Miroculis, imagos. Figs. 164A-K: Wings, male imago. M. (M.) rossi: 164A, fore wing; 164B, hind wing; 164C, hind wing, detail. M. (M.) marauiae: 164D, fore wing; 164E, hind wing. M. (M.) fittkaui: 164F, fore wing; 164G, hind wing. M. (A.) amazonicus: 164H, fore wing; 164I, hind wing. M. (O.) mourei: 164J, fore wing; 164K, hind wing. 164L, M. (M.) rossi: male fore tarsal claws. 164M, M. (M.) fittkaui: apex of female sternum IX. (R = Radial Sector; CP = Costal Projection). Figs. 164A-M, modified from SAVAGE & PETERS, 1983.

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Dorsal portion of eyes on a short broad stalk, similar to Fig. 165R ........................ ...................................................................................................... Miroculis (Yaruma) wandae Dorsal portion of eyes unstalked (Figs. 165N, Q) ..................................................... 3 Upper portion of eyes separated dorsally by a distance equal to 0.5 to 0.6 maximum width of upper portion; styliger plate rounded (Figs. 165E-F); ratio of maximum width to maximum length of fore wings 0.46 to 0.51 (Fig. 164H) ....... ............................................................................................................ Miroculis (Atroari).....8 Eyes broadly separated dorsally by distance about equal to maximum width of upper portion of eye; posteromedian margin of styliger plate with U-shaped indentation (Figs. 165H, J); ratio of maximum width to maximum length of fore wings 0.41 to 0.43 (Fig. 164J) ............................................... Miroculis (Ommaethus).....9 Fore wings with distinct contrast in coloration between dark basal portion and much lighter apical portion; penes longer than forceps (Fig. 165A) ........................ ............................................................................................... Miroculis (Miroculis) bicoloratus Fore wings variable, but never contrasting as above; penes not extending beyond forceps (Figs. 165B-D) ...................................................................................................... 5 Fore and hind wings with bands or maculae around cross veins (Figs. 164F-G) ................................................................................................................................................ 6 Fore and hind wings without maculae around cross veins ...................................... 7 Fore and hind wings with narrow maculae around cross veins (Figs. 164F-G); penes relatively short (subequal to length of segment I of forceps), hooked apically (Fig. 165B) ................................................................ Miroculis (Miroculis) fittkaui Fore and hind wings with broad maculae or bands around cross veins; penes long and narrow (approximately 1.5 times length of segment I of forceps), each lobe with ventral anteromedian lobe (Fig. 165G) ................. Miroculis (Miroculis) nebulosus Length of penes 2.0 to 2.2 times length of segment I of forceps (Fig. 165D); vein MP2 of fore wings joined at base to veins MP1 and CuA by cross vein (Fig. 164A) ........................................................................................................ Miroculis (Miroculis) rossi Length of penes 0.8 to 1.1 times length of segment I of forceps (Fig. 165C); vein MP2 of fore wings free basally or joined at base to vein IMP by cross vein (Fig. 164D) .....................................................................................Miroculis (Miroculis) marauiae Length of penes 1.7 to 1.9 times length of segment I of forceps, each penis lobe with ventral subapical lobe (Fig. 165E); strong brown bands and maculae in fore wings (Fig. 164H) .............................................................. Miroculis (Atroari) amazonicus Length of penes 1.1 to 1.4 times length of segment I of forceps, each penis lobe tapered apically (Fig. 165F); brown bands and maculae very weak in fore wings ................................................................................................... Miroculis (Atroari) duckensis Length of body 3.8 to 6.0 mm; distal 1/3 of segment I and segments II-III of forceps tinted blackish (Fig. 165J); abdominal terga with color pattern as in Fig. 165X; caudal filaments yellowish brown with brownish bands in basal 1/3 to 1/2 ................................................................................................. Miroculis (Ommaethus) mourei Length of body 6.0 to 8.0 mm; forceps light yellowish (Fig. 165H); abdominal terga semihyaline; caudal filaments light yellow ....... Miroculis (Ommaethus) froehlichi

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CLAVES

Abdominal terga II-VI equally dark, with longitudinal blackish brown submedian, sublateral, and lateral lines repeated on all segments (Fig. 165X) ............................. ................................................................................................. Miroculis (Ommaethus) mourei Abdominal terga not equal nor with lines as above ................................................... 2 Abdominal terga II-III and VI dark, IV-V lighter; tergum VI with a V-shaped median macula (Figs. 165T-U, 167M) ............................................................................ 3 Abdominal terga not as above ........................................................................................ 5 Abdominal terga VI-VIII as in Fig. 165U, abdominal gills small and slender, posterolateral lobes medium-sized to reduced (Fig. 167F) ......................................... .................................................................................................Miroculis (Miroculis) marauiae Abdominal terga different than above, abdominal gills variable, posterolateral lobes medium-sized to large (Figs. 167E-G) ................................................................ 4 Abdominal gills large, with posterolateral lobes as in Fig. 167G; caudal filaments yellowish brown, with blackish bands on segments V-IX ........................................... ............................................................................................ Miroculis (Miroculis) brasiliaensis Abdominal gills small, with posterolateral lobes as in Fig. 167E; caudal filaments light brown, without darker bands .................................... Miroculis (Miroculis) fittkaui Abdominal gills small, dorsal portion of gills II-VI not enlarged at base to form a lobe (Fig. 167H) ..................................................................... Miroculis (Atroari) duckensis Abdominal gills large, dorsal portions of gills II-VI with base enlarged to form a lobe (Figs. 167I-J) ............................................................................................................... 6 Abdominal terga V-VIII equally dark or terga IV-V a little lighter (as in Fig. 165V); gill membranes grayish ..................................................... Miroculis (Atroari) amazonicus Abdominal terga V-VIII progressively darker posteriorly (Fig. 167N); gill membranes yellowish brown ............................................................................................. ........................................... Miroculis (Atroari) colombiensis, Miroculis (Miroculis) nebulosus PARA

MIROCULIS

Imagos Machos 1

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2(1) –

Porción superior de los ojos sobre un pedicelo largo, dirigido dorsalmente, de 5 a 20 facetas en la hilera más larga de la porción dorsal, ojos separados en la mitad de la cabeza por una distancia de 0,5 a 1,5 del ancho de la porción superior (Figs. 165L-M) ........................................................................................ Miroculis (Miroculis).....4 Porción superior de los ojos no ubicadas sobre pedicelo (Figs. 165N, Q, 218B), o porción superior sobre un pedicelo corto y ancho, separados en la mitad de la cabeza por una distancia de 0,1 a 0,2 el ancho máximo de la porción superior (Fig. 165R, 218C), con 27 a 40 facetas de tamaño mediano en la hilera más larga ............................................................................................................. 2 Porción dorsal de los ojos sobre un pedicelo corto y ancho, similar al de la Fig. 165R ............................................................................................ Miroculis (Yaruma) wandae Porción dorsal no pedicelada (Figs. 165N, Q) ............................................................. 3

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Porción dorsal de los ojos separados dorsalmente por una distancia igual a 0,5 a 0,6 del ancho máximo de la porción superior; placa estilígera redondeada (Figs. 165E-F); relación ancho máximo/ longitud máxima de las alas anteriores 0,46 a 0,51 (Fig. 164H) .............................................................................. Miroculis (Atroari).....8 Ojos ampliamente separados dorsalmente por una distancia aproximada al ancho máximo de la porción dorsal del ojo; margen posteromediano de la placa estilígera con una indentación en forma de U (Figs. 165H, J); relación ancho máximo/ longitud máxima de las alas anteriores 0,41 a 0,43 (Fig. 164J) ................................... ..................................................................................................... Miroculis (Ommaethus).....9 Alas anteriores con un fuerte contraste de coloración entre la parte basal oscura y la porción apical mucho más clara; penes más largos que los fórceps (Fig. 165A) ............................................................................................... Miroculis (Miroculis) bicoloratus Alas anteriores variables, pero nunca con un contraste tan fuerte entre las porciones basal y apical; penes no extendiéndose más allá de los fórceps (Figs. 165B-D) ......... 5 Alas anteriores y posteriores con manchas alrededor de las venas transversas (Figs. 164F-G) ................................................................................................................................ 6 Alas anteriores y posteriores sin manchas alrededor de las venas transversas ......... 7 Alas anteriores y posteriores con delgadas manchas alrededor de las venas transversas (Figs. 164F-G); penes relativamente cortos (subiguales en longitud al segmento I de los fórceps), terminados en forma de gancho apicalmente (Fig. 165B) ................................................................................................... Miroculis (Miroculis) fittkaui Alas anteriores y posteriores con anchas manchas o bandas alrededor de las venas transversas; penes largos y delgados (aproximadamente 1,5 veces la longitud del segmento I de los fórceps), cada lóbulo con un lóbulo ventral anteromediano (Fig. 165G) .....................................................................................Miroculis (Miroculis) nebulosus Longitud de los penes 2,0 a 2,2 la longitud del segmento I de los fórceps (Fig. 165D); vena MP2 de las alas anteriores unidas en la base a las venas MP1 y CuA por una vena transversa (Fig. 164A) ...................................... Miroculis (Miroculis) rossi Longitud de los penes 0,8 a 1,1 la longitud del segmento I de los fórceps (Fig. 165C); vena MP2 de las alas anteriores libre basalmente o unida a la base de la vena IMP por una vena transversa (Fig. 164D) ..................... Miroculis (Miroculis) marauiae Longitud de los penes 1,7 a 1,9 la longitud del segmento I de los fórceps, cada lóbulo de los penes con un lóbulo subapical ventral (Fig. 165E); en las alas anteriores bandas y manchas castañas muy evidentes (Fig. 164H) .............. Miroculis (Atroari) amazonicus Longitud de los penes 1,1 a 1,4 la longitud del segmento I de los fórceps, cada lóbulo de los penes afinado apicalmente (Fig. 165F); en las alas anteriores bandas y manchas castañas muy débiles ........................................ Miroculis (Atroari) duckensis Longitud del cuerpo 3,8 a 6,0 mm; en los fórceps 1/3 distal del segmento I y segmentos II y III teñidos de negruzco (Fig. 165J); tergos abdominales con un patrón de coloración como en la Fig. 165X; filamentos caudales castaño-amarillentos con bandas castañas en el 1/3 a 1/2 basal ............................... Miroculis (Ommaethus) mourei Longitud del cuerpo 6,0 a 8,0 mm; fórceps amarillento claro (Fig. 165H); tergos abdominales semihialinos; filamentos caudales amarillo claro ................................... ............................................................................................. Miroculis (Ommaethus) froehlichi

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Plate 165. Miroculis, imagos. Figs. 165A-K: Male genitalia, v.v. 165A, M. (M.) bicoloratus; 165B, M. (M.) fittkaui; 165C, M. (M.) marauiae; 165D, M. (M.) rossi; 165E, M. (A.) amazonicus; 165F, M. (A.) duckensis; 165G, M. (M.) nebulosus; 165H, M. (O.) froehlichi; 165I, idem, detail of penes; 165J, M. (O.) mourei; 165K, idem, detail of penes. 165L-R: Male eyes. 165L, M. (M.) bicoloratus, v. l.; 165M, M. (M.) rossi, v. l.; 165N, M. (A.) duckensis, v.l.; 165O, M. (M.) nebulosus, v. l.; 165P, idem, v.d.; 165Q, M. (O.) mourei v.l.; 165R, M. (Yaruma) sp. v. l. 165S-W: male abdominal color pattern. 165S, M. (M.) bicoloratus (II-VII); 165T, M. (M.) fittkaui (III-VII); 165U, M. (M.) marauiae (III-VIII); 165V, M. (A.) amazonicus (IV-VII); 165W, M. (M.) nebulosus (II-VII); 165X, M. (O.) mourei (IV-VI). (numbers between parenthesis = abdominal segment number). Figs. 165A, G, L, N-P, S and W, modified from SAVAGE, 1987; 165B-F, H-K, M, Q-R, T-V and X from SAVAGE & PETERS, 1983.

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Tergos abdominales II-VI igualmente oscuros, con líneas longitudinales castañonegruzcas submedianas, sublaterales y laterales repetidas en todos los segmentos (Fig. 165X) ............................................................................ Miroculis (Ommaethus) mourei Tergos abdominales no parejos ni con líneas como en el caso anterior ................ 2 Tergos abdominales II-III y VI oscuros, IV-V más claros; tergo VI con una mancha mediana en forma de V (Figs. 165T-U, 167M) ............................................ 3 Tergos abdominales no como en el caso anterior ...................................................... 5 Tergos abdominales VI - VIII como en la Fig. 165U, branquias abdominales pequeñas y delgadas, proyecciones posterolaterales de medianas a reducidas (Fig. 167F) ......................................................................................Miroculis (Miroculis) marauiae Tergos abdominales diferentes al caso anterior, branquias abdominales variables, proyecciones posterolaterales de medianas a grandes (Figs. 167E-G) ................... 4 Branquias abdominales grandes, con lóbulos posterolaterales como en la Fig. 167G; filamentos caudales castaño-amarillentos, con bandas negruzcas en los segmentos V-IX .............................................................. Miroculis (Miroculis) brasiliaensis Branquias abdominales pequeñas, con lóbulos posterolaterales como en la Fig. 167E; filamentos caudales castaño-claros, sin bandas más oscuras .......................... ................................................................................................... Miroculis (Miroculis) fittkaui Branquias abdominales pequeñas, porción dorsal de las branquias II-VI no ensanchadas en la base formando un lóbulo (Fig. 167H) ........................................... ................................................................................................... Miroculis (Atroari) duckensis Branquias abdominales grandes, porción dorsal de las branquias II-VI con la base ensanchada formando un lóbulo (Figs. 167I-J) ........................................................... 6 Tergos abdominales V-VIII igualmente oscuros, o tergos IV-V un poco más claros (como en la Fig. 165V); membrana de las branquias grisácea .................................... ............................................................................................... Miroculis (Atroari) amazonicus Tergos abdominales V-VIII progresivamente más oscuros posteriormente (Fig. 167N), membrana de las branquias castaño-amarillentas ............................................ ........................................... Miroculis (Atroari) colombiensis, Miroculis (Miroculis) nebulosus

Subgenus Miroculis (Atroari) SAVAGE & PETERS Miroculis (Atroari) SAVAGE & PETERS, 1983: 549; SAVAGE, 1987a: 103. (Type-species: Miroculis (Atroari) duckensis SAVAGE & PETERS, original designation).

Taxonomy: Miroculis (Atroari) can be differentiated from the other subgenera of Miroculis by having the upper portion of male eyes not stalked, large, with 30-40 mediumsized facets in longest row, upper portions separated by a distance equal to 0,5-0,6 width of upper portion (Fig. 165N). Distribution: Northern Brazil, Colombia, and Venezuela.

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Miroculis (Atroari) amazonicus SAVAGE & PETERS Miroculis (Atroari) amazonicus SAVAGE & PETERS, 1983: 556 (male, nymph); SAVAGE, 1983b: 130.

Taxonomy: Miroculis (Atroari) amazonicus can be differentiated from the other species of the subgenus by the following combination of characters. In the imago: 1) Fore wings with dark brown transverse irregular submedian band and numerous maculae of same color (Fig. 164H); 2) penes from 1.7 and 1.9 times length of segment I of forceps, each lobe with ventral subapical lobe (Fig. 165E); and 3) abdominal color pattern as in Fig. 165V. Nymphs are distinguished from all species of Miroculis by the following combination of characters: 1) Anterolateral margins of labrum flattened; 2) abdominal gills large, with well developed posterolateral lobes (Fig. 167I); 3) color pattern of terga similar to that of imago (Fig. 165V) and 4) gill membranes translucent grayish, with blackish unbranched tracheae (Fig. 167I). Distribution and Biology: Northern Brazil. The adults fly in January. Miroculis (Atroari) colombiensis SAVAGE & PETERS Miroculis (Atroari) colombiensis SAVAGE & PETERS, 1983: 558 (nymph).

Taxonomy: Miroculis (Atroari) colombiensis is known only from nymphs, which can be separated from the other species of the subgenus by the following combination of characters: 1) Anterolateral margins of labrum flattened; 2) abdominal gills large, with well developed posterolateral lobes (Fig. 167J); 3) color pattern as in Fig. 167N; and 4) gill membranes yellowish brown, with blackish tracheae branched near base and apically (Fig. 167J). Nymphs are very similar to nymphs of Miroculis (M.) nebulosus but the male eyes do not have developing stalks. Distribution and Biology: Colombia. The nymphs were collected in a pond in a palm forest. Miroculis (Atroari) duckensis SAVAGE & PETERS Miroculis (Atroari) duckensis SAVAGE & PETERS, 1983: 554 (male, nymph); SAVAGE, 1983b: 130.

Taxonomy: Miroculis (Atroari) duckensis can be differentiated from the other species of the subgenus by the following combination of characters. In the imago: 1) Fore wing with light brown irregular transverse submedian band and numerous maculae of same color; 2) penes between 1.1 and 1.4 times length of segment I of forceps, each penis lobe tapered apically (Fig. 165F); and 3) color pattern of abdominal terga IV-VII as in Fig. 165V. Nymphs are distinguished from all species of Miroculis by the following combination of characters: 1) Anterolateral margins of labrum flattened; 2) abdominal gills small, with posterolateral lobes reduced to scarcely developed (Fig. 167H); 3) gill membrane translucent grayish (Fig. 167H); and 4) color pattern of abdominal terga IV-VII similar to imago (Fig. 165V). Distribution and Biology: Northern Brazil. The nymphs were collected in detritus in a small sandy bottomed stream with some exposed rocks. Adults emerged during June and July.

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Plate 166. Miroculis (Ommaethus) mourei, nymph. Fig. 166A, habitus. Figure modified from SAVAGE & PETERS, 1983.

Subgenus Miroculis (Miroculis) EDMUNDS Miroculis (Miroculis); SAVAGE & PETERS, 1983: 515; SAVAGE, 1987a: 99.

Taxonomy: Miroculis (Miroculis) can be separated from the other subgenera of Miroculis by the dorsal portion of the male eyes which are on a dorsally directed stalk, with 5-20 large facets in the longest row (Fig. 165M). Distribution: Northern Brazil, Colombia, French Guiana, Peru, Surinam, Venezuela and Trinidad.

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Miroculis (Miroculis) bicoloratus SAVAGE Miroculis (Miroculis) bicoloratus SAVAGE, 1987a: 99 (male).

Taxonomy: Miroculis (Miroculis) bicoloratus is known only from male imagos which can be differentiated from the other species of the subgenus by the following combination of characters: 1) Base of fore wings dark and apex light-colored or hyaline; 2) penes about 3 times length of segment I of forceps (Fig. 165A); 3) 8-10 facets in longest row of upper part of compound eyes (Fig. 165L); and 4) abdominal color pattern as in Fig. 165S. Distribution and Biology: Venezuela. The imagos of Miroculis (Miroculis) bicoloratus were collected during the nuptial flight, some 50 m from the closest stream around 15:00 h. Miroculis (Miroculis) brasiliaensis SAVAGE & PETERS Miroculis (Miroculis) brasiliaensis SAVAGE & PETERS, 1983: 535 (nymph).

Taxonomy: Miroculis (Miroculis) brasiliaensis was described only from nymphs, which can be separated from the other species by the following combination of characters: 1) Anterolateral margins of labrum flattened; 2) abdominal terga II-III and VI with dark pattern, IV-V and VII-VIII lighter, and tergum VI with a V-shaped median macula (Fig. 167M); and 3) abdominal gills large with well developed posterolateral lobes (Fig. 167G). Distribution and Biology: Northern Brazil. Apparently the adults emerge around the middle of November. Miroculis (Miroculis) fittkaui SAVAGE & PETERS Miroculis (Miroculis) fittkaui SAVAGE & PETERS, 1983: 529 (male, female, nymph); SAVAGE, 1983b: 130; SAVAGE, 1987a: 103 (male).

Taxonomy: Miroculis (Miroculis) fittkaui can be differentiated from the other species of the subgenus by the following combination of characters. In the male imago: 1) Wing membrane hyaline to light brown, with maculae around cross veins of fore wings (Fig. 164F); 2) penes tubular with apex excavated and 0.9-1.3 times length of segment I of forceps (Fig. 165B); 3) 58 facets in longest row of upper part of compound eyes; and 4) abdominal color pattern as in Fig. 165T. Nymphs are distinguished from all species of Miroculis by the following combination of characters: 1) Anterolateral margins of labrum flattened (Fig. 167A); 2) abdominal terga IIIII and VI dark, IV-V and VII-VIII lighter, and tergum VI with a V-shaped median macula; and 3) gills small with posterolateral lobes weakly developed (Fig. 167E). Distribution and Biology: Northern Brazil, Surinam, and Venezuela. Nymphs were collected on leaves or detritus in small streams with sandy bottoms and some exposed rocks. According to collection dates the adults fly between August and April. Miroculis (Miroculis) marauiae SAVAGE & PETERS Miroculis (Miroculis) marauiae SAVAGE & PETERS, 1983: 517 (male, female, nymph). Taxonomy: Miroculis (Miroculis) marauiae can be differentiated from the other species of the subgenus by the following combination of characters. In the male imago: 1) Wing

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membranes hyaline (Fig. 164D); 2) penes tubular with apex excavated, penes subequal in length to segment I of forceps (Fig. 165C); 3) 8-9 facets in longest row of upper part of compound eyes; and 4) abdominal color pattern as in Fig. 165U. Nymphs are distinguished from all species of Miroculis by the following combination of characters: 1) Anterolateral margins of labrum slightly flattened (Fig. 167B); 2) abdominal terga II-III and VI-VIII with dark pattern, IV-V lighter, and tergum VI with a V-shaped median macula; and 3) abdominal gills small and slender (Fig. 167F). Distribution and Biology: Northern Brazil. The nymphs were collected on leaf packs in rapid current zones in Amazonian streams. The adults fly in January and February.

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M N Plate 167. Miroculis, nymphs. Figs. 167A-D, mouthparts. 167A-C, Clypeus and labrum, d.v.: 167A, M. (M.) fittkaui; 167B, M. (M.)marauiae; 167C, M. (O.) mourei. 167D, left mandible, M. (M.) marauiae. 167EK, gill IV: 167E, M. (M.)fittkaui; 167F, M.(M.) marauiae; 167G, M. (M.) brasiliaensis; 167H, M. (A.) duckensis; 167I, M. (A.) amazonicus; 167J, M.(A.) colombiensis; 167K, M. (O.) mourei. 167L, tarsal claws, M. (M.) marauiae. 167M-N, abdominal color pattern: 167M, M. (M.) brasiliaensis (III-VII); 167N, M. (A.) colombiensis (IV-X). (D = denticle). Figs. 167A-N, modified from SAVAGE & PETERS, 1983.

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Miroculis (Miroculis) nebulosus SAVAGE Miroculis (Atroari) nebulosus SAVAGE, 1987: 104 (male). Miroculis (Miroculis) nebulosus; PETERS et al., in press (female, nymph).

Taxonomy: Miroculis (Miroculis) nebulosus, known from male and female imagos and nymphs, can be distinguished from the other species of the subgenus by the following combination of characters: 1) Wings of males with irregular transverse submedian dark brown bands and numerous maculae of the same color; in females, cross veins surrounded by maculae not forming heavy bands; 2) penes approximately 1.5 times length of segment I of forceps (Fig. 165G); 3) dorsal portion of compound eyes on a short stalk (Figs. 165O-P), with 11-14 facets in longest row of dorsal portion; and 4) abdominal color pattern as in Fig. 165W. Nymphs will key to Miroculis (Atroari) colombiensis but can be distinguished by the developing stalks of the male eyes. Distribution and Biology: Venezuela and Colombia. Males were collected at light in March in Venezuela near a small stream. Nymphs and adults were found in July at a very small Colombian stream with trickling water. There, nymphs were collected from leaf packs and rocks in pools. Miroculis (Miroculis) rossi EDMUNDS Miroculis rossi EDMUNDS, 1963: 34 (male); HUBBARD, 1982a: 266. Miroculis (Miroculis) rossi; SAVAGE & PETERS, 1983: 516 (male).

Taxonomy: Miroculis (Miroculis) rossi is based on three male imagos of which two are in poor condition. The male imago can be differentiated from the other species of the subgenus by the following combination of characters: 1) Wing membranes hyaline; 2) penes approximately 2 times length of segment I of forceps, each lobe tapering gradually toward apex (Fig. 165D) and with diagonal row of approximately 15 thick spines on basal bulge; and 3) 6 facets in longest row of upper portion of compound eyes (Fig. 165M). Distribution and Biology: Peru. The adults were collected in a forested zone in April. Subgenus Miroculis (Ommaethus) SAVAGE & PETERS Miroculis (Ommaethus) SAVAGE & PETERS, 1983: 559. (Type-species: Miroculis (Ommaethus) mourei SAVAGE & PETERS, original designation).

Taxonomy: Miroculis (Ommaethus) can be separated form the other subgenera of Miroculis by the following characters: 1) Upper portion of male eyes ovoid, not stalked, broadly separated by distance equal to width of dorsal portion of eye, and with 27-34 medium sized facets in longest row (Fig. 165Q); and 2) styliger plate with U-shaped depression apically (Figs. 165H, J). Also, fore wings of Ommaethus (Fig. 164J) are narrower (width/length ratio 0.41-0.43) than those of other subgenera (width/length ratio 0.43-0.51). Distribution: Southeastern Brazil and Northeastern Argentina.

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Miroculis (Ommaethus) froehlichi SAVAGE & PETERS Miroculis (Ommaethus) froehlichi SAVAGE & PETERS, 1983: 564 (male); SAVAGE, 1983b: 130.

Taxonomy: The imagos of Miroculis (Ommaethus) froehlichi can be differentiated from the other species of the subgenus by the following combination of characters: 1) Wing membrane light brown with dark brown maculae around cross veins; 2) penes about 0.85 times length of segment I of forceps (Figs. 165H-I); 3) dorsal portion of male compound eyes oval with 27-34 facets in longest row; 4) caudal filaments light yellowish; and 5) abdominal terga semihyaline. Distribution and Biology: Southeastern Brazil. The imagos emerged in October. Miroculis (Ommaethus) mourei SAVAGE & PETERS Miroculis (Ommaethus) mourei SAVAGE & PETERS, 1983: 561 (male, female, nymph); SAVAGE, 1983b: 130.

Taxonomy: Miroculis (Ommaethus) mourei can be separated from the other species of the subgenus by the following combination of characters. In the imago: 1) Wing membranes light brown, with dark brown maculae around cross veins (Fig. 164J); 2) penes about 0.9 times length of segment I of forceps (Figs. 165J-K); 3) dorsal portion of male compound eyes oval, with 27-29 facets in longest row (Fig. 165Q); 4) caudal filaments yellowish brown, basal 1/2 to 1/3 of segments with blackish brown band; and 5) abdominal color pattern as in Fig. 165X. Nymphs are distinguished from all species of Miroculis by the following combination of characters: 1) Anterolateral margins of labrum smoothly rounded to slightly flattened (Fig. 167C); 2) abdominal gills small with posterolateral lobes moderately developed; 3) gill membranes translucent grayish, with blackish, unbranched tracheae (Fig. 167K); and 4) color pattern as in Fig. 166A. Distribution and Biology: Southeastern Brazil. The nymphs were collected in various deep rivers with strong current, or shallow and slow streams. The adults emerged in February and March. Subgenus Miroculis (Yaruma) SAVAGE & PETERS Miroculis (Yaruma) SAVAGE & PETERS, 1983: 546. (Type-species: Miroculis (Yaruma) wandae SAVAGE & PETERS, original designation).

Taxonomy: Miroculis (Yaruma) can be differentiated from the other subgenera of Miroculis by the following character: male eyes on short broad stalk with 32-37 facets in longest row (Fig. 165R), eyes almost meeting dorsally, divided by distance 0.1-0.2 width of dorsal portion. In addition to the type species, another species is known only from a male imago with damaged genitalia; thus, the authors of the subgenus did not name it. Distribution: Northern Brazil. Miroculis (Yaruma) wandae SAVAGE & PETERS Miroculis (Yaruma) wandae SAVAGE & PETERS 1983: 546 (male).

Taxonomy: Miroculis (Yaruma) wandae is known only from male subimagos, which can be differentiated from the other unnamed species of the subgenus by the presence of

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submedian blackish brown maculae on the posterior part of abdominal terga V-VI in M. (Y.) wandae and their absence in the other species. Distribution and Biology: Northern Brazil. The imagos emerge in January and February. Genus Miroculitus SAVAGE & PETERS Miroculitus SAVAGE & PETERS, 1983: 566. (Type-species: Choroterpes emersoni NEEDHAM & MURPHY, original designation).

Taxonomy: The type species of Miroculitus, Choroterpes emersoni, was described by NEEDHAM & MURPHY (1924) based on adult specimens collected from a spider web. In 1946, TRAVER discovered that the type-series was a mixture of specimens from distinct species, and that the body of the holotype was lost; only the male genitalia and wings remained on slides. TRAVER (1947a) redescribed the species based on two male subimagos, which SAVAGE & PETERS (1983) were able to study before establishing a genus for this unique species. Adult Characteristics: 1) Fork of veins MA and MP of fore wings asymmetrical (Fig. 168A); 2) vein ICu1 of fore wings joined at base to CuA (Fig. 168A); 3) costal projection of hind wings well developed, acute, apex tent-shaped and projecting from costal margin basal to point where Sc meets C (Figs. 168B-C); 4) tarsal claws of a pair dissimilar, one apically hooked, the other blunt (Fig. 168D), similar on all legs; and 5) penes tubular, divided, without appendages and with rows of small basal spines (Fig. 168E). Nymphal Characteristics: Unknown. Distribution: Guyana. Biology: Nothing is known about the biology of the single species known of this genus, except that the adults fly in March. MP

MA

E A ICu1

CuA

CP

B

F D

C Plate 168. Miroculitus, imago. Figs. 168A-F: M. emersoni male. 168A, fore wing; 168B, hind wing; 168C, hind wing, detail; 168D, fore tarsal claw; 168E, apex of penes; 168F, styliger plate and forceps. (CP = Costal Projection). Figs. 168A-F modified from SAVAGE & PETERS, 1983.

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Miroculitus emersoni (NEEDHAM & MURPHY) Choroterpes emersoni NEEDHAM & MURPHY, 1924: 47 (male); TRAVER, 1946: 427; TRAVER, 1947a: 154 (male subimago). Choroterpes (Choroterpes) emersoni; HUBBARD, 1982a: 264. Miroculitus emersoni; SAVAGE & PETERS, 1983: 568; SAVAGE, 1983b: 128.

Taxonomy: The imagos of Miroculitus emersoni can be characterized by the following combination of characters: 1) Membrane of fore wings (Fig. 168A) hyaline; 2) middle and hind femora yellowish brown, with submedian and apical blackish brown maculae; and 3) penes dark brown, with rows of basal spines (Fig. 168E); forceps as in Fig. 168F. Distribution: Guyana. Genus Needhamella DOMÍNGUEZ & FLOWERS Needhamella DOMÍNGUEZ & FLOWERS, 1989: 568; FLOWERS & DOMÍNGUEZ, 1991: 51; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 57; DOMÍNGUEZ, FERREIRA & NIETO, 2001: 316; DOMÍNGUEZ & CUEZZO, 2002: 158. (Type-species: Thraulus ehrhardti ULMER, original designation).

Taxonomy: Needhamella is one of the genera included in the Hermanella generic complex by DOMÍNGUEZ & FLOWERS (1989). The relationships of this genus with the other MP

MA

E

A Sc

CP

B

C

Pp SPp

D F

Plate 169. Needhamella ehrhardti, imago. Figs. 169A-F: 169A, fore wing; 169B, hind wing; 169C, hind wing, detail; 169D, prosternum; 169E, genitalia, v.v.; 169F, abdominal terga V-VII. (CP = Costal Projection; Pp = Penes projections; SPp = Styliger Plate projections). Figs. 169A-F, modified from DOMÍNGUEZ & FLOWERS, 1989.

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components of the group were analyzed by FLOWERS & DOMÍNGUEZ (1991), DOMÍNGUEZ et al., (2001) and DOMÍNGUEZ & CUEZZO (2002). Adult Characteristic: 1) Fork of veins MA and MP of fore wings asymmetrical (Fig. 169A); 2) costal projection of hind wings sharply pointed (Figs. 169B-C); 3) vein Sc of hind wings terminating in cross vein; 4) tarsal claws of a pair dissimilar, one apically pointed, other blunt; 5) penes (Fig. 169E) divided in apical 1/2, ventral projections thin and curved, each penis lobe widened laterally in basal 1/3; 6) styliger plate with projections near base of forceps (Fig. 169E); 7) prosternum relatively wide, narrower in apical 1/2 (Fig. 169D); and 6) female sternum IX apically cleft. Nymphal Characteristics (general aspect as in Fig. 170A): 1) Labrum as wide as head, with setae as in Fig. 170B; 2) outer margin of mandible angular (Fig. 170C); 3) maxilla with prominent tusk on inner apical margin (Fig. 170D); 4) submentum with 10-12 setae laterally (Fig. 170E); 5) apical denticle of tarsal claws much larger than preceding denticles (Fig. 170F), and claw with two small apical denticles; and 6) plate-like gills terminated in a finger-like process, present on abdominal segments I-VII (Fig. 170G). Distribution: Argentina, Brazil, and Uruguay. Biology: Most of the adults collected have been attracted to light. The nymphs were collected along the margins of small streams and rivers of medium size, under rocks or in the leaf packs. Needhamella ehrhardti (ULMER) Thraulus ehrhardti ULMER, 1920a: 28 (male, female); ULMER, 1920c: 117; NEEDHAM & MURPHY, 1924: 44; ULMER, 1943: 33 (male, female). Traverella ehrhardti; EDMUNDS, 1950b: 551; TRAVER, 1959b: Figs. 15, 18; THEW, 1960b: 123; TRAVER, 1963: 28 (male, female); ALLEN, 1973a: 1287; HUBBARD, 1982a: 267. Hermanella sp.; EDMUNDS, JENSEN & BERNER, 1976: Figs. 140, 310. Needhamella ehrhardti; DOMÍNGUEZ & FLOWERS, 1989: 570 (male, female, nymph, egg); HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 216; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 58 (male, nymph).

Taxonomy: Needhamella ehrhardti was described in the genus Thraulus by ULMER (1920a) for males and females from Brazil (Estado de Santa Catarina); later, based on other specimens ULMER (1943) provided additional data, especially on the genitalia. In 1950, EDMUNDS transferred this species, and Thraulus maculipennis ULMER to the genus Traverella and EDMUNDS et al. (1976) included figures of the genitalia and gills of this species (as Hermanella sp.). DOMÍNGUEZ & FLOWERS (1989) designated this the type species of genus Needhamella. Needhamella ehrhardti can be characterized by the following combination of characters. In the imago: 1) All cross veins of fore wings (Fig. 169A) lacking shading; 2) each lobe of penes with thin projection directed basally (Fig. 169E); and 3) abdominal color pattern as in Fig. 169F. In the nymph: 1) Gills grayish black, with tracheae darker; and 2) abdominal terga brownish orange, with blackish pattern as in male imago (Fig. 169F). Distribution and Biology: As in generic discussion.

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B

C

T

A

D

D

E

F

G

Plate 170. Needhamella ehrhardti, nymph. Figs. 170A-F: 170A, habitus; 170B, labrum, d. v.; 170C, left mandible; 170D, maxilla; 170E, labium, right ventral, left dorsal; 170F, fore tarsal claw; 170G, gill I. (D = denticle; T = tusk). Fig. 170A, drawing by A. Dupuy. Figs. 170B-G, modified from DOMÍNGUEZ & FLOWERS, 1989.

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Genus Nousia NAVÁS Nousia NAVÁS, 1918: 213; NAVÁS, 1925: 308; ULMER, 1933: 201; TRAVER, 1946: 420; PETERS & EDMUNDS, 1972: 1412; PESCADOR & PETERS, 1985: 91; SUTER, 1986: 347; SAVAGE, 1987b: 207; CAMPBELL & SUTER, 1988: 270; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 59. (Type-species: Nousia delicata NAVÁS, original designation). Atalonella NEEDHAM & MURPHY, 1924: 35; LESTAGE, 1931b: 53 [partim]; TRAVER, 1946: 420; HARKER, 1950: 30; HARKER, 1954: 243; PETERS & EDMUNDS, 1972: 1411; TSUI & PETERS, 1975: 540; PESCADOR & PETERS, 1980b: 43. (Type-species: Atalonella ophis NEEDHAM & MURPHY, subsequent designation by PETERS & EDMUNDS, 1972).

Taxonomy: The genus Nousia was established by NAVÁS (1918) for the species N. delicata from Chile. In 1924, NEEDHAM & MURPHY established Atalonella for A. ophis from Chile and A. fusca from Australia, but did not indicate which was the type species of the genus. PETERS & EDMUNDS (1972) subsequently designated A. ophis as the type species of Atalonella. NAVÁS (1925) synonymized Atalonella with Nousia, but TRAVER (1946) was unconvinced because NAVÁS (1918) noted the absence of an intercalary in the median fork of the hind wings of Nousia. PESCADOR & PETERS (1985), studied material determined by NAVÁS, and revised the taxonomy of the genus and synonymized the genus Atalonella. Adult Characteristics: 1) Fork of vein MA of fore wings symmetrical (Fig. 171A) to slightly asymmetrical; 2) vein MP2 of fore wings moderately recurved, attached to MP1 at base with cross vein (Fig. 171A); 3) hind wings without costal projection (Fig. 171B); 4) length of vein Sc of hind wings 3/4 maximum length of wing (Fig. 171B); 5) claws of a pair similar, each apically hooked with an opposing hook (Fig. 171C); 6) penis lobes fused in basal 2/3, remainder divided and tubular (Figs. 171D-J), each lobe with short sclerotized process on inner margin; 7) female sternum IX apically cleft (Figs. 171K-L). Nymphal Characteristics (Fig. 172A): 1) Clypeus slightly narrower than labrum, clypeus with lateral margins parallel (Fig. 173A); 2) anteromedian emargination of labrum broad, U-shaped, with 5 irregular denticles (Fig. 173B); 3) outer margin of mandibles smoothly curved with few fine long setae on basal half, and median seta tuft (Fig. 173C); 4) segment II of maxillary palpi approximately 4/5 length of segment I, segments II and III subequal in length (Fig. 173D); 5) submentum with long lateral setae (Fig. 173E); 6) tarsal claws with denticles progressively larger apically or apical denticle much larger (Figs. 173FH); 7) posterolateral spines on abdominal segments VI-IX; and 8) abdominal gills I-VII similar, with well developed dorsal and ventral portions tapered apically (Figs. 173I-L). Distribution: Chile (Magallanes Province (ca. 53ºS, XII region) north to Choapa prov. (31°37S; 71°9W, IV region), with easternmost record in Argentina (Río Negro Province (ca. 70ºW)). Biology: The nymphs of this genus inhabit a wide variety of streams, rivers and lakes of Southern Argentina and Chile. The individuals are normally found under rocks and logs, covered with algae or not, or among debris. Adults were observed flying from October through May. The males were observed swarming in the afternoon and evening, in companies of approximately 30 individuals. They can be found at a wide range of elevations (from sea level to approximately 2500 m).

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MA

A

Sc ICu1

MP MP1 CuA 2

D

E

H

I

C

B

F G

K

J

M

R

O

N

S

T

L

P

U

Q

V

W

Plate 171. Nousia, imagos. Figs. 171A-W. Figs. 171A-C, N. delicata: 171A, fore wing; 171B, hind wing; 171C, male fore claw. Figs. 171D-J, penes, v.v.: 171D, N. bella; 171E, N. crena; 171F-G, N. delicata: 171F, general view of genitalia; 171G, penes; 171H, N. grandis; 171I, N. maculata; 171J, N. minor. Figs. 171KL, apex of female sternum IX: 171K, N. delicata; 171L, N. grandis. Figs. 171M-W, abdominal terga V-VI: 171M, N. bella; 171N-O, N. crena: 171N male, 171O female; 171P-Q, N. delicata: 171P male, 171Q female; 171R-S, N. grandis: 171R male, 171S, female; 171T-U, N. maculata: 171T male; 171U female; 171V-W, N. minor: 171V male; 171W female. Figs. 171A-W modified from PESCADOR & PETERS, 1985.

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TO

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SOUTH AMERICAN NOUSIA

Imagos 1 – 2(1) – 3(2) – 4(3)

5(4) –

Maculae on abdominal terga greatly reduced or absent (Figs. 171R-S); apical cleft of female sternum IX shallow, approximately 1/7 maximum width of sternum (Fig. 171L); penes as in Fig. 171H ............................................................ Nousia grandis Maculae on abdominal terga extensive (Figs. 171M-Q, T-W); apical cleft of female sternum IX deep, approximately 1/3 maximum width of sternum (Fig. 171K); penes not as above (Figs. 171D-G, I-L) ........................................................................ 2 Body length less than 6.5 mm; maculae on abdominal terga as in Figs. 171V-W; penes as in Fig. 171J ...................................................................................... Nousia minor Body length more than 6.5 mm; maculae on abdominal terga as in Figs. 171M-Q, T-U; penes as in Figs. 171D-G, I .................................................................................... 3 Eyes of male separated on meson of head; maculae on abdominal terga as in Figs. 171P-Q .......................................................................................................... Nousia delicata Eyes of male meet on meson of head; maculae on abdominal terga as in Figs. 171M-O, T-U ....................................................................................................................... 4 Penes with weakly developed mid-lateral projection (Fig. 171I); maculae on abdominal terga as in Figs. 171T-U ...................................................... Nousia maculata Penes with well developed mid-lateral projection (Figs. 171D-E); maculae on abdominal terga as in Figs. 171M-O .............................................................................. 5 Maculae on abdominal terga as in Fig. 171M; penes with broad mid-lateral projection (Fig. 171D) ..................................................................................... Nousia bella Maculae on abdominal terga as in Figs. 171N-O; penes with narrow mid-lateral projection (Fig. 171E) .................................................................................... Nousia crena

Nymphs 1 – 2(1) – 3(2) – 4(3) –

Abdominal gills slender (Fig. 173L); tibiae with postmedian brown band .............. ............................................................................................................................ Nousia minor Abdominal gills moderately broad (Figs. 173I-K); tibiae without postmedian band ................................................................................................................................................ 2 Maculae on abdominal terga reduced (Figs. 171R-S); lateral margins of submentum each with less than 10 setae ................................................ Nousia grandis Maculae on abdominal terga pronounced; lateral margins of submentum each with more than 10 setae ................................................................................................... 3 Abdominal gills dark gray to black (Fig. 173I); abdominal sterna dark brown, lateral margins pale yellow ............................................................................. Nousia bella Abdominal gills cloudy white; abdominal sterna yellow ............................................ 4 Maximum width of hind femora less than 1/3 maximum length; main tracheal trunk of abdominal gills with a few weakly developed branches (Fig. 173K) ........ ....................................................................................................................... Nousia maculata Maximum width of hind femora more than 1/3 maximum length; main tracheal trunk of abdominal gills with numerous well developed branches (Fig. 173J) ................................................................................................................................................ 5

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CLAVES

Apical denticles on claws distinctly narrower than maximum width of apex of claw (Fig. 173G); maculae on abdominal terga as in Figs. 171P-Q ................ Nousia delicata Apical denticles on claws broader than maximum width of apex of claw (Fig. 173F); maculae on abdominal terga not as above (Figs. 171N-O) ...... Nousia crena PARA LOS

NOUSIA

SUDAMERICANOS

Imagos 1

– 2(1) – 3(2) – 4(3) – 5(4) –

Manchas en tergos abdominales muy reducidos o ausente (Figs. 171R-S); hendidura apical del esterno IX de hembra poco profunda, aproximadamente 1/7 del ancho máximo del esterno (Fig. 171L); penes como en la Fig. 171H ............... .......................................................................................................................... Nousia grandis Manchas extensivas en tergos abdominales (Figs. 171M-Q, T-W); hendidura apical del esterno IX de la hembra profunda, aproximadamente 1/3 del ancho máximo del esterno (Fig. 171K); penes no como arriba (Figs. 171D-G, I-L) ...................... 2 Longitud del cuerpo menos que 6,5 mm; manchas en tergos abdominales como en Figs. 171V-W; penes como en Fig. 171J .............................................. Nousia minor Longitud del cuerpo más de 6,5 mm; manchas en tergos abdominales como en Figs. 171M-Q, T-U; penes como en Figs. 171D-G, I ................................................. 3 Ojos del macho separados en el dorso de la cabeza; manchas en tergos abdominales como en Figs. 171P-Q ....................................................... Nousia delicata Ojos del macho unidos sobre el dorso de la cabeza; manchas en tergos abdominales como en Figs. 171M-O, T-U .................................................................... 4 Penes con la proyección mediolateral débilmente desarrollada (Fig. 171I); manchas en tergos abdominales como en Figs. 171T-U .................. Nousia maculata Penes con la proyección mediolateral bien desarrollada (Figs. 171D-E); manchas en tergos abdominales como en Figs. 171M-O ........................................................... 5 Manchas en tergos abdominales como en Fig. 171M; penes con la proyección mediolateral ancha (Fig. 171D) ..................................................................... Nousia bella Manchas en tergos abdominales como en Figs. 171N-O; penes con la proyección mediolateral angosta (Fig. 171E) ................................................................. Nousia crena

Ninfas 1 – 2(1) – 3(2) –

Branquias abdominales delgadas (Fig. 173L); tibias con banda postmediana castaña .............................................................................................................. Nousia minor Branquias abdominales moderadamente anchas (Figs. 173I-K); tibias sin banda postmediana ........................................................................................................................ 2 Manchas en tergos abdominales reducidas (Figs. 171R-S); cada margen lateral del submentón con menos de 10 setas .......................................................... Nousia grandis Manchas en tergos abdominales marcadas; cada margen lateral del submentón con más de 10 setas ........................................................................................................... 3 Branquias abdominales gris-oscuras a negras (Fig. 173I); esternos abdominales castaño-oscuros, márgenes laterales amarillo-palido ................................ Nousia bella Branquias abdominales blancuzcas; esternos abdominales amarillos ..................... 4

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A

Plate 172. Nousia delicata. Fig. 172A, nymphal habitus. Figure modified from P ESCADOR & P ETERS , 1985.

458

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Ancho máximo de fémur posterior menos de 1/3 de la longitud máxima; traquea principal de las branquias abdominales con pocas ramas débilmente desarrolladas (Fig. 173K) .................................................................................................. Nousia maculata Ancho máximo de fémur posterior más de 1/3 de la longitud máxima; traquea principal de las branquias abdominales con numerosas ramas bien desarrolladas (Fig. 173J) ............................................................................................................................. 5 Dentículos apicales de las uñas mucho más delgadas que el ancho máximo del ápice de la uña (Fig. 173G); manchas en tergos abdominales como en Figs. 171P-Q ......................................................................................................................... Nousia delicata Dentículos apicales de las uñas más anchos que el ancho máximo del ápice de la uña (Fig. 173F); manchas en tergos abdominales no como en el caso anterior (Figs. 171N-O) ................................................................................................. Nousia crena

Nousia bella PESCADOR & PETERS Nousia bella PESCADOR & PETERS, 1985: 117 (male, female, nymph). Nousia (Nousia) bella; CAMPBELL & SUTER, 1988: 270; HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 217; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 61 (male).

Taxonomy: Nousia bella was described by PESCADOR & PETERS (1985) from reared adults and associated nymphs from Río Los Morongos (35º07’S; 71º09’W), Curico Province, VII region, Chile. It can be distinguished from the other species of the genus by the following combination of characters. In the imago: 1) Male eyes meeting on meson of head; 2) extensive maculae on abdominal terga, with pattern as in Fig. 171M; 3) penes with well developed midlateral projections (Fig. 171D); and 4) female sternum IX deeply cleft apically, cleft approximately 1/3 maximum width of sternum (as in Fig. 171K). In the nymph: 1) Maximum width of hind femora more than 1/3 its length; 2) lateral margins of submentum with 10-12 long spines; 3) apical denticle of tarsal claws broader than maximum width of apex of claw (as in Fig. 173F); and 4) abdominal gills black, with dorsal and ventral portions moderately broad, and main tracheal trunk with well developed branches (Fig. 173I). Distribution and Biology: Rio Illapel, Choapa Province (31°37’S; 71°9’W, IV region), southward to Dalcahue (42°21’S; 73°42’W) Chiloe Province, X region, Chile, and eastward to Neuquen Province southward to Chubut Province, Argentina. The nymphs were collected on algae-laden rocks of various sizes found in streams and rivers from 200 to 2000 m in altitude. Collection records indicate that the nymphs occur from September through January, and the adults from November through February. Nousia crena PESCADOR & PETERS Nousia crena PESCADOR & PETERS, 1985: 105 (male, female, nymph). Nousia (Nousia) crena; CAMPBELL & SUTER, 1988: 270; HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 217; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 62 (male, nymph).

Taxonomy: PESCADOR & PETERS (1985) described N. crena from reared adults and nymphs. They briefly discussed intraspecific color variations which occur in both the nymphs and adults throughout the species’ geographic range.

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Nousia crena can be distinguished from the other species of the genus by the following combination of characters. In the imago: 1) Male eyes meeting on meson of head; 2) maculae on abdominal terga pronounced and extensive (Figs. 171N-O); 3) penes with narrow, moderately developed mid-lateral projections (Fig. 171E); and 4) female sternum IX deeply cleft apically, cleft slightly deeper than 1/3 maximum width of sternum (Fig. 171K). In the nymph: 1) Maximum width of hind femora more than 1/3 maximum length; 2) lateral margins of submentum with 10-12 long setae; 3) apical denticle on tarsal claws broader than maximum width of apex of claw (Fig. 173F); 4) maculae on abdominal terga pronounced and extensive, with pattern as in Figs. 171N-O; and 5) gills cloudy white with dorsal and ventral portions moderately broad, main tracheal trunk with several well developed lateral branches (Fig. 173J). Distribution and Biology: Río Teno 34°52’S; 71°10’W, Curicó prov. VII region south to Dalcahue 42°21’S; 73°42’W, Chiloe Province, X region, Chile, and Neuquen Province south to Chubut Province, Argentina. The nymphs occur in a wide variety of habitats, from small temporary streams to medium-sized rivers, from sea level to 1800 m. The nymphs have been collected from September through March, and adults from September through April. Mating flight was observed in the afternoon, the swarm composed of approximately 30 males. Nousia delicata NAVÁS Nousia delicata NAVÁS, 1918: 213 (male); ULMER, 1920c: 116; NAVÁS, 1925: 308 (male); NAVÁS, 1930d: 359; LESTAGE, 1931b: 54; NAVÁS, 1933a: 233; TRAVER, 1946: 421; PETERS & EDMUNDS, 1972: 1412; HUBBARD, 1982a: 263; ALBA-TERCEDOR & PETERS, 1985: 222; PESCADOR & PETERS, 1985: 96 (male, female, nymph, egg). Nousia (Nousia) delicata; CAMPBELL & SUTER, 1988: 270; HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 217; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 62 (male, female, nymph). Atalonella ophis NEEDHAM & MURPHY, 1924: 36; NAVÁS, 1925: 310; LESTAGE, 1931b: 52; TRAVER, 1946: 420 (male); DEMOULIN, 1955e: 13 (male, female); TRAVER, 1959b: Fig. 23 (male); PETERS & EDMUNDS, 1972: 1411. Nousia ophis; NAVÁS, 1925: 310. Atonella [sic] ophis; HUBBARD, 1982a: 266.

Taxonomy: PESCADOR & PETERS (1985) reviewed the taxonomy of N. delicata and synonymized the species Atalonella ophis. Nousia delicata can be distinguished from the other species of the genus by the following combination of characters. In the imago: 1) Male eyes separated on meson of head; 2) maculae on abdominal terga extensive, with pattern as in Figs. 171P-Q; 3) penes with weakly developed mid-lateral projections (Figs. 171F-G); and 4) female sternum IX deeply cleft apically, cleft approximately 1/3 maximum width of sternum (Fig. 171K). In the nymph: 1) Maximum width of hind femora more than 1/3 maximum length; 2) lateral margins of submentum with 15-18 long setae; 3) apical denticles on tarsal claws narrower than maximum width of apex of claw (Fig. 173G); 4) maculae on abdominal terga extensive with pattern as in Figs. 171P-Q; and 5) abdominal gills cloudy white, with dorsal and ventral portions moderately broad, main tracheal trunk with several well developed branches (as in Fig. 173J).

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Distribution and Biology: Nousia delicata is one of the most widely distributed species of the genus, occurring from both the western and eastern slopes of the southern Andes from approximately 30ºS (IV region) to the Magellan strait (XII region). The nymphs live in a wide variety of habitats, from temporary to permanent streams, rivers

A

B

C

D

D

D

E

F

G

H

SM

I

J

K

L

Plate 173. Nousia, nymphs. Figs. 173A-E, mouthparts of N. delicata: 173A, labrum, d.v.; 173B, anteromedian emargination of labrum; 173C, left mandible; 173D, maxilla; 173E, labium (left dorsal, right ventral). 173F-H, Fore claws: 173F, N. crena; 173G, N. delicata; 173H, N. maculata. 173I-L, abdominal gill IV: 173I, N. bella; 173J, N. delicata; 173K, N. maculata; 173L, N. minor. (SM = Submentum; D = Denticle). Figs. 173A-L modified from PESCADOR & PETERS, 1985.

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and lakes, from 200 to 2500 m of elevation. The nymphs have been collected from October through April, and the adults from October through May. A few nymphs were found parasitized by a chironomid, Symbiocladius sp. Nousia grandis (DEMOULIN) Atalonella grandis DEMOULIN, 1955e: 21 (female); PETERS & EDMUNDS, 1972: 1411; HUBBARD, 1982a: 263. Nousia grandis; PESCADOR & PETERS, 1985: 109 (male, female, nymph). Nousia (Nousia) grandis; CAMPBELL & SUTER, 1988: 270; HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 217; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 63 (male, female).

Taxonomy: Nousia grandis can be distinguished from the other species of the genus by the following combination of characters. In the imago: 1) Male eyes meeting on meson of head; 2) maculae on abdominal terga absent or greatly reduced (Figs. 171R-S); 3) penes with well developed mid-lateral projections (Fig. 171H), lobes divided apically; and 4) female sternum IX with shallow cleft, cleft slightly deeper than 1/7 maximum width of sternum (Fig. 171L). In the nymph: 1) Maximum width of hind femora less than 1/3 maximum length; 2) lateral margins of submentum with 8-9 long setae; 3) apical denticles on tarsal claws distinctly narrower than maximum width of apex of claw (as in Fig. 173G); 4) pattern of maculae on abdominal terga (if present) similar to Figs. 171R-S; and 5) abdominal gills cloudy white, with dorsal and ventral portions moderately broad, main tracheal trunk with few to several well developed branches (as in Fig. 173J). Distribution and Biology: Widespread, occurring from Cordillera Province, ca 33°S (metropolitan Region), southward to Puerto Eden (49º20’S), Magallanes Province, XII region, Chile, eastward to Neuquen Province southward to Rio Negro Province, Argentina. The nymphs occur in shallow areas of streams with abundant trapped debris and submerged logs, at elevations from 200 m to 1100 m. The nymphs have been collected from September through February and the adults from December through March. A few nymphs have been found parasitized by the chironomid, Symbiocladius sp. Nousia maculata (DEMOULIN) Atalonella maculata DEMOULIN, 1955e: 18 (male, female); PETERS & EDMUNDS, 1972: 1411; HUBBARD, 1982a: 263. Nousia maculata; PESCADOR & PETERS, 1985: 101 (male, female, nymph). Nousia (Nousia) maculata; CAMPBELL & SUTER, 1988: 270; HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 218; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 64 (male, nymph).

Taxonomy: Nousia maculata was originally described by DEMOULIN (1955) from male and female imagos. The species was later redescribed by PESCADOR & PETERS (1985) based on the holotype, reared adults, and the nymph. Nousia maculata can be distinguished from the other species of the genus by the following combination of characters. In the imago: 1) Male eyes meeting on meson of head; 2) maculae on abdominal terga often with pattern as in Figs. 171T-U, but sometimes less extensive; 3) penes with weakly developed mid-lateral projection (Fig. 171I); and 4)

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female sternum IX deeply cleft apically, approximately 1/3 maximum width of sternum (Fig. 171K). In the nymph: 1) Maximum width of hind femora less than 1/3 maximum length; 2) lateral margins of submentum with 10-12 long setae; 3) apical denticles on tarsal claws as broad as maximum width of apex of claw (Fig. 173H); 4) maculae on abdominal terga with pattern similar to Figs. 171T-U, or pattern reduced; and 5) abdominal gills cloudy white, dorsal and ventral portions moderately broad, main tracheal trunk with few weakly developed branches (Fig. 173K). Distribution and Biology: Magallanes Province (ca 53ºS, XII region) north to Petorca Province (32°33’S) V region, Chile, and Neuquen south to Río Negro Province, Argentina. The nymphs of N. maculata are rheophilic and have been collected mostly under rocks and stones in shallow areas of streams and rivers from 200 to 1800 m of elevation. The nymphs have been collected in September through March, and the adults in November through April. Nousia minor (DEMOULIN) Atalonella minor DEMOULIN, 1955e: 16 (male, female); PETERS & EDMUNDS, 1972: 1411; HUBBARD, 1982a: 263. Nousia minor; PESCADOR & PETERS, 1985: 114 (male, female, nymph). Nousia (Nousia) minor; CAMPBELL & SUTER, 1988: 270; HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 218; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 65 (male, nymph).

Taxonomy: Nousia minor was first described by DEMOULIN from male and female imagos and subimagos collected from Río Diguillin (36°52’S) Ñuble Prov. VIII region, Chile. PESCADOR & PETERS (1985) redescribed the species based on reared adults and described the nymph for the first time. Nousia minor can be distinguished from the other species of the genus by the following combination of characters. In the imago: 1) Male eyes meeting on meson of head; 2) extensive dark brown maculae on abdominal terga similar to Figs. 171V-W; 3) penes with weakly developed mid-lateral projection (Fig. 171J); and 4) female sternum IX deeply cleft apically (Fig. 171K). In the nymph: 1) Maximum width of hind femora less than 1/3 maximum length; 2) lateral margins of submentum with 10-11 setae; 3) apical denticle on tarsal claws broader than maximum width of apex of claw (as in Fig. 173F); 4) maculae on abdominal terga extensive, with pattern similar to Figs. 171V-W; and 5) abdominal gills slender with main tracheal trunk with few to no weakly developed branches (Fig. 173L). Distribution and Biology: Maullín (41°37’S), Llanquihue Province, X region, north to Estero La Jaula (ca. 36ºS), Curico Province, VII region, Chile, and east to Neuquen and Chubut Provinces, Argentina. The rheophilic nymphs of N. minor were mostly found underneath small boulders or big stones near the edge of fairly large streams and rivers from 250 to 1000 m elevation. A few nymphs also were collected among emergent vegetation near the shoreline of lakes. The nymphs of N. minor have been collected from November through February, and the adults from October through April (PESCADOR & PETERS, 1985).

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Genus Paramaka SAVAGE & DOMÍNGUEZ Thraulus [partim]; SPIETH, 1943: 10. Hermanella [partim]; DEMOULIN 1966a: 12. Paramaka SAVAGE & DOMÍNGUEZ, 1992: 244; BLANCO-BELMONTE, RUAISE & PETERS, 2003: 117. (Type-species: Thraulus convexus SPIETH, original designation).

Taxonomy: Thraulus convexus SPIETH was described from two male imagos from Surinam (SPIETH, 1943) which TRAVER (1960) transferred to Homothraulus. More recently, based on a phylogenetic study (FLOWERS & DOMÍNGUEZ, 1989), SAVAGE & DOMÍNGUEZ (1992) determined that this species belonged to the Hermanella generic complex and established for it the new genus Paramaka. BLANCO-BELMONTE et al. (2003) were able to rear and associate Paramaka convexa with the nymph described as Hermanella sp. 2 by DEMOULIN (1966). Adult Characteristics: 1) Forks of veins MA and MP of fore wings asymmetrical (Fig. 174A); 2) fore wings with slanting cross vein above fork of MA; 3) vein ICu1 attached at base to CuA by a cross vein; 4) hind wings with costal projection well developed, acute (Figs. 174B-C); 5) vein Sc of hind wings terminating in cross vein; 6) vein MP of hind wings not forked; 7) claws of a pair dissimilar, one hooked apically, other pad-like (Fig. 174D); 8) penes divided on apical 2/3, each lobe with a mesal spinelike projection directed anteromesally (Fig. 174E); and 9) styliger plate projecting dorsally between penes (Fig. 174F). Nymphal Characteristics: 1) Labrum as wide as head, with setae as in Fig. 174H; 2) outer margin of mandible forming almost a 90° angle (Fig. 174J); 3) submentum reduced, with few lateral setae (Fig. 174I); 4) subapical denticle of tarsal claws much larger than other denticles, with a few small apical denticles (Fig. 174K); 5) posterolateral projections on abdominal segments VIII-IX; and 6) abdominal gills (Fig. 174L) platelike, without terminal filament on I, a short filament often present on dorsal portion of III-VI, gills vestigial on VII. Distribution: Surinam (Gran Soela) and Brazil (Pará), French Guiana (Sinnamary river), Venezuela (Bolivar) and Colombia. Biology: The nymphs of this species were collected from sticks and leaves along margins of a small creek (2-2.5 m wide and 0.3-1 m deep). The males were collected in June and August. Paramaka convexa (SPIETH) Thraulus convexus SPIETH, 1943: 10. Homothraulus convexus; TRAVER, 1960b: 73; HUBBARD, 1982a: 265. Hermanella sp. 2. DEMOULIN, 1966a: 12. Paramaka convexa; SAVAGE & DOMÍNGUEZ, 1992: 244; BLANCO-BELMONTE et al., 2003: 119.

Taxonomy: This is the only species described in the genus, and can be characterized by the following characters: 1) Membrane of fore and hind wings hyaline with faint yellow tinge, longitudinal and cross veins of fore wings brown and distinct; 2) abdominal segments II-VI translucent yellow, I and VII-X dark, color pattern is as in

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MP MA

Scv

A

Sc

ICu1 CuA

CP

B

D

C MP

E F G

I H D

J

K

L

Plate 174. Paramaka convexa. Figs. 174A-L. 174A-G, male imago: 174A, fore wing; 174B, hind wing; 174C, hind wing, detail; 174D, fore tarsal claw; 174E-F, genitalia, v.v. showing slightly different angles; 174G, abdominal terga I-X. 174H-L, nymph: 174H, labro, v. v.; 174I, labium, right ventral, left dorsal; 174J, left mandible; 174K, fore claw; 174L, gill IV. (Scv = slanting cross vein; CP = Costal projection; D = denticle). Figs. 174A-F modified from SAVAGE & DOMÍNGUEZ, 1992; 174G-L from BLANCO-BELMONTE et al., 2003.

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Fig. 174G; and 3) genitalia with ventral spine-like projections directed anteromesally as in Figs. 174E-F. In the nymph: 1) Gills light brown to light gray basally, progressively darker distally to about 3/4 of distance from base, pale apically; main tracheal trunk blackish; 2) abdominal terga brown, a darker mark on posterolateral corners of terga IIVII; and 3) lateral margins of labrum flared as in Fig. 174H. Distribution and Biology: As in generic discussion. Genus Penaphlebia PETERS & EDMUNDS Atalophlebia EATON, 1884: 91 [partim], NEEDHAM & MURPHY, 1924: 34 [partim]; TRAVER, 1946: 419 [partim]; TRAVER, 1959b: 3; DEMOULIN, 1955b: 9 [partim]; DEMOULIN, 1955e: 1 [partim]; TRAVER, 1959b: 3 [partim]; THEW, 1960b: 130 [partim]. Penaphlebia PETERS & EDMUNDS, 1972: 1399; TSUI & PETERS, 1975: 548; PESCADOR & PETERS, 1980b: 45; PESCADOR & PETERS, 1991: 2. (Type-species: Atalophlebia chilensis EATON, original designation).

Taxonomy: PETERS & EDMUNDS (1972) established this genus and transferred to it four recognizable Chilean species of Atalophlebia. NAVÁS (1928, 1930d, 1934, 1936a) described seven other species of Atalophlebia from Chile which Pescador & Peters (1991) transferred to Penaphlebia but listed as nomina dubia because types had been destroyed. Pescador & Peters (1991) revision of the genus resulted in the establishment of two subgenera; namely, Penaphlebia (Megalophlebia) and Penaphlebia (Penaphlebia). Adult Characteristics: 1) Fork of vein MA of fore wings symmetrical (Figs. 175C, E); 2) vein Sc of hind wings 9/10 length of hind wings (Figs. 175D, F-G) hind wings without costal projection; 3) claws of a pair similar, apically hooked, with an opposing hook (Fig. 175H); 4) penis lobes divided and tubular (Figs. 175J-O), each lobe with a subapical, ventral spine (Figs. 175M-O); and 5) female sternum IX deeply cleft apically (Fig. 175I). Nymphal Characteristics (general aspect as in Fig. 176A): 1) Clypeus as wide as labrum (Fig. 177A); 2) length of labrum approximately 1/2 maximum width (Fig. 177A); 3) outer margin of mandibles slightly curved with long, thick setae on basal half (Fig. 177B-C); 4) maxilla as in Fig. 177D; 5) hypopharynx as in Fig. 177E; 6) glossae strongly curved over ventrally, submentum with long lateral setae (Fig. 177F); 7) segment III of labial palpi approximately 3/5 length of segment II (Fig. 177F); 8) tarsal claws with either middle (Fig. 177G) or apical denticles larger than others (Fig. 177H); 9) lateral margins of pronotum glabrous; and 10) gills on abdominal segments I-VII similar, plate-like, with narrow apical process (Figs. 177I-J). Distribution: Chile (Magallanes Province [ca. 53oS] north to Coquimbo Province [ca. 31ºS]), eastward to Argentina (Andean range in Chubut, Rio Negro, and Neuquen Provinces). Biology: The nymphs of Penaphlebia occur in moderately swift flowing streams and rivers, and in the shallow littoral zone of lakes. Some nymphs were collected from underneath silt-coated rocks and stones, while others were found under rocks and submerged logs or among debris trapped between rocks. The adults have been reported to fly from September to March.

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A

B

MA

Sc

C MP2 MP1 CuA

MA

E

D

MP CuA 2

F

G J

K

L

H SP

PE VS

M

N

O

I

P

Q

R

S

T

Plate 175. Penaphlebia, imagos. Figs. 175A-B: Cross veins in costal and subcostal areas of fore wings. 175A: P. (P.) chilensis; 175B, P. (P.) flavidula. 175C-G, whole wings. P. (P.) barriai: 175C, fore wing; 175D, hind wing; P. (P.) exigua: 175E, fore wing; 175F, hind wing; P. (M.) vinosa: 175G, hind wing. 175H, male tarsal claws of P. (P.) barriai; 175I, female sternum IX of P. (P.) barriai. 175J-L, male genitalia, v.v. 175J: P. (P.) barriai; 175K, P. (P.) exigua; 175L, P. (M.) vinosa. 175M-O, penes, v.v. 175M, P. (P.) chilensis; 175N, P. (P.) flavidula; 175O, P. (P.) fulvipes. 175P-T: abdominal terga III-VIII. 175P, P. (P.) barriai; 175Q, P. (P.) chilensis; 175R, P. (P.) exigua; 175S, P. (P.) flavidula; 175T, P. (P.) fulvipes. (SP = Styliger Plate; PE = Posteromedian emargination; VS = Ventral spine). Figs. 175A-D, F-J, L-Q and S-T modified from PESCADOR & PETERS, 1991; 175E-F, K and R from DOMÍNGUEZ & PESCADOR, 1982.

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TO

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PENAPHLEBIA

Imagos 1

–

2(1) – 3(2) – 4(3) – 5(4) –

Hind wings ovate (Fig. 175G); segments III and IV of male protarsi equal in length; posterior margin of styliger plate entire (Fig. 175L); apical 1/3 of penis lobes expanded, each lobe with slender subapical spine (Fig. 175L) ........................ ...................................................................... Subgenus Megalophlebia.....Penaphlebia vinosa Hind wings elliptic (Figs. 175D, F); segment III of protarsi longer than segment IV; styliger plate with midposterior emargination (Figs. 175J-K); apical 1/3 of penis lobes slightly to moderately enlarged, each lobe with stubby subapical spine (Fig. 175O) .......................................................... Subgenus Penaphlebia (Penaphlebia).....2 Body length of male and female 8 mm or less; styliger plate with V-shaped posteromedian emargination (Fig. 175K); dark maculae on abdominal terga as in Fig. 175R; maculae on abdominal sterna absent ...... Penaphlebia (Penaphlebia) exigua Body length of male and female more than 8 mm; styliger plate with broad Ushaped posteromedian emargination (Fig. 175J); dark brown maculae on abdominal terga not as above; abdominal sterna with brown maculae .................. 3 Abdominal terga with reduced dark brown to black maculae, maculae heaviest on VII (Fig. 175P); penes as in Fig. 175J; costal membrane of female fore wings hyaline, subcostal membrane flushed with yellow ................... Penaphlebia (Penaphlebia) barriai Abdominal terga I-IX or I-III, VI-VII, and IX with pronounced dark brown to black maculae (Figs. 175Q, S-T); penes as in Fig. 175M; costal and subcostal membranes of female fore wings flushed with brownish yellow (Fig. 175A) ......... 4 Pre-pterostigmatic costal and subcostal veins of male and female fore wings well developed and thickly infuscated with brown (Fig. 175A); penis lobes as in Fig. 175M ................................................................................ Penaphlebia (Penaphlebia) chilensis Pre-pterostigmatic costal and subcostal cross veins of male fore wings weakly developed (Fig. 175B), female fore wings a little more developed, faintly infuscated; penis lobes as in Figs. 175N-O .................................................................. 5 Abdominal terga I-IX with pronounced black maculae (Fig. 175S); submedian maculae on tergum VIII extended beyond half length of segment; penis lobes as in Fig. 175N ................................................................... Penaphlebia (Penaphlebia) flavidula Abdominal terga IV-V and VIII with reduced submedian maculae (Fig. 175T); submedian maculae on tergum VIII not extended beyond half length of segment; penis lobes as in Fig. 175O .......................................... Penaphlebia (Penaphlebia) fulvipes

Mature Nymphs 1 –

Prominent posterolateral projections on abdominal segments VI-IX (Fig. 177M); abdominal gills apically terminated with a blade-like process (Fig. 177J); mandibles with deep dorsal groove (Fig. 177C) ..... Subgenus Megalophlebia.....Penaphlebia vinosa Posterolateral projections on abdominal segments VII-IX (Figs. 177K-L); abdominal gills apically terminated with a filamentous process (Fig. 177I); mandibles without dorsal groove, or shallow (if present) (Fig. 177B) ..................... ............................................................................... Subgenus Penaphlebia (Penaphlebia).....2

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4(3) – 5(2) –

CLAVES

Mid-denticles on tarsal claws larger than apical denticles (Fig. 177G) ................... 3 Mid-denticles on tarsal claws smaller than apical denticles (Fig. 177H) ................ 5 Posterior spines on abdominal terga equal to a little longer than setae; setae on joints of caudal filaments short (Fig. 177N); pronounced dark brown to black maculae on abdominal terga I-IX (Figs. 175R-S) ........................................................ 4 Posterior spines on abdominal terga distinctly longer than setae; setae on joints of caudal filaments long (Fig. 177O); abdominal terga I-II, and VIII without maculae or with greatly reduced dark brown maculae (Fig. 175P) ........................................... ............................................................................................ Penaphlebia (Penaphlebia) barriai Body length less than 8 mm; posterolateral projections on abdominal terga VIIIIX weakly developed (Fig. 177K); abdominal gills whitish ......................................... ............................................................................................. Penaphlebia (Penaphlebia) exigua Body length distinctly longer than 8 mm; posterolateral projections on abdominal terga VIII-IX well developed (Fig. 177L); abdominal gills grayish-black ................ .......................................................................................... Penaphlebia (Penaphlebia) flavidula Setae on posterior margin of abdominal terga twice as long as spines; tarsi bicolorous ....................................................................... Penaphlebia (Penaphlebia) chilensis Setae on posterior margin of abdominal terga equal to slightly longer than spines; tarsi unicolorous ............................................................. Penaphlebia (Penaphlebia) fulvipes PARA

PENAPHLEBIA

Imagos 1

–

2(1)

–

3(2)

Alas posteriores ovales (Fig. 175G); segmentos III y IV de los protarsos masculinos de igual longitud; margen posterior de la placa estilígera entero (Fig. 175L); 1/3 apical de los lóbulos de los penes expandidos, cada lóbulo con espina subapical delgada (Fig. 175L) .............. Subgénero Megalophlebia.....Penaphlebia vinosa Alas posteriores elípticas (Figs. 175D, F); segmento III de los protarsos más largo que segmento IV; placa estilígera emarginada posteriormente (Figs. 175J-K); 1/3 apical de los lóbulos de los penes un poco a moderadamente ensanchados, cada lóbulo con espina subapical gruesa (Fig. 175O) ............................................................ ............................................................................. Subgénero Penaphlebia (Penaphlebia).....2 Longitud del cuerpo del macho y de la hembra 8 mm o menos; placa estilígera con emarginación posteromediana en forma de V (Fig. 175K); manchas oscuras en tergos abdominales como en Fig. 175R; manchas en esternos abdominales ausentes ............................................................................. Penaphlebia (Penaphlebia) exigua Longitud del cuerpo del macho y de la hembra más de 8 mm; placa estilígera con emarginación posteromediana ancha en forma de U (Fig. 175J); manchas castañas oscuras en tergos abdominales no como el caso anterior; esternos abdominales con manchas castañas ....................................................................................................... 3 Tergos abdominales con manchas reducidas de color castaño oscuras a negro, manchas más fuertes sobre tergo VII (Fig. 175P); penes como en Fig. 175J; membrana costal de las alas anteriores de la hembra hialinas, membrana subcostal teñidas de amarillenta .................................................... Penaphlebia (Penaphlebia) barriai

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Tergos abdominales I-IX ó I-III, VI-VII y IX con manchas fuertes de color castaño oscuras a negro (Figs. 175Q, S-T); penes como en Fig. 175M; membranas costal y subcostal de las alas anteriores de la hembra teñidas de amarillo-castaño (Fig. 175A) ........................................................................................................................... 4 Venas costal y subcostal pre-pterostigmaticas de las alas anteriores del macho y de la hembra bien desarrolladas y fuertemente teñidas de castaño (Fig. 175A); lóbulos de los penes como en Fig. 175M ................ Penaphlebia (Penaphlebia) chilensis Venas costal y subcostal pre-pterostigmaticas de las alas anteriores del macho y de la hembra no bien desarrolladas (Fig. 175B), ala anterior de la hembra un poco más desarrollada y débilmente teñidas de castaño; lóbulos de los penes como en Figs. 175N-O. ...................................................................................................................... 5 Tergos abdominales I-IX con manchas fuertes negras (Fig. 175S); manchas submediales en tergo VIII extendiéndose más allá de la mitad de la longitud del segmento; lóbulos de los penes como en Fig. 175N ........................................ Penaphlebia (Penaphlebia) flavidula Tergos abdominales IV-V y VIII con manchas submediales reducidas (Fig. 175T); manchas submediales en tergo VIII no extendiéndose más allá de la mitad la longitud del segmento; lóbulos de los penes como en Fig. 175O ............................. ............................................................................................ Penaphlebia (Penaphlebia) fulvipes

Ninfas Maduras 1

–

2(1) – 3(2)

–

4(3) –

Prominentes proyecciones posterolaterales en segmentos abdominales VI-IX (Fig. 177M); branquias terminadas apicalmente en un proceso en forma de hoja (Fig. 177J); mandíbulas con un surco dorsal profundo (Fig. 177C) ................................... .................................................................... Subgénero Megalophlebia.....Penaphlebia vinosa Proyecciones posterolaterales en segmentos abdominales VII-IX (Fig. 177K-L); branquias terminadas apicalmente en un proceso filamentoso (Fig. 177I); mandíbulas sin surco dorsal, o si presente, poco profundo (Fig. 177B) ............................................. ............................................................................. Subgénero Penaphlebia (Penaphlebia).....2 Dentículos medios de las uñas tarsales más grande que los apicales (Fig. 177G) ...... 3 Dentículos medios de las uñas tarsales más pequeño que los apicales (Fig. 177H) ....... 5 Espinas posteriores en tergos abdominales de iguales a poco más largas que las setas; setas en las articulaciones de los segmentos de los filamentos caudales cortas (Fig. 171N); tergos abdominales I-IX con fuertes manchas castañas a negras (Fig. 175R-S) .......................................................................................................... 4 Espinas posteriores en tergos abdominales mucho más largas que las setas; setas en las articulaciones de los segmentos de los filamentos caudales largas (Fig. 177O); tergos abdominales I-II y VIII con o sin manchas castaño oscuras muy reducidas (Fig. 175P) ..................................................... Penaphlebia (Penaphlebia) barriai Longitud del cuerpo menos de 8 mm; proyecciones posterolaterales en tergos abdominales VIII-IX débilmente desarrolladas (Fig. 177K); branquias abdominales blanquecinas ...................................................................... Penaphlebia (Penaphlebia) exigua Longitud del cuerpo más de 8 mm; proyecciones posterolaterales en tergos abdominales VIII-IX bien desarrolladas (Fig. 177L); branquias abdominales grisnegruzcos ....................................................................... Penaphlebia (Penaphlebia) flavidula

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Plate 176. Penaphlebia chilensis. Fig. 176A, nymphal habitus. Drawing by A. Dupuy.

5(2) –

Setas en los márgenes posteriores de tergos abdominales dos veces más largas que las espinas; tarsos bicolores ......................................... Penaphlebia (Penaphlebia) chilensis Setas en los márgenes posteriores de tergos abdominales de iguales a levemente más largos que las espinas; tarsos de un solo color .......... Penaphlebia (Penaphlebia) fulvipes

Subgenus Penaphlebia (Megalophlebia) PETERS & EDMUNDS Penaphlebia (Megalophlebia) PESCADOR & PETERS, 1991: 29.

Taxonomy: Penaphlebia (Megalophlebia) can be separated from the other subgenus of Penaphlebia by the following combination of characters. In the imago: 1) Hind wings ovate (Figs. 175G); 2) segment III and IV of male protarsi equal in length; 3) apical 1/3 of penis lobes expanded, each lobe with slender subapical spine (Fig. 175L); 4) styliger plate entire

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(Fig. 175L). In the nymph: 1) mandibles with deep dorsal groove (Fig. 177C); 2) abdominal gills apically terminated with a blade-like process (Fig. 177J); 3) dorsum of femora with oblong setae; 4) posterolateral projections on abdominal segments VI-IX (Figs. 177M). Distribution: Western and Eastern slopes of the Andes from approximately 33ºS to 43ºS. Penaphlebia (Megalophlebia) vinosa (DEMOULIN) Atalophlebia vinosa DEMOULIN, 1955e: 7 (male). Penaphlebia vinosa; PETERS & EDMUNDS, 1972: 1399; HUBBARD, 1982a: 266; DOMÍNGUEZ & PESCADOR, 1983: 21. Penaphlebia (Megalophlebia) vinosa; PESCADOR & PETERS, 1991: 29; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 70.

Taxonomy: PESCADOR & PETERS (1991) established the monotypic subgenus Megalophlebia for Penaphlebia vinosa. The characters that distinguish P. (M.) vinosa from the other species of the genus also distinguish the subgenus Megalophlebia from the subgenus Penaphlebia (Penaphlebia) and are as follow: In the imagos: 1) Hind wings large and ovate (Fig. 175G); 2) inner margin of segment I of genital forceps with a strongly developed angular bend (Fig. 175L); and 3) penis lobes with slender subapical spine projecting posteriorly (Fig. 175L). In the nymph: 1) Well developed posterolateral projections on abdominal segments VI-IX (Fig. 177M); 2) abdominal gills apically terminated in a bladelike process (Fig. 177J); 3) mandibles with deep dorsal groove (Fig. 177C); and 4) caudal filaments with long apical and intersegmental setae (Fig. 177O). Distribution and Biology: Southern Chile and Argentina, from ca. 33ºS to Magallanes, ca 53ºS, XII Region. The nymphs of Penaphlebia (M.) vinosa occur in moderately swift flowing streams and rivers, and in the shallow littoral zone of lakes from 200 to 1200 m above sea level. The nymphs were collected underneath silt-coated rocks and stones. Collection records indicate that the adults occur from October to February. Subgenus Penaphlebia (Penaphlebia) PETERS & EDMUNDS Penaphlebia (Penaphlebia) PETERS & EDMUNDS, 1972: 1399; PESCADOR & PETERS, 1980b: 45; PESCADOR & PETERS, 1991: 6.

Taxonomy: Penaphlebia (Penaphlebia) can be separated from the other subgenus of Penaphlebia by the following combination of characters. In the imago: 1) Hind wings elliptic (Figs. 175D, F); 2) segment III of male protarsi longer than segment IV; 3) apical 1/3 of penis lobes slightly to moderately enlarged, with stubby subapical spine posterolaterally projected (Figs. 175J-K); 4) styliger plate with broad U-shaped (Fig. 175J) or Vshaped emargination (Fig. 175K). In the nymph: 1) mandibles without dorsal groove, or shallow (if present) (Fig. 177B); 2) abdominal gills apically terminated with a filamentous process (Fig. 177I); 3) dorsum of femora with ovate or spatulate setae; 4) posterolateral projections on abdominal segments VII-IX (Figs. 177K-L). Distribution: Western and Eastern slopes of the Andes from approximately 30ºS to 43ºS.

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Penaphlebia (Penaphlebia) barriai PESCADOR & PETERS Penaphlebia (Penaphlebia) barriai PESCADOR & PETERS, 1991: 6.

Taxonomy: PESCADOR & PETERS (1991) described P. (P.) barriai based on nymphs and reared imagos from Rio Illapel, Coquimbo Province, Chile. The species can be distinguished from other species of the genus by the following combination of characters. In the imagos: 1) Costal and subcostal cross veins of male and female fore wings well developed and thickly infuscated with dark brown (Fig. 175C); 2) segment III of male protarsi longer than segment IV; 3) abdominal maculae greatly reduced, as in Fig. 175P; 4) styliger plate with broad and shallow U-shaped posteromedian emargination (Fig. 175J); 5) angular bend on inner margin of segment 1 of genital forceps moderately developed (Fig. 175J); and 6) shape of penis lobes similar to Fig. 175J. In the nymph: 1) Mandibles without dorsal groove, or shallow if present (Fig. 177B); 2) middle denticles on tarsal claws larger than apical denticles (Fig. 177G); 3) hair-like setae on posterior margin of abdominal terga at least twice length of spines; 4) posterolateral projections on abdominal segments VIII-IX, well developed; and 5) caudal filaments with long apical and intersegmental setae (Fig. 177O). Distribution and Biology: North-Central Chile, from Valaparaiso (33º02´S) V Región to Limari 30º37´S, IV Región. The rheophylic nymphs of this species occur mostly under rocks and submerged logs in streams and rivers from 300 to 1100 m above sea level. Adults were collected in October and November. A swarm composed of few individuals was observed in the morning. Penaphlebia (Penaphlebia) chilensis (EATON) Atalophlebia chilensis EATON, 1883-1888: 91, pl. 10 Fig. 16g (male); nec ULMER, 1904: Figs. 12-13 (female); ULMER, 1920c: 115; LESTAGE, 1930a: 249; NAVÁS, 1930d: 358; LESTAGE, 1931b: 51; ULMER, 1938: 106; ULMER, 1942: 124 (male); TRAVER, 1946: 419; DEMOULIN, 1955b: 9; DEMOULIN, 1955e: 2 (male, female); THEW, 1960b: 132; KIMMINS, 1960: 294. Penaphlebia chilensis; PETERS & EDMUNDS, 1972: 1399; TSUI & PETERS, 1975: Fig. 58 (male); HUBBARD, 1982a: 266; DOMÍNGUEZ & PESCADOR, 1983: 21 (nymph); PESCADOR & PETERS, 1991: 11; DOMÍNGUEZ, HUBBARD & PESCADOR. 1994: 67.

Taxonomy: EATON (1883) first described P. chilensis from a male imago and subimago from an unknown locality in Chile. DEMOULIN (1955) subsequently redescribed the male imago and PESCADOR & PETERS (1991) redefined the species based on nymphs and reared adults including a brief discussion on minor color variations between populations. Penaphlebia chilensis can be distinguished from the other species of the genus by the following combination of characters. In the imagos: 1) Costal and subcostal cross veins of male and female fore wings well developed, thickly infuscated with dark brown (Fig. 175A); 2) maculae on abdominal terga I-III, VI-VII, and IX pronounced, with submedian maculae on tergum VIII slightly extended beyond half length of segment (Fig. 175Q); 3) angular bend on inner margin of segment I of genital forceps moderately developed; 4) styliger plate with a broad U-shaped posteromedian emargination (similar to Fig. 175J); and 5) shape of apex of penis lobes as in Fig. 175M, with subapical spines posterolaterally projecting. In the nymph: 1) Mandibles without dorsal

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Plate 177. Penaphlebia, nymphs. Figs. 177A-F, mouthparts. 177A-B, D-F, P. (P.) barriai; 177C, P. (M.) vinosa. 177A, clypeus and labrum; 177B-C, left mandibles; 177D, maxilla; 177E, hypopharynx; 177F, labium (left dorsal, right ventral). Fore tarsal claws: 177G, P. (P.) flavidula; 177H, P. (P.) chilensis. Abdominal gill IV: 177I, P. (P.) barriai; 177J, P. (M.) vinosa. Abdominal terga VI-X: 177K, P. (P.) exigua; 177L, P. (P.) chilensis; 177M, P. (M.) vinosa. Setation of caudal filaments: 177N, P. (P.) flavidula; 177O, P. (P.) barriai. (D = Denticle; DG = Dorsal Groove; PP = Posterolateral Projection; Roman numbers = segment numbers). Figs. 177A-N modified from PESCADOR & PETERS, 1991.

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groove, or shallow if present (as in Fig. 177B); 2) apical denticles of tarsal claws larger than middle denticles (Fig. 177H); 3) hair-like setae on posterior margins of abdominal terga approximately twice length of spines; 4) posterolateral projections on abdominal segments VIII-IX, well developed (Fig. 177L); 5) gill portions apically terminated in a filamentous process (as in Fig. 177I); and 6) caudal filaments with thick and long apical and intersegmental setae (Fig. 177O). Distribution and Biology: Central Chile, from Chiloe (ca 43ºS) to Choapa (31º43´S), east to the Andean range of Southern Argentina. The nymphs of P. chilensis were collected under medium sized rocks found in moderately swift flowing streams and rivers from 50 to 1100 m above sea level. A few nymphs have also been reportedly collected among debris trapped between rocks. Based on collection records, the adults fly from October to February. Mating flights were observed just before dusk at the shore of Andean lakes in Patagonia. Swarms were composed of from 40-50 individuals to a few hundred. Penaphlebia (Penaphlebia) exigua DOMÍNGUEZ & PESCADOR Penaphlebia exigua DOMÍNGUEZ & PESCADOR, 1983: 21 (male, female, nymph); PESCADOR & PETERS, 1991: 17; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 68.

Taxonomy: The unusually small Penaphlebia exigua was described by DOMÍNGUEZ & PESCADOR (1983) from nymphs and imagos collected from the isolated Somuncurá plateau, in Patagonia. Penaphlebia exigua can be distinguished from other species of the genus by the following combination of characters. In the imagos: 1) Body small: length 6.4-6.9 mm (male), 7.7 mm (female); 2) vein MP2 of at least one of fore wings basally attached to vein CuA (Fig. 175C); 3) black maculae pronounced on abdominal terga I-IX (Fig. 175R); 4) subapical spines of penis lobes posteriorly projecting (Fig. 175K); and 5) styliger plate with V-shaped posteromedian emargination (Fig. 175K). In the nymph: 1) Body length from 6.9-7.2 mm; 2) mandibles usually without dorsal groove, or shallow if present (as in Fig. 177B); 3) middle denticles on claws larger than apical denticles (Fig. 177G); 4) posterolateral spine on abdominal segments VIIIIX, weakly developed (Fig. 177K); 5) hair-like setae on posterior margin of abdominal terga subequal to length of spines; and 6) caudal filaments with short and thin apical setae (Fig. 177N). Distribution and Biology: Argentina (Patagonia), Meseta de Somuncurá. The nymphs of P. exigua were collected in a small stream at 700 m above sea level in Somuncurá Plateau with a thick growth of Nostoc and moss. Adults were collected in February. Penaphlebia (Penaphlebia) flavidula PESCADOR & PETERS Penaphlebia flavidula PESCADOR & PETERS, 1991: 20; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 69.

Taxonomy: PESCADOR & PETERS (1991) described P. flavidula from nymphs and reared adults. It can be distinguished from other species of the genus by the following combination of characters. In the imagos: 1) Subapical spine of penis lobes posteriorly projecting (Fig. 175N); 2) styliger plate with broad U-shaped posteromedian emargin-

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ation (as in Fig. 175J); 3) pronounced dark brown to black maculae on abdominal terga I-IX (Fig. 175S); and 4) segment III of male protarsi longer than segment IV. In the nymph: 1) Mandibles without dorsal groove, or shallow if present (Fig. 177B); 2) middle denticles of tarsal claws larger than apical denticles (Fig. 177G); 3) hair-like setae and spines on posterior margin of abdominal terga subequal in length; 4) well developed posterolateral projections on abdominal segments VIII-IX (Fig. 177L); 5) caudal filaments with short apical setae (Fig. 177N). Distribution and Biology: Central and Southern Chile (to 45º34´S) and Andean range in Rio Negro, Neuquén and Chubut in Argentina. The nymphs of P. flavidula inhabit various types of bottom substrates from small rocks, organic debris trapped between stones or rocks, submerged logs to floating roots of trees or shrubs growing on the banks of streams and rivers at altitudes of 50 to 1100 m. Collection records indicate that the adults of P. flavidula occur from September to March. Penaphlebia (Penaphlebia) fulvipes (NEEDHAM & MURPHY) Atalophlebia fulvipes NEEDHAM & MURPHY, 1924, 24: 34; LESTAGE, 1930a: 249; nec LESTAGE, 1931b: 52 (male); TRAVER 1946: 420; DEMOULIN, 1955b: 9 (male); DEMOULIN, 1955e: 6 (male). Penaphlebia fulvipes; PETERS & EDMUNDS, 1972: 1399; HUBBARD, 1982a: 266; DOMÍNGUEZ & PESCADOR, 1983: 21; PESCADOR & PETERS, 1991: 26; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 69. Atalophlebia sepia THEW, 1960b: 130 (male). Penaphlebia sepia; PETERS & EDMUNDS, 1972: 1399; HUBBARD, 1982a: 266; DOMÍNGUEZ & PESCADOR, 1983: 21.

Taxonomy: NEEDHAM & MURPHY (1924) described P. fulvipes from a single male imago. DEMOULIN (1955b) later redescribed the imago and PESCADOR & PETERS (1991) subsequently redefined the species based on a series of nymphs and reared adults. Additionally, both authors placed P. sepia as junior synonym of P. fulvipes. Penaphlebia fulvipes can be distinguished from the other species by the following combination of characters. In the imagos: 1) Subapical spine of penis lobe projecting posteriorly (Fig. 175O); 2) styliger plate with broad U-shaped posteromedian emargination (as in Fig. 175J); 3) pronounced black maculae on terga I-III, VI-VII, and IX, submedian maculae on tergum VIII reduced (Fig. 175T); and 4) segment III of male protarsi longer than segment IV. In the nymph: 1) Mandibles without dorsal groove or shallow if present (Fig. 177B); 2) apical denticles of tarsal claws larger than median denticles (Fig. 177H); 3) tarsi bicolored, brown with narrow basal pale yellow band; 4) well developed posterolateral projections on abdominal segments VIII-IX (Fig. 177L); 5) posterior spines of abdominal terga subequal in length to hair-like setae; 6) gills on abdominal segments I-VII terminating apically in filamentous process (as in Fig. 177I); and 7) caudal filaments with long thick setae (Fig. 177O). Distribution and Biology: From Valdivia (39º47´S) X Region to Ultima Esperanza (51º43´S) XII Region in Chile and Andean range in Chubut and Neuquén in Argentina. The nymphs of P. fulvipes were found underneath submerged logs and tree limbs in moderately flowing streams at low altitudes. The adults have been collected in December.

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Penaphlebia anastasii (NAVÁS) Nomen Dubium Atalophlebia anastasii NAVÁS, 1930d: 357 (male); ULMER, 1938: 107; PETERS & EDMUNDS, 1972: 1401; CAMOUSSEIGHT, 1980: 27; HUBBARD, 1982a: 263; ALBA-TERCEDOR & PETERS, 1985: 218; PESCADOR & PETERS, 1991: 3 (nomen dubium).

This species is presently unidentifiable and according to PESCADOR & PETERS (1991) must be considered a nomen dubium. Distribution: Chile. Penaphlebia athanasii (NAVÁS) Nomen Dubium Atalophlebia athanasii NAVÁS, 1930d: 356 (male); ULMER, 1938: 107; PETERS & EDMUNDS, 1972: 1401; HUBBARD, 1982a: 263; ALBA-TERCEDOR & PETERS, 1985: 218; PESCADOR & PETERS, 1991: 3 (nomen dubium).

This species is presently unidentifiable and according to PESCADOR & PETERS (1991) must be considered a nomen dubium. Distribution: Chile. Penaphlebia discolor (NAVÁS) Nomen Dubium Atalophlebia discolor NAVÁS, 1928: 125 (male, female); LESTAGE, 1931b: 52; NAVÁS, 1933a: 233; PETERS & EDMUNDS, 1972: 1401; CAMOUSSEIGHT, 1980: 27; HUBBARD, 1982A: 263; ALBA-TERCEDOR & PETERS, 1985: 218; PESCADOR & PETERS, 1991: 3 (nomen dubium).

This species is presently unidentifiable and according to PESCADOR & PETERS (1991) must be considered a nomen dubium. Distribution: Chile. Penaphlebia fenestrata (NAVÁS) Nomen Dubium Atalophlebia fenestrata NAVÁS, 1930d: 358 (male); ULMER, 1938: 107; PETERS & EDMUNDS, 1972: 1401; HUBBARD, 1982a: 263; ALBA-TERCEDOR & PETERS, 1985: 218; PESCADOR & PETERS, 1991: 3 (nomen dubium).

This species is presently unidentifiable and according to PESCADOR & PETERS (1991) must be considered a nomen dubium. Distribution: Chile. Penaphlebia hyalina (NAVÁS) Nomen Dubium Atalophlebia hyalina NAVÁS, 1930d: 358 (male, female); ULMER, 1938: 107; PETERS & EDMUNDS, 1972: 1401; HUBBARD, 1982a: 263; ALBA-TERCEDOR & PETERS, 1985: 218; PESCADOR & PETERS, 1991: 3 (nomen dubium).

This species is presently unidentifiable and according to PESCADOR & PETERS (1991) must be considered a nomen dubium. Distribution: Chile.

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Penaphlebia rofucensis (NAVÁS) Nomen Dubium Atalophlebia rofucensis NAVÁS, 1934a: 25 (male, female); ULMER, 1938: 107; PETERSEN & GAEDIKE, 1968: 962; PETERS & EDMUNDS, 1972: 1401; PESCADOR & PETERS, 1991: 3 (nomen dubium). Atolophlebia [sic] rofucensis; HUBBARD, 1982a: 263.

This species is presently unidentifiable and according to PESCADOR & PETERS (1991) must be considered a nomen dubium. Distribution: Chile. Penaphlebia valdiviae (NAVÁS) Nomen Dubium Atalophlebia valdiviae NAVÁS, 1936a: 138 (male); ULMER, 1938: 107; PETERS & EDMUNDS, 1972: 1401; HUBBARD, 1982a: 263; PESCADOR & PETERS, 1991: 3 (nomen dubium).

This species is presently unidentifiable and according to PESCADOR & PETERS (1991) must be considered a nomen dubium. Distribution: Chile. Genus Perissophlebiodes SAVAGE Perissophlebia SAVAGE, 1982: 209. (Type-species: Perissophlebia flinti SAVAGE, original designation). Perissophlebiodes SAVAGE, 1983a: 204. (Type-species: Perissophlebia flinti SAVAGE, objective synonymy).

Taxonomy: SAVAGE (1982) established this genus, under the name Perissophlebia, based on the unusual characteristics of nymphs. He later discovered that the name was a junior homonym of Perissophlebia TILLYARD and proposed the name Perissophlebiodes. The nymph of Perissophlebiodes (Fig. 178A) presents an unusal general aspect and the mouthparts are as in Figs. 178B-G. The material used by SAVAGE for the original description was poorly preserved, and for this reason some characters were not observable. Recently, another nymph became available, and it was clear that the hind wing pads were absent (DOMÍNGUEZ et al., 2001). Adult Characteristics: Unknown. Nymphal Characteristics: 1) Labrum (Fig. 178B) approximately same width as clypeus, with deep anteromedian emargination, with reduced denticles (Fig. 178C); 2) median area of outer margin of mandibles with rounded protuberance (Fig. 178D); 3) anterolateral margin of galea-lacinia of maxillae with small, round protuberance (Fig. 178E); 4) labial palpi with thick setae (Fig. 178G); 5) hind wing pads absent; 6) abdominal gill on segments I-VII lancelolate, narrow, dorsal and ventral portions similar (Fig. 178H); 7) fore legs heavily covered with long setae (Fig. 178I); 8) tarsal claws with three large denticles on inner margin and a row of smaller subapical denticles (Fig. 178J). Distribution: Southern and Central Brazil. Biology: The nymphs were collected in a clear stream with rapid current and rocky bottom.

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Plate 178. Perissophlebiodes flinti, nymph. Figs. 178A-J: P. flinti. 178A, habitus; 178B, labrum d.v.; 178C, detail of anteromedian emargination of labrum; 178D, left mandible; 178E, maxilla; 178F, hypopharynx; 178G, labium (left dorsal, right ventral); 178H, gill II; 178I, fore leg; 178J, tarsal claw. (D = Denticles; P = Protuberance; Roman numbers = segment numbers). Figs. 178A-J modified from SAVAGE, 1982.

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Perissophlebiodes flinti (SAVAGE) Perissophlebia flinti SAVAGE, 1982: 212 (nymph). Perissophlebiodes flinti; SAVAGE, 1983a: 204.

Taxonomy: Perissophlebiodes flinti is the only known species in the genus, and can be recognized by the characters listed in the generic diagnosis. Distribution and Biology: As in generic discussion. Genus Rhigotopus PESCADOR & PETERS Genus C PESCADOR & PETERS, 1980b: 47. Rhigotopus PESCADOR & PETERS, 1982: 4; SAVAGE, 1987b: 207; DOMÍNGUEZ et al., 1992: 21; HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 219; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 71; MISERENDINO, 1996: 22. (Type-species: Rhigotopus andinensis PESCADOR & PETERS, original designation).

Taxonomy: The genus Rhigotopus was established based on male and female imagos by PESCADOR & PETERS (1982) who included a brief description of the eggs. In 1996, MA

Sc

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Plate 179. Rhigotopus andinensis, imago. Figs. 179A-F. 179A, fore wing; 179B, hind wing; 179C, male genitalia, v.v.; 179D, male tarsal claws; 179E, abdominal terga III-VII; 179F, apex of female sternum IX. (PS = Penis spine-like projection). Figs. 179A-F modified from PESCADOR & PETERS, 1982.

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MISERENDINO reared and associated the nymphs of this species, and also provided some interesting data on its biology. Adult Characteristics: 1) Fork of vein MA of fore wings symmetrical (Fig. 179A); 2) vein Sc of hind wings at least 3/4 maximum length of hind wings (Fig. 179B); 3) hind wings without costal projection; 4) tarsal claws of a pair similar, both apically hooked, each with an opposing hook (Fig. 179D); 5) penis lobes divided, each lobe with lateral spine-like projection and ventral submedian thickening (Fig. 179C); and 6) female sternum IX deeply cleft (Fig. 179F). Nymphal Characteristics: 1) Labrum slightly narrower than clypeus (Fig. 180A), anteromedian emargination as in Fig. 180B, length of labrum approximately 0.4 maximum width; 2) mandibles with slightly curved outer margin, and long lateral setae (Fig. 180C); 3) maxilla as in Fig. 180D; 4) hypopharynx as in Fig. 180E; 5) labium with segment III of palpi broadened with heavy row of spine-like setae (Fig. 180F); 6) anterior and lateral A

C B D

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G F H Plate 180. Rhigotopus andinensis, nymph. Figs. 180A-F: mouthparts. 180A, labrum, d.v. ; 180B, anteromedian emargination of labrum, enlarged; 180C, left mandible; 180D, maxilla; 180E, hypopharynx; 180F, labium (left dorsal, right ventral); 180G, fore tarsal claw; 180H, gill IV. (D = denticle; Roman numbers = segment numbers). Figs. 180A-H modified from MISERENDINO, 1996.

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margins of pronotum with long setae; 7) tarsal claws with denticles little larger apically (Fig. 180G); 8) posterolateral projections on abdominal segments IV-IX; 9) gills on abdominal segments I-VI similar, gills on segment VII greatly reduced, dorsal portion of gills I-VI ending in three filaments and larger than similar ventral portion (Fig. 180H). Distribution: Chile (Llanquihue and Cautín provinces, X Región) and Southwestern Argentina (Chubut province). Biology: The adults were collected in the evening, from November to March. The nymphs were found under rocks, in small streams and large rivers, at depths from 20 cm to 60 cm with water temperatures ranging from 12-15°C. Rhigotopus andinensis PESCADOR & PETERS Rhigotopus andinensis PESCADOR & PETERS, 1982: 5 (male, female, egg); HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 219; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 71; MISERENDINO, 1996: 22 (nymph).

Taxonomy: Rhigotopus andinensis is the only species known in the genus, and can be recognized by the characters listed in the generic diagnosis. Distribution and Biology: As in generic description. Genus Secochela PESCADOR & PETERS Genus E PESCADOR & PETERS, 1980b: 50. Secochela PESCADOR & PETERS, 1982: 7; HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 23; DOMÍNGUEZ et al., 2001: 40; PESCADOR & GONSER, 2001: 348 (nymph). (Type-species: Secochela illiesi PESCADOR & PETERS, original designation).

Taxonomy: The genus Secochela was established by PESCADOR & PETERS (1982) based on adults from Chile. These authors also included a brief description of the egg chorion. Its phylogenetic relationships were analyzed by PESCADOR & PETERS (1980b, as “Genus E”). The nymph was reared by T. GONSER and described by PESCADOR & GONSER (2001). Adult Characteristics: 1) Fork of vein MA of fore wings symmetrical (Fig. 181A); 2) vein Sc of hind wings 9/10 maximum length of wing (Fig. 181B); 3) hind wings without costal projection; 4) tarsal claws of a pair dissimilar, one apically hooked, other obtuse and pad-like (Fig. 181D); 5) penis lobes long, only fused basally, each lobe with a short apical spine and setae and serrated along inner subapical margin (Fig. 181C); and 6) female sternum IX entire apically (Fig. 181F). Nymphal Characteristics: 1) Clypeus narrower than labrum and lateral margins of clypeus divergent apically (Fig. 182A); 2) anteromedian emargination of labrum broad, Ushaped with undefined indentations (Fig. 182B); 3) outer margin of mandibles smoothly curved and glabrous except for a thin submedian tuft (Fig. 182C); 4) segment I and II of maxillary palpi subequal in length, segment III slightly shorter with long thick setae; inner margins of segments II and III with spinous setae (Fig. 182D); 5) lateral margins of submentum of labium glabrous (Fig. 182E); 6) tarsal claws with unequal-sized basal denticles, remaining denticles progressively larger apically (Fig. 182F); 7) posterolateral projections on abdominal segments II - IX; 8) gills on abdominal segments I - VII similar, dorsal and ventral portions slender and tapered apically (Fig. 182G).

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MA

A

Sc

B S

E

D

C

F

Plate 181. Secochela illiesi, imago. Figs. 181A-F. 181A, fore wing; 181B, hind wing; 181C, male genitalia, v.v.; 181D, male fore claw; 181E, abdominal terga IV-VII; 181F, female sternum IX. (MA = fork of MA; S = Setae). Figs. 181A-F modified from PESCADOR & PETERS, 1982.

Distribution: Chile (Llanquihue Prov. (41º15´S) to Linares Prov. (35º51´S). Biology: The nymphs were collected in cold water streams. The adults were collected from December to March. Secochela illiesi PESCADOR & PETERS Secochela illiesi PESCADOR & PETERS, 1982: 8 (male, female, egg); PESCADOR & GONSER, 2001: 348 (nymph).

Taxonomy: Secochela illiesi is the only known species in the genus. The abdominal color pattern is given in Fig. 181E. Slight color variation on wing veins and sterna was reported by PESCADOR & PETERS (1982). Distribution and Biology: As in generic description.

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A

C

D

B

D

E

F

G

Plate 182. Secochela illiesi, nymph. Figs. 182A-G. 182A, clypeus and labrum, d.v.; 182B, anteromedian emargination of labrum, enlarged; 182C, left mandible; 182D, maxilla; 182E, labium, left dorsal, right ventral; 182F, fore claw; 182G, gill IV. (D = denticle). Figs. 182A-G modified from PESCADOR & GONSER, 2001.

Genus Simothraulopsis DEMOULIN “Thraulus” (in part); TRAVER, 1947a: 150; 1960b: 73. Simothraulopsis DEMOULIN, 1966a: 15; DOMÍNGUEZ et al., 1997: 142. (Type-species: Simothraulopsis surinamensis DEMOULIN, by original designation, a synonym of Simothraulopsis demerara (TRAVER) (DOMÍNGUEZ et al., 1997)).

Taxonomy: The genus Simothraulopsis was established by DEMOULIN (1966a), based on six nymphs from Surinam. In 1947, TRAVER described Thraulus demerara, based on male and female imagos from Guyana. Later, TRAVER (1960b) stated that T. demerara did not belong in Thraulus, but failed to reassign it to another genus. DOMÍNGUEZ et al. (1997) studied nymphs associated with adults of S. surinamensis and T. demerara and they proved to be conspecific. The close relationships of this genus with Homothraulus and Farrodes are discussed in DOMÍNGUEZ et al. (1997). Adult Characteristics: 1) Fork of vein MA of fore wings asymmetrical (Fig. 183A), stem of MA straight; 2) costal projection of hind wings strongly developed, located about 2/3 distance from wing base to apex (Figs. 183B-C); 3) vein Sc of hind wings ending at base of costal projection (Figs. 183B-C); 4) vein MP of hind wings unforked (Fig. 183C); 5) tarsal claws of a pair dissimilar, one apically hooked, other obtuse, pad-like (Fig. 183E); 6) penes divided in apical half with one ventrally directed spine on each lobe (Figs. 183FG); 7) eyes of male meeting on dorsum of head (Fig. 183D); 8) apex of female sternum IX obtuse, truncated to indented apically (Fig. 183H).

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Nymphal Characteristics: 1) Lateral margins of labrum rounded, with widest part located 2/3 distance from base (Figs. 184A, C); 2) mandibular submedian tuft present (Fig. 184D); 3) labium as in Fig. 184G, setae on palpi segment III as in Fig. 184H; 4) tarsal claws with apical denticle greatly enlarged (Fig. 184J); 5) posterolateral projections on abdominal segments VIII-IX only; 6) gills on abdominal segments I-VII similar, lanceolate (Fig. 184I) of moderate width (width ≤ 1/6 length). CP

MA

Sc

C

MP

A MP

B

SP

D

E

F

G

H

I

Plate 183. Simothraulopsis demerara, imago. Figs. 183A-I. 183A, fore wing; 183B, hind wing; 183C, same, enlarged; 183D, eyes of male imago; 183E, male tarsal claw; 183F, genitalia, v.v.; 183G, genitalia, l.v.; 183H, apex of female sternum IX; 183I, abdominal terga II-IX. (CP = Costal Projection; SP = Styliger Plate). Figs. 183A-I modified from DOMÍNGUEZ et al., 1997.

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A

D

E

B

III II

C

I

F

G

D

J

H

I

Plate 184. Simothraulopsis demerara, nymph. Figs. 184A-J. 184A, labrum d.v.; 184B, detail anteromedian emargination of labrum; 184C, labrum (outline); 184D, left mandible; 184E, maxilla; 184F, hypopharynx; 184G, labium (left dorsal, right ventral); 184H, detail of labial pali III; 184I, gill IV; 184J, tarsal claw. (D = denticle; Roman numbers = segment numbers). Figs. 184A-J modified from DOMÍNGUEZ et al., 1997.

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Distribution: Colombia, Guyana, French Guiana, Surinam, Venezuela and Northern Brazil. This genus shows a vicariant distribution with Homothraulus, its sister group, distributed in the Paraná Basin. Biology: Little known. Adults were collected at light during most of the year, so it is possible that they have a continuous emergence season. Simothraulopsis demerara (TRAVER) Thraulus demerara TRAVER, 1947a: 150; 1960b: 73. Simothraulopsis demerara; DOMÍNGUEZ et al., 1997: 146. Simothraulopsis surinamensis DEMOULIN, 1966a: 18 (nymph); HUBBARD, 1982a: 266.

Taxonomy: Simothraulopsis demarara is the only species described in the genus. It can be recognized by the characteristic abdominal color pattern (Fig. 183I) and the band of color on veins C and Sc of the hind wings (Figs. 183B-C). The nymphs have a color pattern similar to the adults, and the gills are as in Fig. 184I. Distribution and Biology: As in generic description. Genus Terpides DEMOULIN Terpides DEMOULIN, 1966a: 15 (nymph); PETERS & HARRISON, 1974: 178 (adult); DOMÍNGUEZ et al., 2001: 28, 37; DOMÍNGUEZ et al., 2002: 463; ORTH et al, 2000: 33. (Type-species: Terpides guyanensis DEMOULIN, original designation).

Taxonomy: The genus Terpides was established by DEMOULIN (1966a) for the species T. guyanensis based only on nymphs from Surinam. Later, PETERS & HARRISON (1974) described the adults from a new species from St. Vincent, West Indies. There are several undescribed species in continental South America. Adult Characteristics: 1) Upper portion of male eyes located on a short, wide stalk; 2) vein MP2 of fore wings attached to MP1 by cross vein; 3) vein ICu1 of fore wings attached at base to vein CuA (Fig. 185A); 4) hind wings (Figs. 185B-C) small (between 1/8 to 1/5 length of fore wings), with costal projection rounded, weakly developed; 5) tarsal claws of a pair similar, with apical hook and an opposing blunt lobe (Fig. 185D); 6) genital forceps three-segmented (Fig. 185E); 7) penes short and tubular, fused basally, without projections (Fig. 185E); and 8) female sternum IX deeply and broadly cleft apically (Fig. 185F). Nymphal Characteristics (Fig. 186A): 1) Head hypognathous; 2) labrum (Fig. 186B) with broad anteromedian emargination, and with 6 (rarely 5) small, rounded denticles (Fig. 186C); 3) lingua of hypopharynx without lateral processes (Fig. 186D); 4) large apically curved unpectinate seta near large pectinate seta on inner anterior angle of maxillae (Fig. 186E); 5) outer margin of mandible rather straight (Fig. 186F); 6) glossae of labium curved ventrally (Fig. 186G); 7) tarsal claws with large medial denticle; several basal smaller denticles progressively larger apically and several small, equal-sized denticles apically (Fig. 186H); 8) abdominal gills I and VII smaller than gills II-VI, gills I-VII similar, with dorsal portion longer and wider (Fig. 186I), 9) posterolateral spines on abdominal segments III-V or VI and VIII-IX; and 10) caudal

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MA

E

A ICu1

CP CuA MP1

MP2

C

B

D F Plate 185. Terpides, imago. Figs. 185A-F, T. jessiae: 185A, fore wing; 185B, hind wing; 185C, hind wing, detail; 185D, fore claws; 185E, male genitalia, v.v.; 185F, female sternum IX. (CP = Costal Projection). Figs. 185A-F, modified from PETERS & HARRISON, 1974.

filaments with swimming setae (basal 2/5-3/5 of filaments with long setae on inner margin of cerci and lateral margins of terminal filament). Distribution: Argentina, Bolivia, Brazil, Colombia, Ecuador, French Guiana and Surinam. Also West Indies. Biology: Nymphs have been found in fast flowing sections of rivers, or in slow waters near the margins. They are very good swimmers, at first glance looking more like a Baetidae than a Leptophlebiidae. Nymphs were collected from small relatively pristine streams to medium-sized rivers, generally in shaded areas. PETERS & HARRISON (1974) mentioned that the nymphs are remarkably hardy. Subimagos emerge in the late afternoon and the subimaginal molt occurred before the following morning. Terpides guyanensis DEMOULIN Terpides guyanensis DEMOULIN , 1966a: 17 (nymph); HUBBARD , 1982a: 266; S AVAGE & P ETERS , 1978: 296.

Taxonomy: This species is known only from the nymphal stage. It can be separated from the other species of the genus by the following combination of characters: 1) Apex of tibia I with a brown spot; 2) fore femora with subapical brown spots; 3) abdominal terga with two pairs of submedian spots. Distribution and Biology: Surinam (Brokopondo). Nymphs were collected from Podostemaceae in riffle areas, with temperature of 28ºC-29ºC.

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B

C D

D

A

E

G

D

H

F

I

Plate 186. Terpides, nymphs. Figs. 186A-I. 186A, Terpides sp. habitus. 186B-I, T. jessiae: 186B, clypeus and labrum, d.v.; 186C, anteromedian emargination of labrum, enlarged; 186D, hypopharynx, v.v.; 186E, maxilla; 186F, mandible; 186G, labium: left dorsal, right ventral; 186H, fore claw; 186I, gill IV. (D = denticle). Fig. 186A, drawing by C. Molineri. Figs. 186B-I, modified from PETERS & HARRISON, 1974.

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Genus Thraulodes ULMER Thraulodes ULMER, 1920a: 33; ULMER, 1920c: 116; ULMER, 1921: 263; NEEDHAM & MURPHY, 1924: 40; ULMER, 1933: 203; TRAVER, 1935: 551 (in NEEDHAM, TRAVER & HSU, 1935); TRAVER, 1944: 11; TRAVER, 1946: 428; DEMOULIN, 1955a: 16; TRAVER, 1964: 33; TRAVER & EDMUNDS, 1967: 352; ALLEN & BRUSCA, 1978: 413; DOMÍNGUEZ & SAVAGE, 1987: 44; DOMÍNGUEZ, 1987a: 47; SAVAGE, 1987b: 207; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 71. (Type-species: Thraulus laetus EATON, original designation).

Taxonomy: The genus Thraulodes is probably one of the most abundant and the most geographically widespread leptophlebiids in South America and its distribution extends to the Nearctic Region. The genus was established by ULMER (1920) for a male imago from Colombia originally described as Thraulus laetus by EATON in 1883. TRAVER & EDMUNDS (1967), and ALLEN & BRUSCA (1978) called for the establishment of speciesgroups, the first based on adult genitalic characters and the second based on the gill structure of the nymphs. TRAVER & EDMUNDS established two groups, excluding several species impossible to group with this method. The groups of species proposed by these authors are not the same and are not followed here. We use most of the terminology proposed by TRAVER & EDMUNDS (1967) in our treatment of the genus and give a schematic of a penis lobe to show terms used to describe the genitalia (Fig. 187A). When we speak of cross veins basal to the bulla and of the coloration of the costal and subcostal areas, we refer always to the fore wing. Adult Characteristics: 1) Eyes of male meet on meson of head; 2) veins within Rs of fore wings not forming two triangles (Figs. 187B, D); 3) vein MP of hind wings forked (Figs. 187C, E); 4) costal projection of hind wings usually rounded (rarely acute) and located near mid-point of anterior border (Figs. 187C, E); 5) penes divided to base, each lobe with subapical spine directed medially (Sp in Figs. 187A, J, N); 6) forceps sockets fused, forming a single cavity (Figs. 187J, N); 7) female sternum IX broadly cleft apically (Fig. 187G); and 7) claws of a pair dissimilar, one hooked and the other blunt (Fig. 187H). Nymphal Characteristics: 1) General aspect as in Fig. 190A; 2) labrum angled laterally, with anterior margin entire (Fig. 191A); 2) mandibles angularly curved, with setae on distal half (Fig. 191B); 3) submentum without lateral setae (Fig. 191E); 4) lateral margins of pronotum glabrous (Fig. 190A); 5) tarsal claws with subequal-sized denticles to denticles progressively larger apically (Figs. 191I-J); 6) abdominal gills on segments I-VII similar, lanceolate, narrow in some species (Fig. 191F) and broader in others (Figs. 191GH); 7) posterolateral projections on abdominal segments III-IX. Distribution: America, between 30-40ºN and 25-28ºS. Biology: The nymphs of Thraulodes are found in almost all streams and rivers and at a wide range of altitudes. In Northwestern Argentina, for example, the same species has been recorded from altitudes ranging from 400 to 1700 m. They are generally found in well-aerated waters, although can tolerate some fluctuations in temperature and oxygen level. The nymphs are generally associated with stony bottoms, although some are found on submerged vegetation.

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a

Sp lp

rf

B

CP

CuP

A

ICu2

C

CuP ICu2

E

MP

bs

D

CP

F

G

J

L I

H

K

N P

M

O

Plate 187. Thraulodes, imagos. Figs. 187A-P. 187A, General aspect of penis lobe (v.v.), showing terminology used for genus. Figs. 187B-F, wings: 187B, T. bolivianus, fore wing; 187C, idem, hind wing; 187D, T. liminaris, fore wing; 187E, idem, hind wing; 187F, T. colombiae, hind wing. 187G-H, T. consortis: 187G, apex of female sternum IX; 187H, male tarsal claws. 187I-J: T. bolivianus. 187I, abdominal terga II-VII; 187J, penes, v.v. 187K-L: T. bomplandi. 187K abdominal terga II-VII; 187L, left lobe of penis, v.v. 187M-N: T. cochunaensis. 187M, abdominal terga II-VIII; 187N, genitalia, v.v. 187O-P: T. consortis. 187O, abdominal terga II-VII; 187P, genitalia, v.v. (CP = Costal Projection; Arrows = Radial sector; a = apicolateral area; bs = basal lobe; lp = lateral pouch; rf = recurved fold; Sp = subapical spine). Figs. 187A-E, and 187G-P modified from DOMÍNGUEZ, 1987a; 187F from EATON, 1884.

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The subimagos generally emerge at dusk and are attracted to lights. The mating flight is generally composed of relatively few specimens, and takes places from the evening to after dark. KEYS

TO

SOUTH AMERICAN THRAULODES

31

Male Imagos 1 – 2(1) – 3(2) – 4(1) – 5(4) – 6(5) – 7(6) – 8(6) – 9(8) – 10(8) – 31

Abdomen entirely dark brown to orange-brown ........................................................ 2 Abdomen much lighter, with segments II-VI, III-VI, II-V, or II-VI whitish or translucent ........................................................................................................................... 4 Costal and subcostal areas shaded, costal cross veins present basal to bulla; femora with two bands ..................................................................................................... 3 Costal and subcostal areas hyaline, costal cross veins absent basal to bulla; femora with one band .......................................................................................... Thraulodes regulus Vein ICu2 of fore wings (Fig. 187D) attached basally to CuP; spine of each lobe of penes (Fig. 188K) slender and long ............................................ Thraulodes liminaris Vein ICu2 of fore wings detached from CuP (as in Fig. 187B); spine of each lobe of penes (Fig. 187P) broad and robust ............................................Thraulodes consortis Costal and subcostal areas brown, especially in apical 1/3; a large reddish macula on each side of tergum VII; penes as in Fig. 189L ................. Thraulodes venezuelana Costal and subcostal areas yellowish or translucent; without large reddish maculae on tergum VII ..................................................................................................................... 5 One band or apical macula on fore femur .................................................................. 6 Two bands or one band and one macula on fore femur ......................................... 14 Costal and subcostal areas with light yellowish tinge ................................................. 7 Costal and subcostal areas hyaline .................................................................................. 8 Three to four weakly developed costal cross veins basal to bulla; penes (Fig. 188H) very slender, without lateral pouch, recurved fold present only in basal half of penes .............................................................................................. Thraulodes itatiajanus Costal cross veins basal to bulla absent; penes (Fig. 189E) of moderate width, lateral pouch present, recurved fold parallel to margins and reaching to extreme inner margin ......................................................................................... Thraulodes schlingeri Penes relatively long and slender, without lateral pouch (Figs. 188R, 189G) ....... 9 Penes robust, lateral pouch present (Figs. 188P, 189K) ........................................... 10 Abdominal terga I and VII-X brown, II-VI translucent whitish with two small blackish maculae near the anterolateral angle (Fig. 189F) ..... Thraulodes telegraphicus Abdominal terga II-VI and base of VII transparent whitish, remainder of VII yellowish, VIII-X yellowish-brown, pale grayish dorsolateral maculae on segments II-VII (Fig. 188Q) .............................................................................. Thraulodes papilionis Costal cross veins basal to bulla absent ...................................................................... 11 Four weakly developed costal cross veins basal to bulla ......................................... 12

We did not include in this key Thraulodes colombiae known only from females and T. limbatus, T. subfasciatus, and T. vitripennis because of insufficient descriptions.

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11(10) Abdominal terga II-V translucent, with central area of II-IV tinged with red; two black maculae close to anterior margins on segments II-VIII (Fig. 188L), terga VI-X reddish with anterior margins of VI-VII and posterior margin and posterolateral corners of segment VIII with white areas; genitalia as in Fig. 188M ................ Thraulodes marreroi – Abdominal terga II-V and basal 1/3 of VI translucent with transverse reddishbrown bands, intersegmental areas and region adjacent to pleural fold pale yellowish white; posterior 2/3 of tergum VI a deeper brownish color than II-V, but paler than VIII-X, two faint gray maculae close to anterior margin on terga IIV; genitalia as in Fig. 188P .......................................................................Thraulodes osiris 12(10) Fore wing length more than 12 mm; basal 1/2 of main longitudinal veins yellowish, darker toward apex; genitalia as in Fig. 188O ................. Thraulodes mucuy – Fore wing length less than 8 mm; veins yellowish, genitalia not as above (Figs. 188G, 189K) ...................................................................................................................... 13 13(12) Abdominal terga VIII-X and posterior 1/2 of VII dark reddish brown, anterior margin of VII and VIII blackish; genitalia as in Fig. 189K ............ Thraulodes ulmeri – Abdominal terga VII-X orange-brown, with whitish areas on segments VII-VIII and X (Fig. 188F); genitalia as in Fig. 188G ................................... Thraulodes guanare 14(5) One complete and one incomplete band on fore femur ......................................... 15 – Two bands on fore femur ............................................................................................... 16 15(14) Three to four weakly developed costal cross veins basal to bulla; abdominal terga II-VI translucent, with maculae as in Fig. 188C; genitalia as in Fig. 188D ....... Thraulodes flinti – Costal cross veins basal to bulla absent; abdominal terga II-V translucent whitish with maculae as in Fig. 187M; genitalia as in Fig. 187N ....... Thraulodes cochunaensis 16(14) Costal and subcostal areas yellowish; 5 to 8 costal cross veins basal to the bulla; .............................................................................................................................................. 17 – Costal and subcostal areas without pigmentation; costal cross veins absent basal to bulla ..................................................................................................................................... 18 17(16) Abdominal terga II-VII with U-shaped maculae (Fig. 187K) formed by two parallel maculae converging posteriorly; genitalia as in Fig. 187L ............................. ............................................................................................................... Thraulodes bomplandi – Abdominal terga II-VI with triangular median maculae near the posterior margin and submedian maculae near the anterior margin; genitalia as in Fig. 188E .......... ................................................................................................................. Thraulodes furficulus 18(16) Large species, fore wings length more than 10.5 mm; abdominal color pattern as in Fig. 187I ........................................................................................... Thraulodes bolivianus – Small species, fore wings length 8 mm or less; abdominal color pattern not as above (Figs. 188A, 189A, H) ......................................................................................... 19 19(18) Abdominal terga II-VII translucent, with small median maculae as in Fig. 188A; penes (Fig. 188B) with a lateral projection at the base of each lobe ........................ ................................................................................................................. Thraulodes daidaleus – Abdominal terga II-VI translucent, abdominal color pattern not as above; penes without lateral projections .............................................................................................. 20 20(19) Abdominal terga II-VI with distinct median or posteromedian maculae (Figs. 189A, H) ............................................................................................................................. 21

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–

Abdominal terga II-VI without maculae or with small submedian maculae .......... ................................................................................................................... Thraulodes laetus 32 21(20) Abdominal terga III-V with broad maculae reaching posterior margin and meeting lateral maculae (Fig. 189A); genitalia as in Fig. 189B ................................... ............................................................................................................ Thraulodes paysandensis – Abdominal terga III-V with separate or medially joined submedian maculae not reaching posterior margin and separated from lateral maculae (Fig. 189H); genitalia as in Fig. 189I ........................................................................ Thraulodes traverae Nymphs 1 – 2(1) – 3(2)

33

–

Abdominal gills broad, tapering abruptly to apex (Figs. 191G-H) .......................... 2 Abdominal gills narrow, tapering gradually to apex (Fig. 191F) .............................. 6 Abdominal gills with main trachea branched (Fig. 191G) ........................................ 3 Abdominal gills with main trachea not branched (Fig. 191H) ................................. 4 Fore tarsal claw with six denticles progressively larger apically (Fig. 191J) ............. .................................................................................................................. Thraulodes liminaris Fore tarsal claw with six or seven denticles progressively larger apically, except the apical denticle slightly larger (Fig. 191I) ...........................................Thraulodes consortis Small species, length of body of mature nymphs less than 7 mm .... Thraulodes traverae Large species, length of body of mature nymphs more than 10 mm ................... 5 Femora with median brown macula ............................................... Thraulodes bolivianus Femora with median and apical brown maculae ........................ Thraulodes itatiajanus Tarsal claw with six or seven denticles progressively larger towards the apex ....... ............................................................................................................ Thraulodes cochunaensis Tarsal claw with three large and three or four smaller denticles ....... Thraulodes daidaleus

CLAVES

PARA LOS

– 4(2) – 5(4) – 6(1)

THRAULODES

SUDAMERICANOS 34

Imagos Macho 1 – 2(1) –

Abdomen castaño-oscuro a castaño-anaranjado ......................................................... 2 Abdomen más claro, con segmentos II-VI, III-VI, II-V ó II-VI blanquecinos o translúcidos .......................................................................................................................... 4 Area costal y subcostal sombreadas, venas transversas costales basales a la bulla presentes, dos bandas en los fémures ............................................................................ 3 Areas costal y subcostal hialinas, venas transversas costales basales a la bulla ausentes, una banda en los fémures ...................................................... Thraulodes regulus

This species is apparently very similar to T. trijunctus (BANKS), but can be separated from it by the presence in T. laetus of two small maculae near the lateral margin of the terga (“isolated spots” according to TRAVER). As it was not possible to see the type nor an illustration, T. trijunctus is excluded from the key. 33 The differences among species sometimes are very subtle. For this reason it is safer to rely on adults. 34 No incluimos en esta clave Thraulodes colombiae conocido solo de hembras, ni T. limbatus, T. subfasciatus, y T. vitripennis debido a que las descripciones son insuficientes. 32

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Vena ICu2 de las alas anteriores (Fig. 187D) unida basalmente a CuP; espina de cada lóbulo de los penes (Fig. 188K) delgada y larga .................. Thraulodes liminaris – Vena ICu2 de las alas anteriores no unida basalmente a CuP (como en la Fig. 187B); espina de cada lóbulo los penes ancha y robusta (Fig. 187P) ...........Thraulodes consortis 4(1) Areas costal y subcostal castañas, especialmente en el 1/3 apical; una gran mancha rojiza en cada lado del tergo VII; penes como en la Fig. 189L ............... Thraulodes venezuelana – Areas costal y subcostal amarillentas o translúcidas; sin grandes manchas rojizas en el tergo VII .................................................................................................................... 5 5(4) Una banda en el fémur anterior ...................................................................................... 6 – Dos bandas o una banda y una mancha en el fémur anterior ................................ 14 6(5) Areas costal y subcostal amarillenta-clara ..................................................................... 7 – Areas costal y subcostal hialinas ..................................................................................... 8 7(6) Tres a cuatro débiles transversas costales basales a la bulla; penes (Fig. 188H) muy delgados, sin bolsillo lateral, pliegue recurvado presente sólo en la mitad basal de los penes .............................................................................................. Thraulodes itatiajanus – Sin transversas costales basales a la bulla; penes (Fig. 189E) no muy delgados, bolsillo lateral presente, pliegue recurvado paralelo al margen y llegando hasta el extremo del margen interno ............................................................. Thraulodes schlingeri 8(6) Penes relativamente largos y delgados, sin bolsillo lateral (Figs. 188R, 189G) ............ 9 – Penes robustos, bolsillo lateral presente (Figs. 188P, 189K) ................................... 10 9(8) Tergos abdominales I y VII-X castaños, II-VI blanquecino-translúcidos, estos últimos con dos pequeñas manchas negruzcas cerca del ángulo anterolateral (Fig. 189F) ....... ............................................................................................................ Thraulodes telegraphicus – Tergos abdominales II-VI y base de VII blanquecino-transparentes, resto de VII amarillento, VIII-X castaño-amarillento, pálidas manchas grisáceas dorsolaterales en segmentos II-VII (Fig. 188Q) .................................................... Thraulodes papilionis 10(8) Sin transversas costales basales a la bulla ................................................................... 11 – Cuatro débiles transversas costales basales a la bulla ............................................... 12 11(10) Tergos abdominales II-V translúcidos, con el área central de II-IV teñidas de rojizo, dos manchas negruzcas cerca de los márgenes anteriores de los segmentos II-VIII (Fig. 188L ), tergos VI-X castaños con los márgenes anteriores de los segmentos VIVII y margen posterior y esquinas posterolaterales del segmento VIII con áreas blanquecinas; genitalia como en la Fig. 188M ................................. Thraulodes marreroi – Tergos abdominales II-V y 1/3 basal de VI translúcido con bandas transversas castaño-rojizas, áreas intersegmentales y la región adyacente a la unión con las pleuras amarillo-blanquecino claro, 2/3 posteriores del tergo VI castañas, más fuerte que II-V, pero más pálido que VIII-X, dos manchas más bien tenues cerca del margen anterior de los tergos II-V; genitalia como en la Fig. 188P ................... ........................................................................................................................Thraulodes osiris 12(10) Alas anteriores mayores a los 12 mm; mitad basal de las venas longitudinales principales amarillentas, más oscuras hacia el ápice; genitalia como en la Fig. 188O ...................................................................................................................... Thraulodes mucuy – Alas anteriores menores a los 8 mm; venas amarillentas, genitalia no como arriba (Figs. 188G, 189K) ........................................................................................................... 13

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13(12) Tergos abdominales VIII-X y 1/2 posterior de VII castaño-rojizo oscuro, márgenes anteriores de VII y VIII negruzcos; genitalia como en la Fig. 189K ..... ...................................................................................................................... Thraulodes ulmeri – Tergos abdominales VII-X castaño-anaranjados, con areas blanquecinas en los segmentos VII-VIII y X (Fig. 188F); genitalia como en la Fig. 188G ...................... ................................................................................................................... Thraulodes guanare 14(5) Una banda completa y una incompleta en el fémur anterior ................................. 15 – Dos bandas en el fémur anterior .................................................................................. 16 15(14) Tres o cuatro delicadas transversas costales basales a la bulla; tergos abdominales IIVI translúcidos, con manchas como en la Fig. 188C; genitalia como en la Fig. 188D ........................................................................................................................ Thraulodes flinti – Sin transversas costales basales a la bulla; tergos abdominales II-V blanquecinotranslúcidos con manchas como en la Fig. 187M; genitalia como en la Fig. 187N ............................................................................................................ Thraulodes cochunaensis 16(14) Areas costal y subcostal amarillenta; 5 a 8 transversas costales basales a la bulla .............................................................................................................................................. 17 – Areas costal y subcostal sin pigmentación; transversas costales basales a la bulla ausentes .............................................................................................................................. 18 17(16) Tergos abdominales II-VII con manchas en forma de U (Fig. 187K) formadas por dos largas manchas paralelas, convergentes en dirección posterior; genitalia como en la Fig. 187L ................................................................................... Thraulodes bomplandi – Tergos abdominales II-VI con manchas triangulares mediales cerca del margen posterior y manchas submediales cerca del margen anterior; genitalia como en la Fig. 188E ............................................................................................... Thraulodes furficulus 18(16) Especie grande, alas anteriores más de 10,5 mm de longitud; patrón de coloración abdominal como en la Fig. 187I ..................................................... Thraulodes bolivianus – Especies pequeñas, alas anteriores 8 mm o menos de longitud; patrón de coloración abdominal no como en el caso precedente (Figs. 188A, 189A, H) .............................................................................................................................................. 19 19(18) Tergos abdominales II-VII translúcidos, con manchas medianas pequeñas como en la Fig. 188A; penes (Fig. 188B) con una proyección lateral en la base de cada lóbulo ..................................................................................................... Thraulodes daidaleus – Tergos abdominales II-VI translúcidos, patrón de coloración abdominal no como en el caso precedente; penes sin proyecciones laterales .......................................... 20 20(19) Tergos abdominales II-VI con manchas medianas o posteromedianas (Figs. 189A, H) ......................................................................................................................................... 21 – Tergos abdominales II-VI sin manchas o con pequeñas manchas submedianas ... ................................................................................................................... Thraulodes laetus 35

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Esta especie es aparentemente muy semejante a T. trijunctus (BANKS), de la que sería separable por la presencia en T. laetus de 2 pequeñas manchas cercanas al margen lateral de los tergos (“isolated spots” según TRAVER). Como no fué posible ver los tipos ni existe ninguna ilustración disponible, preferimos dejar por el momento a T. trijunctus fuera de la clave.

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21(20) Tergos abdominales III-V con manchas anchas que alcancanzan el margen posterior y que se unen con las manchas laterales (Fig. 189A); genitalia como en la Fig. 189B .......................................................................................... Thraulodes paysandensis – Tergos abdominales III-V con manchas separadas o unidas a la mancha submediana, no alcanzando el margen posterior y separadas de las manchas laterales(Fig. 189H); genitalia como en la Fig. 189I ....................... Thraulodes traverae Ninfas

36

1

Branquias abdominales anchas, que se afinan abruptamente cerca del ápice (Figs. 191G-H) ............................................................................................................................... 2 Branquias abdominales delgadas, que se van afinando gradualmente hacia el ápice (Fig. 191F) ............................................................................................................................ 6 Branquias abdominales con traquea principal ramificada (Fig. 191G) ................... 3 Branquias abdominales con traquea principal sin ramificaciones (Fig. 191H) ..... 4 Uña tarsal I con 6 dentículos que se van agrandando paulatinamente hacia el ápice (Fig. 191J) ............................................................................................... Thraulodes liminaris Uña tarsal I con 6 ó 7 dentículos que se van agrandando paulatinamente hacia el ápice, excepto el distal que es levemente mayor (Fig. 191I) ........Thraulodes consortis Especies pequeñas, longitud del cuerpo de la ninfa madura menor a 7 mm ......... ................................................................................................................... Thraulodes traverae Especie grande, longitud del cuerpo de la ninfa madura mayor de 10 mm .......... 5 Fémures con mancha castaña mediana ......................................... Thraulodes bolivianus Fémures con manchas castañas mediana y apical ...................... Thraulodes itatiajanus Uña tarsal con seis ó siete dentículos que se van agrandando paulatinamente hacia el ápice .............................................................................................. Thraulodes cochunaensis Uña tarsal con tres dentículos grandes y tres ó cuatro más pequeños ..................... ................................................................................................................. Thraulodes daidaleus

– 2(1) – 3(2) – 4(2) – 5(4) – 6(1) –

Thraulodes bolivianus DOMÍNGUEZ Thraulodes bolivianus DOMÍNGUEZ, 1986: 149 (male, female, nymph).

Taxonomy: Thraulodes bolivianus is known from male and female imagos and nymphs. This species can be differentiated from the other species of the genus by the following combinations of characters. In the imago: 1) Costal membrane basal to bulla without cross veins (Fig. 187B); 2) costal and subcostal areas translucent; 3) fore femora with basal band and a broader median band occupying middle third of femur; 4) abdominal color pattern as in 187I; and 5) apicolateral area of each penis lobe of male genitalia (Fig. 187J) not forming “ear”, spine short and robust, lateral pouch present on lateral margin and recurved, widened angularly near apex. In the nymph: 1) Abdominal gills (Fig. 191H) without secondary tracheae; and 2) apical denticle of fore tarsal claws slightly larger than others. 36

Las diferencias entre las especies son a veces muy sutiles. Por esta razón es más confiable utilizar los adultos.

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497

D

E A C G I

F H

K J

L

M

O

N

P

Q

R

Plate 188. Thraulodes, imagos. Figs. 188A-R. 188A-B: T. daidaleus. 188A, abdominal terga IV-VII; 188B, genitalia, v.v. 188C-D: T. flinti. 188C, abdominal terga II-VII; 188D, genitalia, v.v. 188E: T. furficulus, genitalia, v.v. 188F-G: T. guanare. 188F, abdominal terga II-VII; 188G, genitalia, v.v. 188H: T. itatiajanus, genitalia, v.v. 188I: T. laetus, genitalia, v.v. 188J-K: T. liminaris. 188J, abdominal terga II-VII; 188K, genitalia, v.v. 188L-M: T. marreroi. 188L, abdominal terga II-VII; 188M, genitalia, v.v. 188N-O: T. mucuy. 188N, abdominal terga II-VII; 188O, penes, v.v. 188P: T. osiris, genitalia, v.v. 188Q-R: T. papilionis. 188Q, abdominal terga II-VII; 188R, genitalia, v.v. Figs. 188A-B, E, H and P-R modified from TRAVER & EDMUNDS, 1967; 188C-D and JK from DOMÍNGUEZ, 1987a; 188F-G, and L-O from CHACON et al., 1999 and 188I from EATON, 1884.

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Distribution and Biology: Bolivia (Cochabamba).The nymphs were collected in the Pasare river, at 2000 m in the Yungas of Bolivia, which is a large river about 10 m wide with large rocks. The nymphs were found below small stones in the river margins and in backwater zones. The adults were collected in October. Thraulodes bomplandi (ESBEN-PETERSEN) Thraulus bomplandi ESBEN-PETERSEN, 1912: 336 (male). Thraulus bonplandi [sic]; NAVÁS, 1917: 188 (female). Thraulodes bomplandi; ULMER, 1920a: 34; ULMER, 1920c: 116; ULMER, 1921: 263; NEEDHAM & MURPHY, 1924: 41; ULMER, 1943: 24 (male); TRAVER, 1946: 433; TRAVER, 1959b (misidentified, see T. daidaleus); TRAVER & EDMUNDS, 1967: 362 (male); HUBBARD, 1982A: 266; DOMÍNGUEZ, 1987a: 49 (male); HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 220; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 73.

Taxonomy: This species is known only from adults and can be differentiated from the other species of the genus by the following combination of characters: 1) Costal membrane basal to bulla with 5-8 delicate cross veins; 2) costal and subcostal zones shaded yellowish; 3) femora with two bands; 4) abdominal terga II-VI with U-shaped maculae and some smaller comma-shaped maculae near U-shaped maculae (Fig. 187K); and 5) male genitalia as in Fig. 187L. Distribution: Argentina (Bompland, Misiones Prov.). Thraulodes cochunaensis DOMÍNGUEZ Thraulodes cochunaensis DOMÍNGUEZ, 1987a: 51 (male, female, nymph, egg); HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 220; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 74.

Taxonomy: This species is known from all stages and can be differentiated from the other species of the genus by the following combination of characters. In the imago: 1) Costal membrane basal to bulla without cross veins; 2) costal and subcostal zones translucent; 3) fore femora with one complete apical band and one incomplete band in medial zone; 4) abdominal terga VI-X brown and terga II-V translucent whitish with blackish maculae (Fig. 187M); and 5) apicolateral area of penes not “ear”-shaped, spine very short and robust, without lateral pouch, and with recurved fold subparallel to inner margin (Fig. 187N). In the nymph: 1) Abdominal gills (Fig. 191F) slender with main trachea tinted blackish, unbranched; and 2) abdominal color pattern as in imago (Fig. 187M). Distribution and Biology: Northeastern Argentina (Provinces of Tucumán, Salta and Jujuy) and Southern Bolivia. This species is very common and is found widely distributed in Northwestern Argentina and Southern Bolivia. It is usually encountered associated with Thraulodes consortis. The nymphs are normally found under rocks in shallow areas of a variety of clean rivers, at altitudes ranging from 400 to 1700 m a.s.l. Adults have been collected from October to May, but probably emerge throughout most of the year. The subimagos emerge at dusk.

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Thraulodes colombiae (WALKER) Ephemera colombiae WALKER, 1853: 537. Palingenia? HAGEN, 1861: 304. Leptophlebia colombiae; EATON, 1871: 84; EATON, 1873: 395. Adenophlebia colombiae; EATON, 1881: 194. Thraulus colombiae; EATON, 1884: 110 (female). Thraulodes colombiae; ULMER, 1920a: 34; ULMER, 1920c: 116; LESTAGE, 1924c: 43; NEEDHAM & MURPHY, 1924: 40; TRAVER, 1946: 436; KIMMINS, 1960: 302; TRAVER & EDMUNDS, 1967: 363 (female); HUBBARD, 1982a: 266.

Taxonomy: Thraulodes colombiae, known from one female subimago, has been placed in six different genera to date, probably because the male is still unknown and the type has not been illustrated except for the unusual hind wing. It is unlikely that it belongs to Thraulodes. It can be characterized by the following combination of characters: 1) Costal membrane basal to bulla with 1 or 2 weak cross veins; 2) costal and subcostal areas hyaline; 3) femora without bands; and 4) abdominal terga yellowish red with blackish posterior margins on segments I-VIII and wide black oblique rays on terga II-VII. The female is distinct because of the shape and location of the costal projection of hind wing (Fig. 187F) and its large size (fore wings= 15 mm) (TRAVER, 1946). Distribution: Colombia. Thraulodes consortis DOMÍNGUEZ Thraulodes consortis DOMÍNGUEZ, 1987a: 55 (male, female, nymph, egg); HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 220; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 74.

Taxonomy: This species is known from all stages and can be differentiated from the other species of the genus by the following combination of characters. In the imago: 1) Costal membrane basal to bulla with 6-8 cross veins; 2) costal and subcostal areas shaded brown; 3) fore femora with two bands; 4) abdominal terga orange- brown, with blackish macula situated mid-way between median dorsal line and the lateral border (in some cases separated into two independent maculae) (Fig. 187O); and 5) each penis lobe with apicolateral area forming an “ear”, spine short and robust, without lateral pouch, and with recurved folds parallel to inner margins (Fig. 187P). In the nymph: 1) Abdominal gills broad, grayish, main tracheae blackish, lateral tracheae less distinct (Fig. 191G); 2) apical denticle of fore tarsal claw somewhat larger than other denticles (Fig. 191I); and 3) abdominal color pattern as in imago (Fig. 187O). Distribution and Biology: Argentina (Provinces of Tucumán, Salta and Jujuy) and Southern Bolivia. This species is widely distributed in Northwest Argentina and Southern Bolivia. It is found in different types of rivers, most commonly with rocky bottoms and clean water. Nymphs can be found throughout the year. Nuptial flight has never been observed, but a pair in copula was captured flying around 16:00 h. The subimagos emerge around 18:00 h.

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B

A

C

G

E

D

F

J

H

K

I

L

Plate 189. Thraulodes, imagos. Figs. 189A-L. 189A-B: T. paysandensis. 189A, abdominal terga II-VII; 189B, genitalia, v.v. 189C: T. regulus, genitalia, v.v. 189D-E: T. schlingeri. 189D, abdominal terga II-VII; 189E, genitalia, v.v. 189F-G: T. telegraphicus. 189F, abdominal terga II-VII; 189G, genitalia, v.v. 189HI: T. traverae. 189H, abdominal terga II-VII; 189I, genitalia, v.v. 189J: T. trijunctus, genitalia, v.v. 189K: T. ulmeri, genitalia, v.v. 189L: T. venezuelana, genitalia, v.v. Figs. 189C, E and I-K modified from TRAVER & EDMUNDS, 1967; 189L from ULMER, 1943.

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Thraulodes daidaleus THEW Thraulodes bomplandi; TRAVER, 1959: Fig.12 (misidentified). Thraulodes daidaleus THEW, 1960b: 119 (male, female); TRAVER & EDMUNDS, 1967: 364 (male, female, nymph); HUBBARD, 1982a: 267; HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 220; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 75.

Taxonomy: This species, known from adults and nymphs, can be differentiated from the other species of the genus by the following combination of characters. In the imago: 1) Costal membrane basal to bulla without cross veins; 2) costal and subcostal areas without pigmentation; 3) femora with two bands; 4) abdominal terga (Fig. 188A) II-VII translucent and median line whitish, with triangular blackish markings restricted to posterior part of segment II-V and extended to anterior border of segments VII-X, and a macula near anterior border and half-way between median line and lateral border, with another similar macula situated laterally; and 5) outer lateral projection at base of each penis lobe (Fig. 188B). In the nymph: 1) Tarsal claws with 3 large and 4 small denticles; 2) abdominal gills slender, tracheae unbranched; and 3) abdominal color pattern as in imago (Fig. 188A). Distribution and Biology: Brazil and Uruguay. The adults were collected flying in January. Thraulodes flinti DOMÍNGUEZ Thraulodes flinti DOMÍNGUEZ, 1987a: 57 (male); HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 220; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 75.

Taxonomy: This species, known only from imagos, can be differentiated from the other species of the genus by the following combination of characters: 1) Costal membrane basal to bulla with 4 very delicate cross veins; 2) costal and subcostal areas without pigmentation; 3) one macula and one band on fore femur; 4) abdominal segments I and VII-X yellowish brown and remainder translucent with maculae as in Fig. 188C; and 5) outer border of penes without lateral pouch, and recurved fold very tenuous and parallel to inner margin (Fig. 188D). Distribution and Biology: Northeastern Argentina (Misiones Prov.). The adults have been collected in November. Thraulodes furficulus TRAVER Thraulodes furficulus TRAVER, 1946: 435 (male); THEW, 1960b: 122; TRAVER & EDMUNDS, 1967: 365 (male); HUBBARD, 1982a: 267.

Taxonomy: Thraulodes furficulus is known only from male imagos which can be differentiated from the other species of the genus by the following combination of characters: 1) Costal membrane basal to bulla with 7 cross veins; 2) costal area shaded yellowish; 3) two bands on femora; 4) abdominal terga I and VII-X reddish brown, II-VI transparent yellow with median triangles near posterior border of each segment and submedian reddish brown maculae near anterior border; and 5) penis lobes (Fig. 188E) with short broad spine and recurved fold broader near apex. Distribution: Guyana (Georgetown).

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Thraulodes guanare CHACÓN, SEGNINI & DOMÍNGUEZ Thraulodes guanare CHACÓN, SEGNINI & DOMÍNGUEZ, 1999: 252 (male, female).

Taxonomy: This species, known from adults of both sexes, can be differentiated from the other species of the genus by the following combination of characters: 1) Costal membrane basal to bulla with 4 cross veins; 2) costal and subcostal areas without pigmentation; 3) main longitudinal veins yellowish; 4) one band on femora; 5) abdominal terga II- VI yellowish-translucent, with two light blackish spots close to anterior margins on segments IIIVI, terga IV- VI slightly washed with brown in medial area; terga VII- X orange-brown, with whitish areas on segments VII -VIII and X (Fig. 188F); 6) penis lobes (Fig. 188G) relatively long and stout, spine long and narrow; apicolateral area forming an “ear”; external margin forming lateral pouch, and recurved fold parallel to internal margin, not forming a “lapel”. Distribution and Biology: Venezuela (Edo Portuguesa). The adults of this species were attracted to light in December, along with specimens of T. marreroi, Campsurus sp., Leptohyphes sp, and Tricorythodes sp. at the margins of the Guanare river. This river is approximately 30 m wide, with warm waters, and located in a tropical dry forest at 200 m altitude. Thraulodes itatiajanus TRAVER & EDMUNDS Thraulodes itatiajanus TRAVER & EDMUNDS, 1967: 368 (male); HUBBARD, 1982a: 267; DA-SILVA, 2003: 5 (nymph).

Taxonomy: Thraulodes itatiajanus was described only from male imagos, but recently (DA-SILVA, 2003), described the nymph which he associated with the imago by body size, distribution, and abdominal color pattern. This species can be differentiated by the following combination of characters. In the imago: 1) Costal membrane basal to bulla with 3-4 very weak cross veins; 2) costal area weakly tinted yellowish; 3) one band on femora; 4) anterior half of abdominal tergum I blackish brown, the posterior half and segments II-VI translucent yellowish, and terga VII-X pink; large submedian maculae near anterior border of segments III-IV, reduced to one line on segment II and replaced by a series of dark dots on segments V-VI; small stigmatic maculae on segments II-VII with a darker macula near these on segments III-VII; and 5) penes (Fig. 188H) very slender and recurved fold present only on the basal half of penes. In the nymph: 1) Abdominal gills broad, grayish, main tracheae darkened, lateral tracheal branches not visible; 2) femora with median and apical brown spots; 3) abdominal terga dark brown on anterior margins, light brown on posterior margins, terga IX and X dark brown medially near posterior margin (after DA-SILVA, 2003). Distribution: Brazil (Itatiaia National Park). Thraulodes laetus (EATON) Calliarcys laetus EATON, 1883: pl. 13. Thraulus laetus; EATON, 1884: 110 (male). Thraulodes laetus; ULMER, 1920c: 116; ULMER, 1921: 263; NEEDHAM & MURPHY, 1924: 41; KIMMINS, 1960: 302; TRAVER & EDMUNDS, 1967: 369 (male); HUBBARD, 1982a: 267. Thraulodes loetus [sic]; LESTAGE, 1924c: 43. Thraulodes laeta; ULMER, 1943: 24.

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Taxonomy: Thraulodes laetus was provisionally described in Calliarcys by EATON (1883) and then in Thraulus (EATON 1884). ULMER (1920) transferred it to Thraulodes as the type species of the genus. Thraulodes laetus is known only from male imagos and can be differentiated from the other species of the genus by the following combination of characters: 1) Costal membrane basal to bulla without cross veins; 2) costal and subcostal

A

Plate 190. Thraulodes consortis. Fig. 190A, nymphal habitus. Drawing by A. Dupuy.

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areas without pigmentation; 3) two bands on femora; 4) abdominal segments II-VI whitish, with “a series of small triangular spots on each side of the dorsum” (EATON, 1884), segments VII-X blackish red dorsally; and 5) penes as in Fig. 188I. Distribution: Colombia. Thraulodes limbatus NAVÁS Thraulodes limbatus NAVÁS, 1936b: 733 (male); TRAVER & EDMUNDS, 1967: 351; HUBBARD, 1982a: 267.

Taxonomy: Thraulodes limbatus was briefly described by NAVÁS (1936b) and the type is probably lost so it is impossible to characterize the species. Without explanation, TRAVER & EDMUNDS (1967) suggested that it belonged to Ulmeritus. Distribution: Southern Brazil (Sta. Catarina). Thraulodes liminaris DOMÍNGUEZ Thraulodes liminaris DOMÍNGUEZ, 1987a: 60 (male, female, nymph, egg); HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 221; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 76.

Taxonomy: The nymphs of T. liminaris are difficult to separate from those of Thraulodes consortis, so it is imperative to consider the adults to obtain a correct determination. Thraulodes liminaris can be differentiated from the other species of the genus by the following combination of characters. In the imago: 1) Costal membrane basal to bulla with 11-14 cross veins; 2) costal and subcostal areas shaded reddish brown; 3) vein ICu2 of fore wing joined at base to vein CuP (Fig. 187D); 4) two bands on femora; 5) abdominal terga II-V translucent whitish in medial zone surrounded by an orangebrown area with a lateral branch which extends laterally to spiracle (Fig. 188J); and 6) apicolateral area of each penis lobe (Fig. 188K) forming an “ear”, spine long and slender, outer border without lateral pouch, and recurved fold parallel to inner margin. In the nymph: 1) Abdominal gills broad and violet gray with main trachea darker than lateral tracheal branches; 2) denticles on tarsal claw of fore leg (Fig. 191J) enlarged gradually toward apex; and 3) abdominal color pattern as in adult (Fig. 188J). Distribution and Biology: Northwestern Argentina (Salta Prov.). Thraulodes liminaris is restricted to rocky streams with clear water. The nymphs are found in large numbers and great density on stones in zones above 1400 m. Thraulodes marreroi CHACÓN, SEGNINI & DOMÍNGUEZ Thraulodes marreroi CHACÓN, SEGNINI & DOMÍNGUEZ, 1999: 255 (male).

Taxonomy: This species, known from male imagos, can be differentiated from the other species of the genus by the following combination of characters: 1) Costal membrane basal to bulla without cross veins; 2) costal and subcostal areas without pigmentation; 3) longitudinal veins of fore wings yellowish; 4) apical 1/5 of femora covered by a tricolored spot: black basally, reddish in middle and light brown close to apex; 5) terga II-V translucent, with central area of segments II-IV tinged with red; black spots close to anterior margins on segments II-VII; terga VI-X reddish, with anterior

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margins of segments VI-VII, and posterior margin and posterolateral corners of segment VIII with white areas (Fig. 188L); and 6) penes (Fig. 188M) long, stout, each lobe with long and narrow spine, apicolateral area forming a small “ear”, external margin forming a small lateral pouch and recurved fold slightly angulated.

B

A

C

III

D

II

E

I

D

F

G

H

I

J

Plate 191. Thraulodes, nymphs. Figs. 191A-J. 191A-E: mouthparts, Thraulodes consortis. 191A, labrum, d.v.; 191B, left mandible; 191C maxilla; 191D, hypopharynx; 191E, labium (left dorsal, right ventral); 191F-H: gill II. 191F, T. cochunaensis; 191G, T. consortis; 191H, T. bolivianus. 191I-J: fore tarsal claw. 191I, T. consortis; 191J, T. liminaris. (D = denticle; Roman numbers = segment numbers). Figs. 191AJ modified from DOMÍNGUEZ, 1987a.

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Distribution and Biology: Venezuela (Edo Portuguesa). The adults of this species were attracted by light in December, along with specimens of T. guanare, Campsurus sp., Leptohyphes sp, and Tricorythodes sp. at the margins of the Guanare river. This river is approximately 30 m wide, with warm waters, and is located in a tropical dry forest at 200 m altitude. Thraulodes mucuy CHACÓN, SEGNINI & DOMÍNGUEZ Thraulodes mucuy CHACÓN, SEGNINI & DOMÍNGUEZ, 1999: 250 (male).

Taxonomy: This species, known from male imagos, can be differentiated from the other species of the genus by the following combination of characters: 1) Costal membrane with 4 cross veins basal to bulla; 2) costal and subcostal areas without pigmentation; 3) basal 1/2 of main longitudinal veins of fore wings yellowish; 4) fore and hind femora with one band; 5) abdominal terga II-VI yellowish, translucent, washed with brown on central area; two brownish black spots on anterior part of segments III-VI; terga VII-IX reddish brown, with lateral areas of VII-VIII whitish and lateral margins of IX yellowish translucent (Fig. 188N); 6) penes short; each penis lobe with short, stout spine (Fig. 188O), external margin forming lateral pouch and recurved fold forming “lapel”. Distribution and Biology: Venezuela (Edo Merida). Males were collected in December at a light trap. The light was near a turbulent, clear, mountain forest stream (altitude 2,000 m) approximately 5 m wide with water temperature about 12 °C. Thraulodes osiris TRAVER & EDMUNDS Thraulodes osiris TRAVER & EDMUNDS, 1967: 371 (male, female); HUBBARD, 1982a: 267.

Taxonomy: Thraulodes osiris is known from imagos of both sexes and can be differentiated from the other species of the genus by the following combination of characters: 1) Costal membrane basal to bulla without cross veins; 2) fore wings of male hyaline, those of female weakly tinted yellowish; 3) one band on femora (in the only female specimen with legs); 4) abdominal segments I and VII-X reddish brown and II-VI translucent, with pale transverse reddish brown bands, bands weaker on sterna than on terga; and 5) male genitalia as in Fig. 188P, with recurved fold apparently parallel to inner margin, lateral pouch present, and a short broad spine. Distribution: Ecuador (Napo). Thraulodes papilionis TRAVER & EDMUNDS Thraulodes papilionis TRAVER & EDMUNDS, 1967: 374 (male); HUBBARD, 1982a: 267.

Taxonomy: Thraulodes papilionis is known only from male imagos and can be distinguished from the other species of the genus by the following combination of characters: 1) Costal membrane basal to bulla without cross veins; 2) costal and subcostal areas without pigmentation except very close to wing base; 3) femora with one band; 4) abdominal segments II-VI and anterior portion of VII transparent whitish, VII yellowish, VIII-X yellowish brown, pale grayish dorsolateral maculae on terga II-VII, weak mediodorsal line with oblique submedian maculae on III-IV and dark stigmatic maculae

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on II-VII (Fig. 188Q); and 5) penes (Fig. 188R) slender, long; each lobe with recurved fold broader medially and long broad spines. Distribution: Colombia (Tolima). Thraulodes paysandensis TRAVER Thraulodes paysandensis TRAVER, 1964: 33 (male, female); TRAVER & EDMUNDS, 1967: 374 (male, female); HUBBARD, 1982a: 267; DOMÍNGUEZ, 1987a: 64 (male, female, egg); HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 221; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 76.

Taxonomy: Thraulodes paysandensis is known only from imagos and can be distinguished from the other species of the genus by the following combination of characters: 1) Costal membrane basal to bulla without cross veins; 2) costal and subcostal areas without pigmentation; 3) femora with two bands; 4) abdominal terga (Fig. 189A) translucent, I and II almost totally tinted grayish brown, posterior part of segments III-VI colored and extreme lateral area of this zone with a circular area occupying most of the width of segment; and 5) apicolateral area of each penis lobe (Fig. 189B) not forming “ear”, spine long and slender, lateral pouch present, and recurved fold broadened angularly near apex. Distribution and Biology: Northwestern Argentina (Misiones and Entre Rios Provs.) and Uruguay (Paysandú, Artigas y Salto). Adults were collected between November and January. This is probably the most abundant species of Thraulodes in the area. Thraulodes regulus TRAVER & EDMUNDS Thraulodes regulus TRAVER & EDMUNDS, 1967: 375 (male); HUBBARD, 1982a: 267.

Taxonomy: This species, known only from male imagos, can be distinguished from the other species of the genus by the following combination of characters: 1) Costal membrane basal to bulla without cross veins; 2) costal and subcostal areas without pigmentation except for a black macula near wing base; 3) femora with one band; 4) abdominal tergum I blackish brown, terga II-X with broad transverse brown bands, bands darker on II-IV, somewhat weaker on VI-VIII and pale on V; sublateral maculae on terga III-V; and 5) penes (Fig. 189C) rather short, each lobe with a lateral pouch, recurved fold widened medially, and large stout spines. Distribution: Peru (Huallaga Prov.). Thraulodes schlingeri TRAVER & EDMUNDS Thraulodes schlingeri TRAVER & EDMUNDS, 1967: 376 (male, female, egg); HUBBARD, 1982a: 267; FERREIRA & FROEHLICH, 1992: 541; LOPES, FROEHLICH & DOMÍNGUEZ, 2003: 197 (nymph).

Taxonomy: Thraulodes schlingeri was known originally from imagos of both sexes, and the nymph was described recently by LOPES et al., 2003. This species can be distinguished from the other species of the genus by the following combination of characters. In the imago: 1) Costal membrane basal to bulla without cross veins; 2) fore wings weakly colored yellowish; 3) femora with one band; 4) abdominal segments II-V translucent yellowish, VI-X pink; with submedian grayish maculae on segments II-V, and prominent stigmatic maculae on segments I-VII (Fig.

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189D); 5) penis lobes (Fig. 189E) each with a lateral pouch, a recurved fold parallel to inner margin, and slender curved spines. In the nymphs: 1) Abdominal gills broad with well developed tracheal branches (similar to Fig. 191G); 2) tarsal claws with 8 to 9 denticles, progressively larger apically; and 3) abdominal color pattern similar to adult (Fig. 189D). Distribution and Biology: Peru (Huallaga Prov.) and Brazil (São Paulo State). In Brazil, the nymphs were collected in riffles in a stony bottom stream at 700 m. Water temperature fluctuated from 14ºC (July) to 24ºC (November). Nymphs were present all months of the year. Nymphal growth was rapid and the cycle was completed in approximately two months. Adults were present all year, except in September. Thraulodes subfasciatus NAVÁS Thraulodes subfasciatus NAVÁS, 1934b: 164; TRAVER & EDMUNDS, 1967: 351; HUBBARD, 1982: 267.

Taxonomy: The original description of Thraulodes subfasciatus is very incomplete and does not include figures of the genitalia. Because the type specimen could not be located, it is impossible to characterize the species at this time. According to TRAVER & EDMUNDS (1967) this species probably belongs to Ulmeritus. Distribution: Brazil (Rio de Janeiro). Thraulodes telegraphicus NEEDHAM & MURPHY Thraulodes telegraphicus NEEDHAM & MURPHY, 1924: 41 (male, female); TRAVER, 1946: 428 (male); EDMUNDS, 1950b: 554; TRAVER & EDMUNDS, 1967: 379 (male, female); HUBBARD, 1982a: 267. Thraulodes plicatus NEEDHAM & MURPHY, 1924: 42 (male, female).

Taxonomy: Thraulodes telegraphicus is known from imagos of both sexes and can be distinguished from the other species of the genus by the following combination of characters: 1) Costal membrane basal to bulla without cross veins; 2) costal and subcostal areas without pigmentation, except near wing base; 3) femora with one band; 4) abdominal terga I and VII-X brown, remaining terga translucent whitish; terga with two small blackish maculae situated near the anterolateral angle (Fig. 189F); 5) apicolateral area of each penis lobe (Fig. 189G) forming an “ear”, spine long and slender, outer border without lateral pouch, and recurved fold present subparallel to inner margin. Distribution: Peru (Dept. Junín). Thraulodes traverae THEW Thraulodes traverae THEW, 1960b: 120 (male, female); TRAVER & EDMUNDS, 1967: 380 (male, female, nymph); HUBBARD, 1982a: 267.

Taxonomy: Thraulodes traverae, known from imagos of both sexes and nymphs, can be differentiated from the other species of the genus by the following combination of characters. In the imago: 1) Costal membrane basal to bulla without cross veins; 2) costal and subcostal areas without pigmentation; 3) femora with two bands; 4) abdominal terga I and VI-X yellowish brown, II-VI translucent yellowish; two blackish maculae situated near posterior margins of terga I-VII, one on each side of median line, and blackish maculae at posterolateral angles and anterolaterally

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(Fig. 189H); and 5) apicolateral area of each penis lobe forming an “ear”, spine long and slender, and outer border forming a lateral pouch (Fig. 189I). In the nymph: 1) Abdominal gills broad, trachea unbranched; 2) seven denticles on tarsal claws progressively increasing in size apically; and 3) abdominal color pattern as in male imago (Fig. 189H). Distribution and Biology: Southern Brazil (Santa Catarina) and Uruguay (Artigas). The adults were collected from September to January. Thraulodes trijunctus (BANKS) Thraulus trijunctus BANKS, 1918: 10. Thraulodes trijuncta; ULMER, 1943: 28; TRAVER, 1960b: 73. Thraulodes trijunctus; THEW, 1960b: 120; TRAVER & EDMUNDS, 1967: 381 (male, female); HUBBARD, 1982a: 267.

Taxonomy: Thraulodes trijunctus, originally described by BANKS (1918) from one male imago, was later redescribed by TRAVER & EDMUNDS (1967) using males and females collected in Arequipa (Peru). T. trijunctus is not distinguished in the key from T. laetus (see footnote 32) but may possibly be distinguished by the following combination of characters: 1) Costal membrane basal to bulla without cross veins; 2) costal and subcostal areas without pigmentation; 3) femora with two bands; 4) abdominal terga II-VI translucent yellowish, tergum I dark brown, terga VII-X with dark brown maculae occupying the greater part of each segment; 5) penes (Fig. 189J) short, each lobe with lateral pouch, recurved fold broader apically, and spine short and robust. Distribution: Peru (Lima). Thraulodes ulmeri EDMUNDS Thraulodes ulmeri EDMUNDS, 1950b: 552 (male, female); TRAVER & EDMUNDS, 1967: 381 (male, female); HUBBARD, 1982a: 267.

Taxonomy: Thraulodes ulmeri is known from imagos of both sexes and can be distinguished by the following combination of characters: 1) Costal membrane basal to bulla with 4 cross veins; 2) costal and subcostal areas without pigmentation except near wing base; 3) femora with one band; 4) abdominal segments II-VI translucent yellowish, VII-X reddish brown; prominent submedian maculae near anterior margin of terga, maculae larger on segments III-V, and stigmatic maculae dark; and 5) each penis lobe (Fig. 189K) with lateral pouch, recurved fold parallel to inner margin, and short spine. Distribution: Brazil (Santa Catarina). Thraulodes venezuelana ULMER Thraulodes venezuelana ULMER, 1943: 23 (male); TRAVER & EDMUNDS, 1967: 382 (male); HUBBARD, 1982a: 267.

Taxonomy: Thraulodes venezuelana, known only from a male imago, can be distinguished by the following combination of characters: 1) Costal membrane basal to bulla with 2 or 3 cross veins; 2) costal and subcostal areas shaded with brown, especially in apical third; 3) femora with two bands; 4) abdominal terga brown, except segments III-VI translucent with posterior margin blackish; rounded submedian maculae near anterior margin of segments

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III-VI, large blackish red maculae laterally on tergum VII, and blackish brown stigmatic maculae on segments II-VI; and 5) penes as in Fig. 189L (after ULMER, 1943). Distribution: Venezuela. Thraulodes vitripennis (BLANCHARD) Ephemera (Cloe) vitripennis BLANCHARD in GAY, 1851: 107. Cloeon vitripennis; EATON, 1871: 107. Thraulus(?) vitripennis; EATON, 1887: 297. Thraulodes vitripennis; ULMER, 1920c: 116; ULMER, 1920a: 34; ULMER, 1921: 262 (male); LESTAGE, 1924c: 57; NEEDHAM & MURPHY, 1924: 41; LESTAGE, 1931b: 58; TRAVER & EDMUNDS, 1967: 351; HUBBARD, 1982a: 267.

Taxonomy: Thraulodes vitripennis was described by BLANCHARD (1851) in Ephemera and later transferred by ULMER (1921) to Thraulodes, because vein MP of the hind wing was forked. The abdomen is missing in the only known specimen. We agree with TRAVER & EDMUNDS (1967) about the dubious generic placement of this species in Thraulodes because there are several other Neotropical genera with forked vein MP of the hind wings, and Thraulodes does not occur in Chile. Distribution: Chile. Genus Thraulus EATON Thraulus EATON, 1881: 195. (Type-species: Thraulus bellus EATON, original designation).

Taxonomy: The two species from South America currently assigned to Thraulus obviously do not belong to this genus. It does not occur in South America. Thraulus siewertii (WEYENBERGH) Nomen Dubium Cloe siewertii WEYENBERGH, 1883: 170. Potamanthus siewertii; EATON, 1887: 296; HUBBARD, 1982a: 272. Thraulus siewertii; LESTAGE, 1924c: 57; LESTAGE, 1931a: 74.

Taxonomy: This species is almost certainly not a Thraulus, but its correct placement is uncertain. We consider it a nomen dubium. Distribution: Argentina. Thraulus vogleri (WEYENBERGH) Nomen Dubium Cloe(?) vogleri WEYENBERGH, 1883: 171. Potamanthus vogleri; EATON, 1887: 296 (male, female); HUBBARD, 1982a: 272. Thraulus vogleri; LESTAGE, 1924c: 57; LESTAGE, 1931a: 74.

Taxonomy: This species is almost certainly not a Thraulus, but its correct placement is uncertain. It must be considered a nomen dubium. Distribution: Argentina.

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Genus Tikuna SAVAGE, FLOWERS & PORRAS Tikuna SAVAGE, FLOWERS & PORRAS, 2005: 1. (Type-species: Choroterpes atramentum TRAVER, original designation).

Taxonomy: The genus Tikuna was established by SAVAGE et al. (2005) for the species C. atramentum TRAVER, from Costa Rica. These authors stated that it belonged to the Terpides lineage, which includes the genera Terpides and Fittkaulus. Recently, PETERS et al. (2005) transferred also Choroterpes (Choroterpes) bilineata to the newly established genus Tikuna. Adult Characteristics: 1) Upper portion of male eyes located on short, wide stalk (Fig. 192A); 2) vein ICu1 attached at base to CuA (Fig. 192B); 3) hind wings small, approximately 1/8 length of fore wings, costal projection thumb-like and rounded (Figs. 192C-D); 4) tarsal claws of a pair similar, with an apical hook and an opposing lobe (Fig. 192E); 5) genital forceps three-segmented (Fig. 192F); 6) penes short; basal 1/2 fused, apical 1/2 unfused, mid-lateral margins with projections, apices without appendages (Fig. 192F); 7) female sternum IX deeply and broadly cleft apically (Fig. 192G). Nymphal Characteristics: 1) Head hypognathous; 2) labrum (Fig. 192H) with anteromedian emargination well developed; 3) lingua of hypopharynx without lateral processes (Fig. 192I); 4) large apically curved unpectinate seta near large pectinate seta on inner anterior angle of maxillae (Fig. 192J); 5) glossae curved ventrally (Fig. 192K); 6) fore claws weakly hooked and narrow apically, with basal group of 5-6 denticles, increasing in size apically; and a subapical row of 5 narrow pointed denticles, increasing in size apically (Fig. 192L); 7) abdominal gill I lanceolate, dorsal and ventral portions same size (Fig. 192M); gills II-VI broader basally with inner and outer lobes on dorsal and ventral portions, ventral portion smaller than dorsal (Fig. 192N); gill VII similar, without lobes (Fig. 192O); and 8) caudal filaments with swimming setae (basal 2/5-3/5 of filaments with long setae on inner margin of cerci and lateral margins of terminal filament). Distribution: Central America and Northern South America. Biology: The nymphs were collected from small intermittent streams with moderate to dense shade, from areas of riffles to small pools. The adults were collected at light, and apparently have a long emergence period, as adults have been collected during most of the year. Tikuna bilineata (NEEDHAM & MURPHY) Choroterpes bilineata NEEDHAM & MURPHY, 1924: 48 (male, female); TRAVER, 1947a: 156 (male, female). Choroterpes (Choroterpes) bilineata; HUBBARD, 1982a: 264. Tikuna bilineata PETERS et al., 2005: 52.

Taxonomy: T. bilineata was described in the genus Choroterpes by NEEDHAM & MURPHY (1924) based on male subimagos and female imagos. After SAVAGE et al. (2005) established the new genus Tikuna for Choroterpes atramentum, it became clear that C. bilineata was closely related to this species, and for this reason PETERS et al. (2005) transferred it to Tikuna. T. bilineata can be separated from the other species of the genus by the following combination of characters: 1) Membrane of fore wings with costal strip of yellowish to light tan pigment;

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MP MA

B A C

CP

Sc

ICu1 CuA

E D

F

G H

I

J

K

D

L

M

N

O

Plate 192. Tikuna atramentum. Figs. 192A-G, imago: 192A, eyes, l. v.; 192B, fore wing; 192C, hind wing; 192D, hind wing, detail; 192E, tarsal claw, female imago; 192F, genitalia, v.v.; 192G, female sternum IX. Figs. 192H-O, nymph: 192H, labrum; 192I, hypopharynx, v.v.; 192J, maxilla; 192K, labium, left dorsal, right ventral; 192L, fore tarsal claw; 192M, gill I; 192N, gill V; 192O, gill VII. (CP = Costal Projection; D = denticle). Figs. 192A, E-O modified from SAVAGE et al., 2005.

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2) cross veins in apical costal area anastomosed; and 3) anterior portion of thoracic nota and abdominal terga 1-8 with prominent, narrow continuous submedian blackish stripes. Distribution and Biology: Peru, Surinam and Venezuela. Unpublished records from Ecuador, Brazil and Colombia. Genus Traverella EDMUNDS Traverella EDMUNDS, 1948: 141; EDMUNDS, 1950b: 551; TRAVER, 1960a: 24; TRAVER, 1960b: 73; THEW, 1960b: 122; TRAVER, 1963: 25; TSUI & PETERS, 1972: 331; ALLEN, 1973a: 1287; FLOWERS & DOMÍNGUEZ, 1991: 51; DOMÍNGUEZ, 1995b: 423; DOMÍNGUEZ & CUEZZO, 2002: 149. (Type-species: Thraulus albertanus MCDUNNOUGH, original designation).

Taxonomy: Traverella was established by EDMUNDS (1948) after rearing adults of Thraulus albertanus MCDUNNOUGH. In the same work he transferred Thraulus presidianus TRAVER to Traverella. In 1950, following the suggestion of ULMER, EDMUNDS added four more species to Traverella: Thraulus maculipennis ULMER (now in Hermanella), Thraulus ehrhardti ULMER (now in Needhamella), Thraulus primanus EATON (now in Hydrosmilodon), and Thraulus versicolor EATON. In 1960, TRAVER transferred Thraulus bradleyi NEEDHAM & MURPHY, Thraulus montium ULMER, and Thraulus valdemari ESBEN-PETERSEN to Traverella. In 1972 KILGORE & ALLEN described Traverella castanea and ALLEN added Traverella lewisi in 1973. In 1991, FLOWERS & DOMÍNGUEZ conducted a cladistic analysis of the Hermanella complex, and concluded that the Traverella group was monophyletic, and composed by Traverella s.s., Traverella “San Juan” (later Traverella (Zonda) ) and Traverella “bradleyi”. In 1995b, DOMÍNGUEZ established the new subgenus Traverella (Zonda), and described the new species T. (Zonda) calingastensis. LUGO-ORTIZ & MCCAFFERTY added T. longifrons, T. holzenthal and T. promifrons in 1996, and RANDOLPH & MCCAFFERTY (2000) transferred Thraulus sallei NAVÁS to Traverella. In 2002 DOMÍNGUEZ & CUEZZO made a cladistic analysis using egg chorion characters, with results that confirmed the proposed relationships between the Hermanella and Traverella members of the complex. In his revision of nymphs of this genus, ALLEN (1973a) used the following characters to distinguish the nymphs of the species: 1) Form, direction, and degree of development of the frontoclypeal projection; 2) color of the body and legs, and maculae on the legs and abdominal terga; 3) body size; 4) number of spines on the anterior face of fore and hind femora; and 5) geographic distribution. Based on large series of nymphs of T. (Z.) calingastensis, it was shown that the size and form of frontoclypeal projection change with successive molts (Fig. 195H), and therefore this character is valid only when comparing equivalent instars. We caution that, except in the last nymphal stage (in which the wing pads are black), the nymphs are very difficult to determine by traditional methods mainly due to the allometric growth of certain body parts. To the list of dubious characters above, we add the number of tarsal denticles. Another variable generic character is the presence of a spine on the inner anterior angle of the maxillae, which becomes visible only in the last nymphal stages. With respect to the adults, TRAVER (1963) constructed an artificial key to separate the South American species, and principally used the coloration of the wings and abdominal terga. Adult Characteristics: 1) Vein MA of fore wings asymmetrical, with slanting cross vein above MA fork (Figs. 193A, D); 2) vein MP of hind wings unforked (Figs. 193B-C, E); 3) vein Sc of hind wings ending in cross vein near costal projection (Figs. 193B-C, E),

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4) penes divided in apical half, with ventral subapical projection on each lobe (Figs. 193F, J, K); 5) styliger plate with conical projections apically near bases of forceps; 6) tarsal claws of a pair dissimilar, one hooked and the other blunt (Fig. 193H); 7) female sternum IX apically cleft (Fig. 193I). Nymphal Characteristics: 1) Labrum as wide as head, with rounded margins and long setae (Fig. 195A); 2) clypeus with a variable median projection (Fig. 195A); 3) maxillae with (Fig. 195G) or without (Fig. 195C) tusk on antero-median margin; setae on segment III of palpi arranged in rows; segment I of palpi cup shaped and joined laterally to maxilla; 4) outer margin of mandibles angular (Fig. 195B); 5) labium as in Fig. 195D; 6) abdominal gills on segments I-VI or I-VII, with fringed margins (Fig. 195F); 7) posterolateral spines on abdominal segments VIII-IX; and 8) denticles on tarsal claws subequal in size (Fig. 195E). Distribution: New World, from the Province of Saskatchewan, Canada (52ºN) to the Province of San Juan, Argentina (31ºS). The species of Traverella (Zonda) are found close to the Andes, while the species of Traverella (Traverella) are found in the lowlands. Biology: Nymphs of this genus, as well as others of the Hermanella complex, are apparently filter-feeders, utilizing the wide, setaceous labrum and maxillae to filter particles from the current. The nymphs of T. calingastensis were always found in areas of stronger current, with many specimens under the same rocks. The observed mating flight took place in the morning, close to the river, but over land at an altitude of approximately 4-5 m. The swarm was composed of around 15-20 males. KEY

SOUTH AMERICAN TRAVERELLA

TO

Imagos 1 – 2(1) – 3(2) –

CLAVE

Vein ICu1 of fore wings attached to vein CuA (as in Fig. 193D); male styliger plate with long, narrow paired projections ................................. Traverella (Traverella).......2 Vein ICu1 of fore wings free basally (Fig. 193A); male styliger plate with broad paired projections (Fig. 193F) ...................................... Traverella (Zonda) calingastensis Wing membranes almost hyaline; genitalia as in Fig. 193K ......... Traverella montium Wing membranes with coloration; genitalia similar to Fig. 193J ............................. 3 Wing membranes hyaline with prominent brown areas at base; no costal cross veins basal to bulla .................................................................................. Traverella bradleyi Wing membranes translucent brown; costal area and cross veins blackish brown; 2-5 costal cross veins basal to bulla ................................................. Traverella valdemari PARA LOS

TRAVERELLA

SUDAMERICANOS

Imagos 1 –

Vena ICu1 de las alas anteriores unida a la vena CuA (como en la Fig. 193D); placa estilígera del macho con proyecciones pares, largas y delgadas ................................. .................................................................................................... Traverella (Traverella)....... 2 Vena ICu1 de las alas anteriores libres basalmente (Fig. 193A); placa estilígera del macho con proyecciones pares anchas (Fig. 193F) ........................................... .............................................................................................. Traverella (Zonda) calingastensis

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Membrana de las alas casi hialinas, sin venas transversas costales basales a la bulla, genitalia como en la Fig. 193K ........................................................... Traverella montium Alas con coloración, genitalia similar a la de la Fig. 193J ......................................... 3 Membrana de las alas hialinas, con prominentes áreas castañas en la base, sin venas transversas costales basales a la bulla ...................................... Traverella bradleyi Membrana de las alas castaño-translúcidas, área costal y venas transversas castaño-negruzcas; 2-5 transversas costales basales a la bulla .... Traverella valdemari

3(2)

sCV

MA

ICu1

ICu1

A

Sc

D

CuA

B

E

CuA

CP

C

MP Pp

SPp

Pp

J

F

G

K H

I

Plate 193. Traverella, imagos. Figs. 193A-E, wings. 193A-C, T. (Z.) calingastensis. 193A, fore wing; 193B, hind wing; 193C, hind wing, detail. 193D-E, T. (T.) albertana: 193D, fore wing; 193E, hind wing. 193FI, T. (Z.) calingastensis: 193F, genitalia, v.v.; 193G, abdominal terga II-VII; 193H, tarsal claw; 193I, apex of female sternum IX. 193J, T. (T.) bradleyi, penes, v.v. 193K, T. (T.) montium, penes, v.v. (sCV = slanted cross vein; CP = Costal Projection; Pp = Penes projections; SPp = Styliger Plate projection). Figs. 193A-I modified from DOMÍNGUEZ, 1995b; 193J from EDMUNDS, 1948; 193K from ULMER, 1943.

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Subgenus Traverella (Traverella) EDMUNDS Traverella (Traverella); DOMÍNGUEZ, 1995b: 424; DOMÍNGUEZ, FERREIRA & NIETO, 2001: 316; DOMÍNGUEZ & CUEZZO, 2002: 149.

Taxonomy: Traverella (Traverella) can be separated from the other subgenera of Traverella, in the imago: 1) Vein ICu1 of fore wings attached to vein CuA (Fig. 193D); 2) male styliger plate with long and narrow paired projections. In the nymph: 1) Tusk present at inner anterior angle of maxillae (Fig. 195G); 2) segment III of labial palpi not elongated (0.8-1.2 times length of segment II); 3) gills present on abdominal segments I-VII. Distribution: Canada (52ºN) to Northwestern Argentina (27ºS). Traverella (Traverella) bradleyi (NEEDHAM & MURPHY) Thraulus bradleyi NEEDHAM & MURPHY, 1924: 45 (male); ULMER, 1943: 28. Traverella bradleyi; TRAVER, 1960b: 73; THEW, 1960b: 123; ALLEN, 1973a: 1287; HUBBARD, 1982a: 267; FLOWERS & DOMÍNGUEZ, 1991: 51; DOMÍNGUEZ, 1995b: 424; DOMÍNGUEZ, FERREIRA & NIETO, 2001: 316; DOMÍNGUEZ & CUEZZO, 2002: 149.

Taxonomy: The body of the type-specimen of Traverella bradleyi has been lost and only the wings and genitalia can be studied. The only description of this species is that of NEEDHAM & MURPHY (1924) and it is quite incomplete. Although HUBBARD (1982a) in his catalog of the South American Ephemeroptera listed Brazil as the type-locality for this species, the original description and the labels on the specimen list Argentina (Misiones). T. bradleyi can be differentiated from the other species of the subgenus by the following combination of characters in the imago: 1) Wings without maculae except for one basal macula; and 2) spine-like projections on penes long, slender, and curved apically toward median of the body (Fig. 193J). Distribution and Biology: Argentina (Misiones). The river at the type locality is large, about 15-20 m wide and in parts 3-4 m deep. The bottom is composed of stones and rubble. The surroundings are covered by forest. Traverella (Traverella) montium (ULMER) Thraulus montium ULMER, 1943: 32 (male). Traverella montium; TRAVER, 1960b: 73; ALLEN, 1973a: 1287; HUBBARD, 1982a: 268; FLOWERS & DOMÍNGUEZ, 1991: 51; DOMÍNGUEZ, 1995b: 424.

Taxonomy: The genitalia of Traverella montium is very characteristic and atypical for the genus, and therefore we have serious doubts about its correct placement. Until it is possible to study the type we prefer to leave it in Traverella. The imagos of T. montium can be distinguished from the other species of the subgenus by the following combination of characters: 1) Wing membranes almost colorless; 2) no costal cross veins basal to bulla; and 3) penes divided into three ventral lobes (two lateral sharp lobes and one tongue-like ventral lobe) with two short thick apical projections (Fig. 193K). Distribution: Peru.

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Plate 194. Traverella (Z.) calingastensis. Fig. 194A, nymphal habitus. Drawing by A. Dupuy.

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Traverella (Traverella) valdemari (ESBEN-PETERSEN) Thraulus valdemari ESBEN-PETERSEN, 1912: 335 (male); ULMER, 1920c: 117; ULMER, 1921: 263 (male); ULMER, 1943: 30 (male). Traverella valdemari; TRAVER, 1960b: 73; TRAVER, 1963: 25; ALLEN, 1973a: 1287; HUBBARD, 1982a: 268; FLOWERS & DOMÍNGUEZ, 1991: 51; DOMÍNGUEZ, 1995b: 424.

Taxonomy: The imagos of Traverella valdemari can be distinguished from the other species of the genus by the following combination of characters: 1) Wing membrane with brownish tinge and costal and subcostal areas blackish-brown; and 2) abdominal terga IIVIII yellowish or light brown, with blackish-brown markings joining at the posterior margin. Also with paler quadrangular maculae laterally on terga, the posterolateral angles shaded grayish-brown with a dark longitudinal band parallel to lateral border. Distribution and Biology: Argentina (Misiones, Bompland). The river from the type locality is large, about 15-20 m wide and in parts 3-4 m deep. The bottom is composed of stones and rubble. The surroundings are covered by rain forest.

B

A

C

FCp III

D II

E T

I

D G F

H

Plate 195. Traverella, nymphs. Figs. 195A-F, 195H, T. (Z.) calingastensis. 195A, clypeus and labrum, d.v.; 195B, left mandible; 195C, maxilla; 195D, labium, left dorsal, right ventral; 195E, tarsal claw; 195F, gill II. 195G, T. (T.) albertana, maxilla. 195H, differential development of frontoclypeal projection in different nymphal stages (D = denticle; T = tusk; FCp = frontoclypeal projection; Roman numbers = segment numbers). Figs. 195A-F, H modified from DOMÍNGUEZ, 1995b; 195G, from EDMUNDS, 1948.

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Subgenus Traverella (Zonda) DOMÍNGUEZ Traverella (Zonda); DOMÍNGUEZ, 1995b: 424; DOMÍNGUEZ, FERREIRA & NIETO, 2001: 316; DOMÍNGUEZ & CUEZZO, 2002: 149.

Taxonomy: Traverella (Zonda) can be separated from the other subgenera of Traverella by the following characters, in the imago: 1) Vein ICu1 of fore wings free basally (Fig. 193A); and 2) male styliger plate with broad paired projections (Fig. 193F). In the nymph (Fig. 194A): 1) Tusk at inner anterior angle of maxillae absent or produced as a very small prominence in last instar (Fig. 195C); 2) segment III of labial palpi elongated (more than 1.2 times length of segment II) (Fig. 195D); 3) gills present on abdominal segments I-VI (Fig. 194A). Distribution: Northern and Central Argentina (from 31ºS to 25ºS). Traverella (Zonda) calingastensis DOMÍNGUEZ Traverella (Zonda) calingastensis DOMÍNGUEZ, 1995b: 424; DOMÍNGUEZ, FERREIRA & NIETO, 2001: 316; DOMÍNGUEZ & CUEZZO, 2002: 149.

Taxonomy: This is the only species described in this subgenus, although apparently there are several others known only from nymphs, but not named. The imagos of T. calingastensis can be separated from the other species of the genus by the following combination of characters: 1) Veins C, Sc and R1 orange-brown, remaining veins paler, lighter posteriorly; 2) abdominal terga I-VI translucent, with a median brownish-gray band, darker on lateral margins (Fig. 193G); 3) forceps and penes whitish, spine-like projections yellowish (Fig. 193F). In the nymph: 1) Frontoclypeal projection as in Fig. 195A; 2) abdominal color pattern as in imago (Fig. 193G). Distribution and Biology: Central and Northern Argentina, along the lower part of the Andes. The nymphs of this species were collected in stony, mountain streams in areas of stronger current. Nymphs presented a patchy distribution, with many individuals under some rocks. Adults were collected in January. Nuptial flight took place from 8:30 to 11:00 very close to the rocky areas of the river, but over land. Genus Ulmeritoides TRAVER Ulmeritus (Ulmeritoides) TRAVER, 1959b: 8; THEW, 1960b: 125. Ulmeritus (Pseudulmeritus) TRAVER, 1959b: 8. Ulmeritoides, DOMÍNGUEZ, 1991: 160; FLOWERS & DOMÍNGUEZ, 1991: 52; DOMÍNGUEZ, 1995a: 16; ORTH et al., 2000: 33. (Type-species: Ulmeritus uruguayensis TRAVER, original designation).

Taxonomy: Ulmeritoides was established by TRAVER (1959b) as a subgenus of Ulmeritus and originally included two species: Ulmeritus (Ulmeritoides) uruguayensis and Ulmeritus (Ulmeritoides) leuteotinctus, the former known from a male and female imagos and the latter only from females. THEW (1960) described another species as Ulmeritus (Ulmeritoides) adustus. DOMÍNGUEZ (1991) reared for the first time the nymph of this group and elevated Ulmeritoides to generic status, synonymizing the subgenus Ulmeritus (Pseudulmeritus) with Ulmeritoides and including Ulmeritus flavopedes SPIETH in Ulmeritoides. As only two species from South America

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are known from nymphs, no key is provided. Two other species included in this genus were described from Costa Rica, U. guanacaste DOMÍNGUEZ and U. tifferae DOMÍNGUEZ and are not included here except for illustrative purposes. A key to species of Ulmeritoides is provided in DOMÍNGUEZ (1995). Specimens of undescribed species are also recorded from Guyana, French Guiana and Venezuela. Adult Characteristics: 1) Penes divided to base, apex of penis lobes rounded to almost straight and base of forceps very broad (Figs. 196E, H, K, N, Q, T); 2) vein Rs of fore wings forming two triangles (Fig. 196A); 3) only costal cross veins of fore wings with maculae (Fig. 196A); 4) vein MP of hind wings forked (Fig. 196B); 5) costal projection of hind wings rounded with apex of projection located 3/10 - 4/10 distance from base of wing; 6) vein Sc of hind wings ending less than 4/5 of distance from base of wing to apex; 6) female sternum IX broadly cleft apically (Fig. 196D); 7) tarsal claws dissimilar, one apically hooked and the other blunt (Fig. 196C). Nymphal Characteristics (Fig. 197A): 1) Labrum as broad as clypeus, with deep anteromedian emargination, lateral borders of emargination almost straight and median denticle a little larger (Figs. 198A-B) to much larger (Figs. 198C-D) than others; 2) setae present along entire outer margin of mandible, most dense in apical 1/3 (Fig. 198E); 3) segment III of maxillary palpi 2/3 length of segment II (Fig. 198F); 4) abdominal gills with finger-like margins (Fig. 198I); 5) dorsal row of thick setae from apex of segment II to segment III of labial palpi; 6) posterolateral projections present on abdominal segments VI-IX, very large on VIII-IX, and with lateral spines (Fig. 198J); and 7) denticles of tarsal claws subequal in size (Fig. 198K). Distribution: Argentina, Brazil, Colombia, Costa Rica, Surinam, and Uruguay. Additional records of nymphs of undescribed species: Paraguay, Guyana, French Guiana, Venezuela and Guatemala. Biology: The species of this genus are not common. Populations are relatively small and seem to be mostly associated with lowland subtropical or tropical streams. The nymphs (Fig. 197A) are characteristic, due to the dark, fringed gills that normally cover part of the abdomen and move like a “skirt”, and with mouthparts as in Figs. 198A-H. They have been collected from leaf packs and under logs from a variety of streams, from pool areas to slow to swift current sections. Most adults were collected at light traps. In some cases (e.g., U. spinulipenis) only subimagos were collected at light, from which it can be supposed that dusk is the time of emergence. In others (U. misionensis and U. luteotinctus), only female imagos were attracted by the light trap, many arriving with a single spherical mass of eggs adhered to the abdomen. In this case, the nuptial flight probably takes place during daylight. Based on collection data, adults emerge from November to May. KEY

TO

SOUTH AMERICAN ULMERITOIDES

Male Imagos 1 – 37

37

Fore wings with costal and subcostal areas tinged with brown (Fig. 196A) ........ 2 Fore wings with costal and subcostal areas hyaline ................................................... 4 U. luteotinctus and U. patagiatus are not included in this key. See species discussion

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C CP

A

D

Sc

B

J

MP

F

I

E

G

H

L

O

LG

K

M

N

R

Q

P

U

S

T

Plate196. Ulmeritoides, imagos. Figs. 196A-G: U. misionensis. 196A, fore wing (arrows showing triangles); 196B, hind wing; 196C, male tarsal claw; 196D, apex of female sternum IX; 196E, genitalia, v.v.; 196F, penes, detail; 196G, abdominal terga II-VII. 196H-J: U. haarupi. 196H, genitalia, v.v.; 196I, penes, detail; 196J, abdominal terga II-VII. 196K-M: U. flavopedes. 196K, genitalia, v.v.; 196L, penes, detail; 196M, abdominal terga II-VII. 196N-P: U. huitoto. 196N, genitalia, v.v.; 196O, penes, detail; 196P, abdominal terga II-VII. 196Q-S: U. spinulipenis. 196Q, genitalia, v.v.; 196R, penes, detail; 196S, abdominal terga II-VII. 196T-U: U. uruguayensis. 196T, genitalia, v.v.; 196U, penes, detail. (CP = Costal Projection; LG = lateral groove). Figs. 196A-U modified from DOMÍNGUEZ, 1995a.

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2(1) – 3(2) – 4(1) – 5 (4) –

CLAVE

Apex of penis lobes convex with a small spine on ventral surface of outer corner (Figs. 196H-I); abdominal color pattern as in Fig. 196J ............ Ulmeritoides haarupi Apex of penis lobes concave or straight, ending in a rather acute meso apical projection (Figs. 196E, N); abdominal color pattern not as above (Figs. 196G, P) ................................................................................................................................................ 3 Basal broad region of forceps almost 1/4 length of segment I, inner angle rounded (Fig. 196N); abdominal color pattern as in Fig. 196P . Ulmeritoides huitoto Basal broad region of forceps about 1/6 length of segment I, inner angle acute (Fig. 196E); abdominal color pattern as in Fig. 196G ........... Ulmeritoides misionensis Apex of penis lobes rounded, each penis lobe with a lateral groove (Figs. 196K-L) .............................................................................................................. Ulmeritoides flavopedes Apex of penis lobes not rounded (Fig. 196Q, T); no lateral grooves present ..... 5 Penis lobes with apical margin ending in an acute projection (Figs. 196T-U) ........ ..........................................................................................................Ulmeritoides uruguayensis Penes lobes with several small apical spines (Figs. 196Q-R) ...................................... ........................................................................................................... Ulmeritoides spinulipenis PARA LOS

Imagos Macho 1 – 2(1) – 3(2) – 4(1) – 5(4) –

38

ULMERITOIDES

SUDAMERICANOS

38

Alas anteriores con las áreas costal y subcostal teñida de castaño (Fig. 196A) ....... 2 Alas anteriores con las áreas costal y subcostal hialina .............................................. 4 Ápice de los lóbulos de los penes más o menos redondeados, con una pequeña espina sobre la superficie ventral del ángulo externo (Figs. 196H-I); patrón de coloración abdominal como en la Fig. 196J .................................. Ulmeritoides haarupi Ápice de los lóbulos de los penes rectos, terminando en una proyección meso apical más bien aguda (Figs. 196E, N); patrón de coloración abdominal no como en el caso anterior (Figs. 196G-P) .................................................................................. 3 Región basal ensanchada de los fórceps casi 1/4 de la longitud del segmento I, ángulo interno redondeado (Fig. 196N); patrón de coloración abdominal como en la Fig. 196P ............................................................................................ Ulmeritoides huitoto Región basal ensanchada de los fórceps casi 1/6 de la longitud del segmento I, ángulo interno agudo (Fig. 196E); patrón de coloración abdominal como en la Fig. 196G ................................................................................................. Ulmeritoides misionensis Apice de los lóbulos de los penes redondeados, cada lóbulo de los penes con un surco lateral (Figs. 196K-L) ........................................................... Ulmeritoides flavopedes Apice de los lóbulos de los penes no redondeado (Figs. 196Q, T); lóbulos sin surcos laterales .................................................................................................................... 5 Lóbulos de los penes con el margen apical terminando en una proyección aguda (Figs. 196T-U) ...............................................................................Ulmeritoides uruguayensis Lóbulos de los penes con varias espinas pequeñas sobre el margen apical (Figs. 196Q-R) ........................................................................................... Ulmeritoides spinulipenis

U. luteotinctus y U. patagiatus no se incluyen en la clave. Ver discusión de las especies.

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PP

Plate 197. Ulmeritoides misionensis. Fig. 197A, nymphal habitus. (PP = posterolateral projections). Drawing by C. Molineri.

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Ulmeritoides flavopedes (SPIETH) Thraulodes flavopedes SPIETH, 1943: 11. Atalophlebioides flavopedes; TRAVER, 1946: 426. Ulmeritus (Pseudulmeritus) flavopedes; TRAVER, 1959b: 8. Ulmeritoides flavopedes; DOMÍNGUEZ, 1991: 162; DOMÍNGUEZ, 1995a: 19.

Taxonomy: This unusual species was described by SPIETH (1943) in the genus Thraulodes. TRAVER (1959) later established the subgenus Ulmeritus (Pseudulmeritus) for this single species. When DOMÍNGUEZ (1991) elevated Ulmeritoides to generic status, he tentatively included the species, a placement later confirmed by phylogenetic analysis of the Ulmeritus-Ulmeritoides complex (DOMÍNGUEZ, 1995). This species, known only from male imagos, can be differentiated from other species of the genus by the following combination of characters: 1) Membrane of fore wings hyaline, wing bases brown; 2) apex of penis lobes rounded, each with a lateral groove (Figs. 196K-L); and 3) abdominal terga orange-brown, posterior margins blackish, color pattern as in Fig. 196M. Distribution: Surinam. Ulmeritoides haarupi (ESBEN-PETERSEN) Thraulus haarupi ESBEN-PETERSEN, 1912: 337 (male); NAVÁS, 1917: 188 (male, female). Deleatidium haarupi; ULMER, 1920c: 115; ULMER, 1938: 105; ULMER, 1943: 15. Atalophlebioides haarupi; TRAVER, 1946: 423. Ulmeritus (Ulmeritus) haarupi; TRAVER, 1959b: 5 (female); THEW, 1960b: 123; HUBBARD, 1982A: 268; DOMÍNGUEZ, 1991: 160; HUBBARD, DOMÍNGUEZ & PESCADOR: 1992: 222. Ulmeritus haarupi; DOMÍNGUEZ, 1991: 160; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 82 Deleatidium vittatum THEW, 1960b: 130. Ulmeritoides fidalgoi DOMÍNGUEZ, 1995a: 23. Ulmeritoides haarupi PETERS et al., 2005: 53.

Taxonomy: ESBEN-PETERSEN (1912) described this species in the genus Thraulus from a badly preserved female from Bompland, Argentina, with fore legs and cerci missing, and TRAVER (1959) later described another female from Uruguay. Then THEW (1960b) described Deleatidium vittatum from Uruguay based on two more females, but without illustrations. DOMÍNGUEZ (1995) described Ulmeritoides fidalgoi based upon a good series of males and females from Bompland. Recently, PETERS et al. (2005) examined the type material of Ulmeritus haarupi and Deleatidium vittatum, discovering both fit within the variation in the female series of Ulmeritoides fidalgoi; thus D. vittatum and U. fidalgoi were synonymized with Ulmeritoides haarupi. Ulmeritoides haarupi can be differentiated from the other species of the genus by the following combination of characters: 1) Fore wings hyaline, with costal and subcostal areas tinged with brown and cross veins in this area a heavier brown (similar to Fig. 196A); 2) penis lobes more or less rounded, each with a small spine close to the outer corner (Figs. 196H-I); 3) abdominal terga brown-orange, with pattern as in Fig. 196J, although medial markings of terga inconspicuous on females.

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525

III II

B

I

MD

C

E

F

D

G MD

H

I

PP

K J

PP

Plate 198. Ulmeritoides, nymphs. Figs. 198A-B: U. tifferae (species from Costa Rica, used to show variation of emargination): 198A, labrum d.v.; 198B, detail anteromedian emargination; 198C-K: U. misionensis. 198C, labrum, d.v.; 198D, detail anteromedian emargination; 198E, left mandible; 198F, maxilla; 198G, hypopharynx, v.v.; 198H, labium (left dorsal, right ventral); 198I, gill II; 198J, posterolateral projections on abdominal segments VIII-IX; 198K, tarsal claw. (MD = Median Denticle; PP = posterolateral projections; Roman numbers = segment numbers). Figs. 198A-I modified from DOMÍNGUEZ, 1991.

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Distribution and Biology: Northeastern Argentina, Brazil and Uruguay. The adults were collected between November and February. They were always found near streams and rivers with stony bottoms, where the nymphs probably inhabit. Ulmeritoides huitoto DOMÍNGUEZ & ZUÑIGA Ulmeritoides huitoto DOMÍNGUEZ & ZUÑIGA, 2003: 123.

Taxonomy: This species was described from male imagos from the Colombian Amazon region. It can be separated from the other species of the genus by the following combination of characters: 1) Fore wings hyaline, with costal and subcostal areas brown; 2) apex of penis lobes straight; basal broad region of forceps almost 1/4 length of segment (Figs. 196N-O); 3) abdominal terga yellowish-brown, with pattern as in Fig. 196P. This species seems to be related to U. misionensis, U. tifferae and U. haarupi but can be separated from them by the structure of the genitalia. Distribution and Biology: Colombia (Amazonas). The adults were collected with a light trap, in the Amazonian rainy season, near a black water stream. Ulmeritoides luteotinctus (TRAVER) Ulmeritus (Ulmeritoides) luteotinctus TRAVER, 1959b: 11 (male, female); HUBBARD, 1982a: 268. Ulmeritus luteotinctus; THEW, 1960b: 123. Ulmeritoides luteotinctus; DOMÍNGUEZ, 1995a: 18.

Taxonomy: TRAVER (1959) described Ulmeritoides luteotinctus from female imagos only, and also gave a description of three subimagos (two males and one female), which she believed might belong to this species. After studying the male subimagos mentioned by TRAVER in her original description, we believe that they belong to another species and for this reason we did not include them in the key. Ulmeritoides luteotinctus can be differentiated from the other species of the genus by the following combination of characters in the imago: 1) Fore wings hyaline, veins orangebrown; and 2) abdominal terga and sterna yellowish, slightly tinged with brownish, a darker band on posterior border of each segment. Distribution and Biology: Uruguay and northeastern Argentina. According to the dates of collection, the adults emerge between December and March, near zones of lagoons or open rapids. Only females have been collected at light, many arriving with a single spherical mass of eggs attached to the abdomen. Ulmeritoides misionensis DOMÍNGUEZ Ulmeritoides misionensis DOMÍNGUEZ, 1995a: 25.

Taxonomy: Ulmeritoides misionensis was described from imagos and nymphs of both sexes. This species can be differentiated from others of the genus by the following combination of characters. In the imago: 1) Fore wings hyaline, with costal and subcostal areas light brown, stigmatic area lighter (Fig. 196A); 2) apex of penis lobes straight, ending in an acute apical projection (Figs. 196E-F); 3) abdominal terga yellow-brown, with

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pattern as in Fig. 196G. In the nymph: 1) Median denticle on anteromedian emargination of labrum much larger than others (Figs. 198C-D); 2) fore tibiae with subbasal and subapical black bands; 3) femora of middle leg with median black spot. Distribution and Biology: Northeastern Argentina and Brazil. The adults were collected at light in November. The nymphs were collected in a small, stony stream, under rocks and logs. Ulmeritoides oepa LOPES, DA-SILVA & PY-DANIEL Ulmeritoides oepa LOPES, DA-SILVA & PY-DANIEL, 2003: 195.

Taxonomy: This species was described based only on nymphs. It can be differentiated from the other species of the genus by the following combination of characters: 1) Median denticle on anteromedian emargination of labrum much larger than other denticles; 2) fore tibiae lightly stained with black apically; 3) femora of middle leg without median black markings, but with apical brown marks. Distribution and Biology: Brazil (Roraima). The nymphs were collected from leaves and on stones in riffle areas. Ulmeritoides patagiatus (THEW) Ulmeritus patagiatus THEW, 1960b: 128 (male, female); HUBBARD, 1982a: 268. Ulmeritoides patagiatus; DOMÍNGUEZ, 1995a: 2.

Taxonomy: In the original description of Ulmeritus patagiatus, THEW (1960) did not place this species in any of the existing subgenera of Ulmeritus. We had the opportunity to study the type material, composed of subimagos of both sexes, which undoubtedly belong to Ulmeritoides. Nevertheless, specimens at this stage are not adequate for specific determination, and for this reason the species is not included in the key. When imagos of this species became available will be possible to verify if it is a valid species or should be synonymized. Distribution: Brazil (Santa Catarina). Ulmeritoides spinulipenis DOMÍNGUEZ Ulmeritoides spinulipenis DOMÍNGUEZ, 1995a: 22.

Taxonomy: Ulmeritoides spinulipenis was described from male imagos. They can be differentiated from the other species of the genus by the following combination of characters: 1) Fore wings hyaline, with three small spots near the bulla; 2) penis lobes with small spines, as in Figs. 196Q-R; and 3) abdominal terga with black markings enclosing a lighter central area as in Fig. 196S. This species is very similar in coloration to Ulmeritoides uruguayensis, but can be separated from it based on the structure of the penes. Distribution and Biology: Uruguay and Northeastern Argentina. Only subimagos have been collected at light, from which it can be supposed that dusk is the time of emergence. The adults have been collected between November and January.

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Ulmeritoides uruguayensis (TRAVER) Ulmeritus (Ulmeritoides) uruguayensis TRAVER, 1959b: 8 (male, female); HUBBARD, 1982a: 268. Ulmeritus uruguayensis; THEW, 1960b: 123. Ulmeritoides uruguayensis; DOMÍNGUEZ, 1991: 162; DOMÍNGUEZ, 1995a: 21. Ulmeritus adustus THEW, 1960b: 126 (male, female).

Taxonomy: TRAVER (1959), described this species in Ulmeritus, using it to establish the subgenus Ulmeritoides. THEW (1960), in the original description of Ulmeritus adustus, recognized that it was closely related to U. uruguayensis, but gave as diagnostic characters two small spines present on the lobes of the penes of U. adustus, and minor differences in coloration of the legs and abdomen. After studying the types DOMÍNGUEZ (1995) concluded that there were no differences in the form of the penis lobes, and that the difference in coloration of the holotype of Ulmeritus adustus fell within the intraspecific variation of Ulmeritoides uruguayensis. For these reasons, he synonymized Ulmeritus (Ulmeritoides) adustus with Ulmeritoides uruguayensis. Ulmeritoides uruguayensis can be differentiated from the other species of the genus by the following combination of characters. In the imagos: 1) Fore wings hyaline, with brown spot at base; 2) penis lobes ending in an acute apical projection (Fig. 196T), and without small spines on margin; 3) abdominal color pattern similar to that of U. spinulipenis (as in Fig. 196S). Distribution and Biology: Southeastern Brazil and Uruguay. According to collection dates, the adults emerge between December and April. Genus Ulmeritus TRAVER Ulmeritus TRAVER, 1956a: 2; TRAVER, 1959b: 4; THEW, 1960b: 123; DOMÍNGUEZ, 1991: 158; FLOWERS & DOMÍNGUEZ, 1991: 52; DOMÍNGUEZ, 1995a: 33. (Type-species: Ulmeritus carbonelli TRAVER, original designation).

Taxonomy: TRAVER (1956) established the genus Ulmeritus, designated Ulmeritus carbonelli as type species and transferred Atalophlebioides saopaulense TRAVER to this genus. In 1959, she established three subgenera in Ulmeritus: Ulmeritus (s.s), U. (Ulmeritoides) and U. (Pseudulmeritus). DOMÍNGUEZ (1991) elevated Ulmeritoides to generic status and synonymized Pseudulmeritus with Ulmeritoides. DOMÍNGUEZ (1995) studied the phylogeny of the Ulmeritus-Ulmeritoides group. Adult Characteristics: 1) Penes divided to base, apex of each lobe rounded with a ventral finger-like projection directed toward base of penes (Figs. 199C, G) and basal segment of forceps long, thin, except very broad at base; 2) vein Rs of fore wings forming two triangles (Figs. 199A, E); 3) fore wings with maculae around most cross veins (Figs. 199A, E); 4) vein MP of hind wings forked (Figs. 199B, F); 5) costal projection of hind wings rounded with apex located less than 1/3 distance from base; 6) vein Sc of hind wings ending near apex of wing; 7) female sternum IX broadly cleft apically (Fig. 199H); and 7) tarsal claws dissimilar, one apically hooked, other blunt (Fig. 199I). The abdominal color pattern of the species of this genus is represented here by U. balteatus (Fig. 199D). Nymphal Characteristics (general aspect similar to Ulmeritoides, Fig. 197A): 1) Labrum as broad as clypeus, with deep anteromedian emargination, lateral borders of emargination almost straight and without median denticle (Figs. 200A-B); 2) outer margin of mandibles with a dense row of setae on the apical 1/3, and weak tuft of long setae medially (Fig. 200C); 3) segment III of maxillary palpi subequal in length to segment II

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(Fig. 200D); 4) hypopharynx as in Fig. 200E; 5) dorsal row of thick setae from apex of segment II to segment III of labial palpi; 6) abdominal gills with finger-like margins (similar to Fig. 198I); 7) posterolateral spines present on abdominal segments VI-IX (Fig. 200G); and 8) denticles of tarsal claws subequal in size (Fig. 200H). Distribution: The species of this genus seems to be restricted to lowland subtropical streams, mainly in Southern Brazil, Uruguay and Northeastern Argentina. Biology: The nymphs were collected under rocks, in zones of rapids. Most adults were collected with light traps. Based on collection data, adults fly from September to March. KEYS

TO

ULMERITUS

Male Imagos 1 – 2(1) –

Cross veins of hind wings few (around 20); maculae only around subcostal cross veins .................................................................................................... Ulmeritus saopaulensis Cross veins of the hind wings numerous (33 to 50); maculae around all cross veins (Figs. 199B, F) ..................................................................................................................... 2 Maculae of fore wings forming bands (Fig. 199A); ventral prolongation of penes short, approximately as long as wide (Fig. 199C) .......................... Ulmeritus balteatus Maculae of fore wings not forming bands (Fig. 199E); ventral prolongation of penes long, approximately twice as long as wide (Fig. 199G) ....... Ulmeritus carbonelli

Nymphs 1 – 2 1) –

Segment III of maxillary palpi shorter than segment II ........ Ulmeritus saopaulensis Segment II and III of maxillary palpi subequal in length (Fig. 200D) .................. 2 Abdominal gills grayish brown ........................................................... Ulmeritus balteatus Abdominal gills grayish ....................................................................... Ulmeritus carbonelli

CLAVES

PARA

ULMERITUS

Imagos Machos 1 – 2(1) –

En el ala posterior venas transversas poco numerosas (alrededor de 20); manchas sólo alrededor de las venas transversas subcostales ................. Ulmeritus saopaulensis En el ala posterior venas transversas numerosas (entre 33 y 50); manchas alrededor de todas las venas transversas (Figs. 199B, F) ................................................................ 2 Manchas del ala I que forman bandas (Fig. 199A); prolongación ventral de los penes cortas, aproximadamente tan largas como anchas (Fig. 199C) ......... Ulmeritus balteatus Manchas del ala I sin formar bandas (Fig. 199E); prolongación ventral de los penes larga, aproximadamente 2 veces más larga que ancha (Fig. 199G) ............. Ulmeritus carbonelli

Ninfas 1 – 2(1) –

Segmento III del palpo maxilar más mas corto que el segmento II ......................... .............................................................................................................. Ulmeritus saopaulensis Segmentos II y III del palpo maxilar subiguales en longitud (Fig. 200D) ............. 2 Branquias castaño-grisáceas ................................................................ Ulmeritus balteatus Branquias grisáceas .............................................................................. Ulmeritus carbonelli

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D A

CP

Sc

B MP

C G

CP

E I

F

MP

Sc

H

Plate 199. Ulmeritus, imagos. Figs. 199A-D: U. balteatus. 199A, fore wing (arrows showing triangles); 199B, hind wing; 199C, genitalia, v.v.; 199D, abdominal terga II-VII. 199E-I: U. carbonelli. 199E, fore wing; 199F, hind wing; 199G, genitalia, v.v.; 199H, apex of female sternum IX; 199I, male tarsal claw. (CP = Costal Projection). Figs. 199A-F modified from DOMÍNGUEZ, 1991.

Ulmeritus balteatus THEW Ulmeritus sp. TRAVER, 1956a: 11 (male, female). Ulmeritus balteatus THEW, 1960b: 124 (male, female); DOMÍNGUEZ, 1991: 159; DOMÍNGUEZ, 1995a: 34. Ulmeritus (Ulmeritus) balteatus; HUBBARD, 1982a: 268.

Taxonomy: The illustrations were drawn based on specimens from the localities mentioned by TRAVER (1956) in her description and which correspond to the description of this species. Further discussion of this species can be found in the account of U. carbonelli.

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Ulmeritus balteatus can be separated from the other species of the genus by the following combination of characters: 1) Ventral prolongation of penis lobes short, approximately as long as wide (Fig. 199C); 2) cross veins of hind wings numerous (around 50) and most shaded with maculae (Fig. 199B); and 3) bands formed in fore wings by aggregation of maculae of cross veins (Fig. 199A). In the nymph: 1) Abdominal gills grayish; 2) segments II and III of maxillary palpi subequal in length (Fig. 200D) (see discussion under U. saopaulensis). Distribution: Uruguay and Brazil. Ulmeritus carbonelli TRAVER Ulmeritus carbonelli TRAVER, 1956a: 5 (male, female, nymph); THEW, 1960B: 123; DOMÍNGUEZ, 1991: 160; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 82; DOMÍNGUEZ, 1995a: 34. Ulmeritus (Ulmeritus) carbonelli; TRAVER, 1959b: 6 (male, female); HUBBARD, 1982a: 268; HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 222.

Taxonomy: When TRAVER (1956) described U. carbonelli, she treated two other male subimagos and four females as Ulmeritus sp. until she knew more of the variation of this species. THEW (1960) described U. balteatus based on three male and two female imagos and many subimagos from Nova Teutonia, Brazil, and included the specimens of Ulmeritus sp. of TRAVER from Lavalleja (Uruguay). THEW argued that the localities were 450 miles apart, there must be two species. Moreover, the male imagos showed “definite differences”. We have studied a male imago, female imagos, and, for the first time, nymphs of U. balteatus. The difference to which THEW alluded is the difference in the length of the ventral projection of the penes and the penis width. While this characteristic is repeated in the specimen from Uruguay, we have noticed variations in the wing maculation. Furthermore, the only significant difference found between the nymphs is in the coloration, especially of the abdominal gills. Ulmeritus carbonelli can be separated from the other species of the genus by the following combination of characters. In the imago: 1) Ventral projection of penes long, approximately twice as long as wide (Fig. 199G); 2) cross veins of hind wings numerous (around 35) and almost all shaded (Fig. 199F); and 3) maculae which surround cross veins do not group to form bands (Fig. 199E). In the nymph: 1) Abdominal gills grayish brown; 2) segments II and III of maxillary palpi subequal in length (Fig. 200D) (see discussion under U. saopaulensis). Distribution and Biology: Uruguay. The nymphs were collected under rocks, in streams and rivers of different sizes and current speed. The adults were collected in February and March. Ulmeritus saopaulensis (TRAVER) Atalophlebioides sao-paulense TRAVER, 1946: 424 (male, female). Ulmeritus sao-paulensis; TRAVER, 1956a: 12 (male); DOMÍNGUEZ, 1991: 160; DOMÍNGUEZ, 1995a: 34. Ulmeritus sao-paulense; THEW, 1960b: 23. Ulmeritus (Ulmeritus) sao-paulense; TRAVER, 1959b: 6; HUBBARD, 1982a: 268; DA-SILVA, 1992a: 855 (nymph).

Taxonomy: TRAVER (1946) originally described this species in Atalophlebioides, but in 1956 she established the genus Ulmeritus and included this species. In 1992, DA-SILVA reared and described the nymph. Nevertheless, it is important to note that one of the

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characters described in this paper (segment III of maxillary palpi smaller than segment II) is characteristic of the genus Ulmeritoides and must be used with caution. Ulmeritus saopaulensis can be differentiated from the other species of the genus by the following combination of characters. In the imago: 1) Ventral projections of penes shorter and more lateral than those of U. carbonelli; 2) few cross veins in hind wings (about 20) and cross veins shaded only in costal area; and 3) cross veins of fore wings less broadly shaded than in other species and not forming bands. In the nymph: 1) Abdominal gills dark gray; 2) segment III of maxillary palpi shorter than segment II. Distribution: Brazil. A

B

C

MD

III II

E I

D

H

PP

F

PP

G

Plate 200. Ulmeritus, nymphs. Figs. 200A-H: U. carbonelli. 200A, labrum d.v.; 200B, detail anteromedian emargination; 200C, left mandible; 200D, maxilla; 200E, hypopharynx; 200F, labium (left dorsal, right ventral); 200G, posterolateral projections on abdominal segments VIII-IX; 200H, tarsal claw. (MD = Median denticle; PP = posterolateral projections; Roman numbers = segment numbers). Figs. 200A-F modified from DOMÍNGUEZ, 1991.

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FAMILY MELANEMERELLIDAE Taxonomy: The classification of this monotypic family has been quite confusing. The genus Melanemerella was placed by ULMER (1920a) in the Ephemerellidae because of the presence of short intercalaries not joining the posterior margin, and the basal direction of the 3 anal and cubital veins of the fore wings. Since then, the taxonomic position of the genus has remained uncertain and much debate has been generated because it was known only from a female imago. LESTAGE (1925) commented on the wing shape and venation of Melanemerella, concluding that it was a “specialized Ephemerellidae”. DEMOULIN (1955g) transferred Melanemerella to Tricorythidae and established for it the subfamily Melanemerellinae. EDMUNDS et al. (1963) returned Melanemerella to Ephemerellidae, but retained the subfamily proposed by DEMOULIN. ALLEN (1965) remarked that the relationship of Melanemerella would remain tentative until the male imago and nymphal stages were discovered, and stated “The characters of the wings of M. brasiliana suggest that it should be maintained as a separate subfamily in the Ephemerellidae”. WANG & MCCAFFERTY (1996) decided to move Melanemerella to Leptophlebiidae. Recently MOLINERI & DOMÍNGUEZ (2003) described the nymphs and eggs of this genus, showed it to be an ephemerelloid mayfly, and proposed family status for this taxon. Genus Melanemerella ULMER Melanemerella ULMER, 1920a: 43; LESTAGE 1925: 258; DEMOULIN 1955g: 214; ALLEN 1965: 265; WANG & MCCAFFERTY 1996: 99; MOLINERI & DOMÍNGUEZ 2003: 267. (Type-species: Melanemerella brasiliana ULMER, original designation).

Taxonomy: Melanemerella was described by ULMER (1920) from a female, recently the nymphs and eggs were discovered (MOLINERI & DOMÍNGUEZ, 2003). Until the male of this genus is known, its taxonomic placement and relationships will remain somewhat enigmatic. The egg has a unique polar cap ringed by flap-like attachment structures. Adult Characteristics: 1) Fore wings (Fig. 202A) broad, with 2-3 marginal intercalary veins between longitudinal veins along length of posterior margin; 2) hind wings (Figs. 202B-C) ovoid, with marginal intercalaries; 3) costal projection of hind wings blunt, located in basal 2/3 of wing; and 4) cross veins numerous in both wings. Nymphal Characteristics: 1) Lateral margins of submentum rounded (Fig. 202F); 2) labial palpi 3-segmented, distal segment reduced (Fig. 202F); 3) labrum with shallow anteromedian emargination (Fig. 202D); 4) maxilla with tiny palp (Fig. 202E); 5) mandibles long and narrow (Figs. 202G-H); 6) femora of all legs wide and with dorsal transverse row of spines near base (Figs. 202I-K); 7) maximum width of fore femora equaling maximum length, acute median projection on leading edge (Fig. 202K); 8) middle coxae with a dorsal acute projection (Fig. 202J); 9) gills on abdominal segments II-VI, ventral lamellae of gills II-V bifid and fringed with numerous small lobes (Figs. 202N-Q); 10) abdominal gills II operculate (Fig. 201A); 11) paired posteromedial projections on abdominal terga (Fig. 201A). Distribution: Brazil.

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Melanemerella brasiliana ULMER Melanemerella brasiliana ULMER, 1920a: 44 (female); LESTAGE 1925: 301; DEMOULIN, 1955g: 214; HUBBARD, 1982a: 262; WANG & MCCAFFERTY 1996: 100. MOLINERI & DOMÍNGUEZ 2003: 271 (nymph and egg).

Taxonomy: The only known female of Melanemerella brasiliana has the following characteristics: 1) Body black; 2) wings transparent, blackish with violet reflections, except stigmatic area colorless; 3) fore legs blackish, with yellowish tarsi; 4) hind legs with black femora, tibiae and tarsi light yellowish except articulations darker. Nymphs: 1) Brownishgray to yellowish-brown, with whitish and brownish marks; 2) wing pads blackish. Distribution: Brazil (Espirito Santo and São Paulo).

A

Plate 201. Melanemerella brasiliana. Fig. 201A, nymphal habitus. Drawing by C. Molineri.

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A C D

B

E

G

H

F O

M

I

P

N

J

K

Q

L R

Plate 202. Melanemerella brasiliana. Figs. 202A-C, female imago: 202A-C. 202A, fore wing; 202B, female hind wing; 202C, idem, detail. Figs. 202D-R, nymph: 202D, labrum, d.v.; 202E, right maxilla, v.v. and detail of palpi; 202F, labium, v.v. at left, d.v. at right; 202G, right mandible, d.v.; 202H, left mandible, d.v.; 202I, hind leg; 202J, middle leg; 202K, fore leg; 202L, fore tarsal claw, detail; 202M, gill II, d.v.; 202N-R, gills IIVI, v.v. Figs. 202A-C modified from ULMER, 1920a; 202D-R from MOLINERI & DOMÍNGUEZ, 2003.

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FAMILY NESAMELETIDAE Taxonomy: This amphinotic family is represented by three genera: Metamonius from southern South America, Ameletoides from Australia, and Nesameletus from New Zealand. Its present definition is based on characters of the adults and nymphs discussed by KLUGE et al. (1995) and KLUGE (2004). Adult characteristics: 1) Fore wings slender and triangular with numerous veins (Fig. 203A); 2) hind wings large (Fig. 203B); 3) vein CuA of fore wings connected to hind margin by numerous veinlets (arrows in Fig. 203A); 4) tarsal claws of a pair similar, both acute and hooked. Nymphal characteristics: 1) Body fusiform (Fig. 203D); 2) epicranial suture ending on or before lateral ocelli (Fig. 3S); 3) mandibles elongated perpendicular to axis of articulation with blade-like incisors (Figs. 203F-G); 4) maxilla as in Fig. 203E; 5) glossae and paraglossae short and broad; 6) antennae short (Fig. 203D); 7) tarsal claws with two longitudinal rows of small denticles; 8) abdominal gills present on segments I-VII; 9) three caudal filaments present; and 10) posterolateral corners of abdominal segments expanded laterally. Distribution: South America (Argentina and Chile), Australia, New Zealand. Metamonius EATON Metamonius EATON, 1883-1888: 208; DEMOULIN, 1955c: 1; DEMOULIN, 1955d: 2; DOMÍNGUEZ et al., 1994: 33; MERCADO & ELLIOTT, 2004: 149. (Type-species: Siphlurus anceps EATON, original designation).

Taxonomy: MERCADO & ELLIOTT (2004) recently clarified the definition and composition of Metamonius. After extensive collections and based on reared material they concluded that this genus is represented by only one species, M. anceps (EATON). Adult characteristics: 1) Cross veins at stigmatic area moderately anastomosed (Fig. 203A); 2) costal projection of hind wings rounded (Fig. 203B); 3) forceps 4-segmented (Fig. 203C); 4) terminal filament reduced; 5) tarsal claws identical in all legs. Nymphal characteristics: 1) Body streamlined; 2) head hypognathous; 3) abdominal gills oval, with a median chitinized ridge and small denticles along the outer margin (Fig. 203H). Distribution: Argentina and Chile. Biology: MERCADO & ELLIOTT (2004) reported that this species is almost exclusively restricted to small headwater streams covered with nord-patagonic rain forest (dominated by Nothofagus trees), and not perturbed by human activities. Nymphs are very good swimmers and commonly found below rocks, leaf packs or other submerged objects. They have a general behavior similar to Baetidae. The adults emerge in spring from November to January, subimaginal stage last 2-4 days. Metamonius anceps (EATON) Siphlurus? anceps EATON, 1883 (male). Metamonius anceps; EATON, 1885: 209 (male, female); ULMER, 1904: 4; ULMER, 1920c: 134; LESTAGE, 1924c: 58; LESTAGE, 1930b: 438; LESTAGE, 1931b: 49; LESTAGE, 1935a: 95; NAVÁS, 1936a: 140; DEMOULIN, 1955b: 1 (female); DEMOULIN, 1955c: 2 (nymph); DEMOULIN, 1955d: 2 (female); KIMMINS, 1960: 270 (nymph); EDMUNDS et al., 1963: 28 (nymph); HUBBARD, 1982a: 273; MERCADO & ELLIOTT, 2004: 150.

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Nymphe des Baëtis-Typus ULMER, 1904: 6 (nymph). Ameletus? fuegiensis LESTAGE, 1935a: 110. Ameletoides fuegiensis; LESTAGE, 1935b: 348; ULMER, 1938: 107 (nymph). Metamonius? fuegiensis; DEMOULIN, 1955d: 5. Metamonius fuegiensis; DEMOULIN, 1955c: 3 (nymph); HUBBARD, 1982a: 273; HUBBARD et al., 1992: 209; DOMÍNGUEZ et al., 1994: 33 (adult, nymph); MERCADO & ELLIOTT, 2004: 151.

Taxonomy: Metamonius anceps is the only species of the genus, and can be recognized by the characteristics listed in the generic section. Distribution and Biology: See generic section.

A

C B

D

E

F

G

H

Plate 203. Metamonius anceps. Figs. 203A-C, male imago: 203A, fore wing; 203B, hind wing; 203C, genitalia. (arrows indicate sinuous veinlets). Figs. 203D-H, nymph: 203D, habitus; 203E, maxilla; 203F, right mandible; 203G, left mandible; 203H, abdominal gill II. Figs. 203A-B modified from DEMOULIN, 1955d; Fig. 203D, drawing by A. Dupuy.

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FAMILY OLIGONEURIIDAE Taxonomy: The family contains two extant subfamilies (Oligoneurinae and Chromarcyinae) and the extinct subfamily Colocrurinae (MCCAFFERTY, 1990). This essentially pantropical family is believed to have evolved in the South America-Africa-Madagascar landmass (EDMUNDS, 1972). The recently described Cretaceous fossil specimens of the oligoneuriid genus Colocrus from the Santana Formation in Ceará Crato, Brazil (MCCAFFERTY, 1990) lend more support to this hypothesis. Additionally, South America has seven [Colocrus (Colocrinae), Fittkauneuria, Homoeoneuria, Lachlania, Oligoneuria, Oligoneurioides and Spaniophlebia] of the twelve known oligoneuriid genera. Chromarycinae are restricted to Asia, and only Oligoneuriinae are found in South America. Adult Characteristics: 1) Fore wings with three or four compound longitudinal veins behind R1 (Figs. 204A, 205A, 206A, 208A, 209A) made up of pairs of convex and concave longitudinal veins or intercalaries (pairs are sometimes called “geminate” veins); 2) fore legs shorter than mesothoracic legs; 3) terminal filament vestigial or as long as cerci. Nymphal Characteristics (Figs. 204J, 205G, 207A, 208C): 1) Maxillary and labial palpi 2segmented; 2) gill tufts at bases of maxillae; 3) leading edge of fore femora and tibiae with double rows of long setae; 4) gills on abdominal segment I ventral, gills on segments II-VII dorsal. Distribution: North and South America, Europe, Asia, Africa. KEYS TO SOUTH AMERICAN GENERA Adults 1 – 2(1) – 3(1) – 4(3) – 5(4) –

39

Two caudal filaments ......................................................................................................... 2 Three caudal filaments ..................................................................................................... 3 Veins IRs/R3 (paired) diverging from vein R1 near its base (Fig. 210A); apex of male styliger plate convex (Fig. 210C); penes similar to Fig. 210C .................. Spaniophlebia Veins IRs/R3 (paired) diverging from vein R1 near middle of wing (Fig. 206A); apex of male styliger plate with a depression (Fig. 206C); penes similar to Fig. 206D ..... .................................................................................................................................. Lachlania Cross veins absent in fore and hind wings (Figs. 205A-B); male genital forceps absent (Fig. 205E) ..................................................................... Homoeoneuria (Notochora) Cross veins present in fore and/or hind wings (Figs. 204A-B, 208A, 209A-B); male genital forceps present or absent ..................................................................................... 4 Males ..................................................................................................................................... 5 Females ................................................................................................................................. 6 Male genital forceps absent (Fig. 204D); hind tibiae more than 2 times the length of hind femora; terga VIII-IX widely expanded ................................ Fittkauneuria 39 Male genital forceps present (Fig. 209C); hind tibiae shorter than hind femora; terga VIII-IX may present posterolateral extensions, but not very expanded .... Oligoneurioides

Although the males of Oligoneuria and the females of Fittkauneuria are unknown, it is very unlikely that they belong to the same genus, because the wings of Fittkauneuria (Fig. 204A) have numerous cross veins in the MA field, which are absent in Oligoneuria.

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Cross veins present in costal field and between veins R1 and paired veins IRs/R3 only (Fig. 209A) .............................................................................................Oligoneurioides Cross veins present in costal field and between veins R1 and paired veins R4+5/ MA1, and traces of cross veins in MP field (Fig. 208A) ......................... Oligoneuria 39

Nymphs 1 – 2 (1) –

Two caudal filaments (Fig. 207A) ................................................................. Lachlania 40 Three caudal filaments (Figs. 204J, 205G) ................................................................... 2 Gill lamellae on abdominal segments II-VII elongated, three times as long as broad (Fig. 205L) ....................................................................... Homoeoneuria (Notochora) Gill lamellae on abdominal segments II-VII rounded (Fig. 204J) ....... Fittkauneuria

CLAVES

PARA LOS

GÉNEROS

SUDAMERICANOS

Adultos 1 – 2(1) – 3(1) – 4(3) – 5(4) – 6(4) –

40 41

Dos filamentos caudales presentes ................................................................................. 2 Tres filamentos caudales presentes ................................................................................ 3 Venas IRs/R3 (pareadas) alejándose de la vena R1 cerca de la base de ésta (Fig. 210A); ápice de la placa estilígera del macho convexa (Fig. 210C); penes similares a los de Fig. 210C ............................................................................................... Spaniophlebia Venas IRs/R3 (pareadas) alejándose de la vena R1 a partir del centro del ala (Fig. 206A); ápice de la placa estilígera del macho con una depresión (Fig. 206C); penes similares a los de Fig. 206D ............................................................................... Lachlania Venas transversas ausentes en alas anteriores y posteriores (Figs. 205A-B); fórceps genitales del macho ausentes (Fig. 205E) ............................ Homoeoneuria (Notochora) Venas transversas presentes en alas anteriores y posteriores (Figs. 204A-B, 208A, 209A-B); fórceps genitales del macho presentes o ausentes .................................... 4 Machos ................................................................................................................................. 5 Hembras ............................................................................................................................... 6 Fórceps genitales del macho ausentes (Fig. 204D); tibias posteriores más de 2 veces la longitud del fémur posterior; tergos VIII-IX ampliamente expandidos ................. ........................................................................................................................ Fittkauneuria 41 Fórceps genitales del macho presentes (Fig. 209C); tibias posteriores más cortas que los fémures posteriores; tergos VIII-IX pueden tener extensiones posterolaterales, pero no muy expandidas ..............................................................................Oligoneurioides Venas transversas presentes en el campo costal y entre las venas R1 y las venas IRs/ R3 (pareadas) solamente (Fig. 209A) ...........................................................Oligoneurioides Venas transversas presentes en el campo costal y entre las venas R1 y las venas pareadas R4+5/Ma1 y trazas de venas transversas en el campo MP (Fig. 208A) ....... Oligoneuria41

The nymphs of Spaniophlebia also have two caudal filaments, based on unpublished data. Aunque los machos de Oligoneuria y las hembras de Fittkauneuria no se conocen, es muy improbable que ellos pertenezcan al mismo género, ya que las alas de Fittkauneuria (Fig. 204A) tienen numerosas venas transversas en el campo MA, ausentes en Oligoneuria.

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Dos filamentos caudales presentes (Fig. 207A) ......................................... Lachlania 42 Tres filamentos caudales presentes (Figs. 204J, 205G) ............................................. 2 Porción laminar de las branquias de los segmentos abdominales II-VII lanceoladas, tres veces más largas que anchas (Fig. 205L) ....................... Homoeoneuria (Notochora) Porción laminar de las branquias en los segmentos abdominales II-VII redondeadas (Fig. 204J) ............................................................................... Fittkauneuria

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Genus Fittkauneuria PESCADOR & EDMUNDS Fittkauneuria PESCADOR & EDMUNDS, 1994: 263. (Type-species: Fittkauneuria carina PESCADOR & EDMUNDS, original designation).

Taxonomy: The genus Fittkauneuria was described by PESCADOR & EDMUNDS (1994) based on mature nymphs, male imagos and eggs (extracted from mature nymphs). The genus presently includes two species, F. carina PESCADOR & EDMUNDS and F. adusta PESCADOR & EDMUNDS. Adult Characteristics: 1) Fore wing with several cross veins distributed more or less evenly throughout entire wing (Fig. 204A); 2) fore wing with four or more apparent longitudinal veins behind vein R1 (Fig. 204A); 3) vein IMP of fore wings weakly developed (Fig. 204A); 4) three caudal filaments; 5) genital forceps absent; penis lobes bulbous (Figs. 204D-E); 6) female sternum IX apically rounded. Nymphal Characteristics: 1) Frontoclypeal region of head not expanded, strongly tapered anteriorly and with thick dorsal setae and marginal hairs (Fig. 204J); 2) fore tarsi normally developed; 3) abdominal gills II-VII dorsolaterally attached on segments (Fig. 204J); 4) gill lamellae broad and plate-like, length more than one-half length of segment (Figs. 204F-J); 5) posterolateral projections on abdominal segments VIII and IX (Fig. 204J); and 6) terminal filament more than half length of cerci. Distribution: Brazil and Venezuela. Biology: The rheophilic nymphs of Fittkauneuria occur in small streams and medium-sized rivers with substrates ranging from coarse sand to boulders. Nymphs were collected in January through March and the single male imago in March. KEY

TO

FITTKAUNEURIA

Nymphs 1 –

42

Ventral tufts of abdominal gill II-VI well developed (Fig. 204I); dorsal lamellae oval (Figs. 204H-I); length of posterolateral spines on abdominal segment IX distinctly less than 1/3 of maximum width of segment ........... Fittkauneuria carina Ventral tufts of abdominal gills vestigial (Fig. 204G); dorsal lamellae ovate (Figs. 204F-G); length of posterolateral spines on abdominal segment IX distinctly more than 1/3 maximum width of segment ............................... Fittkauneuria adusta Las ninfas de Spaniophlebia, según datos sin publicar, también tienen dos filamentos caudales.

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FITTKAUNEURIA

Ninfas 1

Borla de branquias ventrales abdominales II-VI bien desarrolladas (Fig. 204I); láminas dorsales ovales (Figs. 204H-I); longitud de las espinas posterolaterales en el segmento abdominal IX claramente menor a 1/3 del ancho máximo del segmento .............................................................................................. Fittkauneuria carina C Sc R1

C

R3 IRs

A B

D

F

G

E

H

I

J

Plate 204. Fittkauneuria. 204A-E, male imago. Fittkauneuria carina: 204A, fore wing; 204B, hind wing; 204C, frontal view of head; 204D, genitalia, v.v.; 204E, detail of penes. 204F-J, nymph. F. adusta: 204F, abdominal gill IV, d.v.; 204G, same, v.v. F. carina: 204H, abdominal gill IV, d.v.; 204I, same, v.v.; 204J, habitus. Figs. 204A-J modified from PESCADOR & EDMUNDS, 1994.

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Borla de branquias ventrales abdominales vestigiales (Fig. 204G); láminas dorsales más bien redondeadas (Figs. 204F-G); longitud de las espinas posterolaterales en el segmento abdominal IX claramente mayor a 1/3 del ancho máximo del segmento .............................................................................................. Fittkauneuria adusta

Fittkauneuria adusta PESCADOR & EDMUNDS Fittkauneuria adusta PESCADOR & EDMUNDS 1994: 268 (nymph).

Taxonomy: Fittkauneuria adusta is presently known only in the nymphal stage. The nymph is distinguished from F. carina by the following characters: 1) Ventral gill tufts vestigial (Fig. 204G); 2) dorsal lamellae of gills ovate (Fig. 204F); 3) length of posterolateral spines of abdominal segment IX distinctly more than one-third maximum width of segment; and 4) brown markings of head extensive, covering vertex and extending anteriorly between bases of antennae. Distribution and Biology: Venezuela and Brazil. The nymphs were collected in medium sized-streams and rivers with rubbles, boulders and bedrocks substrates. Fittkauneuria carina PESCADOR & EDMUNDS Fittkauneuria carina PESCADOR & EDMUNDS, 1994: 268 (male imago, nymph).

Taxonomy: PESCADOR & EDMUNDS (1994) described the species from one male imago and nymphs. The nymph is differentiated from F. adusta by the following characters: 1) Ventral tufts present on abdominal gills II-VI (Fig. 204I); 2) dorsal lamellae of gills II-VII oval (Fig. 204H); 3) length of posterolateral spines on abdominal segment IX less than one-third maximum width of segment; and 4) brown markings of head less extensive and limited to area between eyes and ocelli. Distribution and Biology: The species has only been collected in the highlands of Venezuela. Knowledge of the biology of F. carina is limited. The nymphs have been collected in a small stream (30-40 cm in width and 7-10 cm in depth) nestled in a dense jungle, with substrates ranging from sand to boulders (PESCADOR & EDMUNDS, 1994). A few nymphs were collected from a small heavily shaded stream seeping from sandstone and flowing down a rocky chute with numerous shallow sloping falls and rock-edged pools. The adult was collected in mid-March. Genus Homoeoneuria EATON Homoeoneuria EATON, 1881: 197; EDMUNDS, BERNER & TRAVER, 1958: 376, EDMUNDS, JENSEN & BERNER, 1976: 182; PESCADOR & PETERS, 1980a: 358. (Type-species: Homoeoneuria salviniae EATON, original designation).

Taxonomy: EATON (1881) established the genus Homoeoneuria for the female imago of H. salviniae from Dueñas and Aceytano, Guatemala. PESCADOR & PETERS (1980a) subsequently revised the genus and established two subgenera, Homoeoneuria s.s. and Notochora. The subgenus Homoeoneuria, which has never been collected in South America, is represented by five species in North and Central America, and will not be discussed

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here. The monotypic subgenus Notochora is presently known only from Brazil and is represented by the species H. (N.) fittkaui PESCADOR & PETERS. Adult Characteristics: 1) Fore wings without cross veins (Fig. 205A); 2) fore wings with only three apparent longitudinal cross veins behind R1 (Fig. 205A); 3) three caudal filaments; 4) genital forceps absent, penis lobes apically excavated (Figs. 205E-F); 5) vein IMP of fore wings absent (Fig. 205A); and 6) female sternum IX posteromedially extended (Fig. 205D). Nymphal Characteristics: 1) Frontoclypeal region of head not expanded, slightly tapered anteriorly with few marginal hairs (Fig. 205H); 2) fore tarsi vestigial (Fig. 205J); 3) abdominal gills II-VII dorsally attached on segments; 4) gill lamellae narrow and lanceolate, slightly longer than length of segments (Fig. 205L); 5) posterolateral projections on abdominal segments VIII and IX; and 6) terminal filament more than half length of cerci. Distribution: Nearctic and Neotropical. Biology: The nymphs occur in sand-bottom beds of large streams and rivers. Nymphs of Homoeoneuria (N.) fittkaui were collected in a large moderately rapid stream with shifting sand bottom. Subgenus Homoeoneuria (Notochora) PESCADOR & PETERS Homoeoneuria (Notochora) PESCADOR & PETERS, 1980a: 385; DA-SILVA, 1992b: 693. (Type-species: Homoeoneuria (Notochora) fittkaui PESCADOR & PETERS, original designation).

Taxonomy: Homoeoneuria (Notochora) can be differentiated from the other subgenus of Homoeoneuria by the following combination of characters. In the imago: 1) Styliger plate weakly developed; penial arms weakly developed; penes with finger-like process (Figs. 205E-F); 2) female sternum IX posteromedially extended, apically pointed (Fig. 205D). In the nymph: 1) Tarsal claw with subapical denticle on mesothoracic leg. Homoeoneuria (Notochora) fittkaui PESCADOR & PETERS Homoeoneuria (Notochora) fittkaui PESCADOR & PETERS, 1980a: 385 (male, female); DA-SILVA, 1992: 693 (nymph).

Taxonomy: Homoeoneuria (N.) fittkaui was described by PESCADOR & PETERS (1980a) based upon male and female adult specimens including the eggs. The nymph was subsequently described by DA-SILVA (1992). The species can be distinguished from the rest of the Homoeoneuria species by the following combination of characters. In the imagos: 1) Posteromedian pale yellow spot of pronotum subequal to posterolateral spots (Fig. 205C); 2) abdominal terga with no markings; and 3) shape of penes as in Figs. 205E-F. In the nymph: 1) Antennal pedicel with short and thick setae (Fig. 205K); 2) posterior margins of superlingua of hypopharynx strongly curved (Fig. 205I); 3) tibia of prothoracic leg bowed, with maximum width subequal to maximum width of femur; 4) tarsal claw of metathoracic leg with small subapical denticle; and 6) abdominal terga with no markings. Distribution and Biology: The species occurs in northern and northeastern Brazil. The nymphs were collected in a large, moderately rapid stream with shifting sand bottom, at a water depth of about 70 cm (DA-SILVA, 1992). Collection records show that the adults emerge in February.

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C

D A

B

E

F

G

H

I J K

L

Plate 205. Homoeoneuria. Figs. 205A-F, imago: H. (H.) cahabensis: 205A, male fore wing, 205B, male hind wing. H. (Notochora) fittkaui: 205C, male pronotum; 205D, female abdominal sterna VII-IX; 205E, male genitalia, v.v.; 205F, detail of penes, d.v. Figs. 205G-L, nymph. H. cahabensis, nymph: 205G, habitus; 205H, thorax and head, l. v. H. (Notochora) fittkaui: 205I, hypopharynx; 205J, fore tarsus; 205K, antenna; 205L, abdominal gill II. (Leg segments on Fig. 205H: Cx = coxa, Tr = trochanter, Fe = femur, Tb = tibia, Ts = tarsus and CL = claw). Figs. 205A-F and 205H modified from PESCADOR & PETERS, 1980a; 205G from EDMUNDS et al., 1958; 205I-L from DA-SILVA, 1992b.

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Genus Lachlania HAGEN Lachlania HAGEN, 1868: 372; NEEDHAM & MURPHY, 1924: 32; DEMOULIN, 1952d: 3; KOSS & EDMUNDS, 1970: 55; EDMUNDS, BERNER & TRAVER, 1958: 381; PEREIRA, 1987: 1; PEREIRA, 1989A: 1; DA-SILVA & PEREIRA, 1993: 65. (Type-species: Lachlania abnormis HAGEN, original designation). Lachlania (Noya) DEMOULIN, 1955a: 23. Neophlebia NAVÁS, 1912a: 746. (Type-species: Neophlebia garciai NAVÁS, monobasic). Noya NAVÁS, 1912a: 746; ULMER, 1920c: 20; ULMER, 1920a: 129; ULMER, 1921: 249. (Type-species: Neophlebia garciai NAVÁS, objective synonym). Noyopis NAVÁS,1924b: 70. (Type-species: Noyopis fusca NAVÁS, original designation). Alloydia NEEDHAM, 1932: 275. (Type-species: Alloydia cacautana NEEDHAM, original designation).

Taxonomy: HAGEN (1868) established the genus Lachlania based on female imagos from Cuba. The absence of a terminal filament and presence of middle cross veins in the fore wings, according to HAGEN, separate the genus from other oligoneuriid genera. NAVÁS (1912a) established the genus Noya, that was subsequently synonymized by ULMER (1943) with Lachlania. Since then, KOSS & EDMUNDS (1970) and PEREIRA (1987) have reviewed the taxonomy of the genus. South America appears to be the center of diversity of Lachlania. Seven of the 14 nominal species of the genus occur in the region. One unidentified species, a nymph from Brazil, was designated as Lachlania (Noya) sp. by DEMOULIN (1955a). PEREIRA (1987) grouped the 14 known species of Lachlania into two groups based on the number of cross veins between veins R1 and MP2 of the fore wings. Currently, it appears that the above characters are the main basis of differentiating the various species of the genus, notably those occurring in South America. In a study of L. dencyanna and L. cacautana wings, however, KOSS & EDMUNDS (1970) indicated that there is considerable variation on the number of cross veins, and their taxonomic value must be exercised with caution. The development of a reliable key to Lachlania species of South America at this time would be premature until a comprehensive revision of the genus is addressed. Any attempt to identify the South American species of Lachlania requires access to the type specimens to avoid misidentifications and unnecessary creation of synonymies. Additionally, three (e.g., L. garciai, L. radai, L. santosi) of the seven species that occur in the region are known only from female specimens and one species (L. boanovae) is known from a female holotype and nymphal paratypes. Adult Characteristics: 1) Fore wings with relatively few cross veins, concentrated mainly between veins R1 and MA (Fig. 206A); 2) fore wing with four apparent longitudinal veins behind vein R1; 3) two caudal filaments; 4) genital forceps present (Fig. 206C); penis lobes similar to Fig. 206C; 5) vein IMP of fore wing well developed (Fig. 206A); 6) paired veins IRs/R3 diverging from vein R1 near middle of wing; and 7) female abdominal sternum IX apically cleft (Fig. 206D). Nymphal Characteristics: 1) Frontoclypeal region of head moderately expanded (Fig. 207A); 2) fore tarsi normally developed; 3) abdominal gills II-VII dorsally attached

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on segments (Fig. 207A); 4) gill lamellae small, rounded, and less than half length of each segment (Fig. 207A); 5) posterolateral projections on abdominal segments II-IX (Fig. 207A); and 6) terminal filament absent. Distribution: Nearctic and Neotropic. Biology: The nymphs are commonly found in fast currents, firmly attached to snags and sticks wedged between rocks or clinging to the undersides of rocks. Submerged vegetation caught in crevices of rocks is also a favorite nymphal microhabitat. In sandy bottom streams, the nymphs usually congregate in partly submerged sticks or leaf packs. Adult emergence has been reported in August and early September among Nearctic species and from January to May in the Neotropics. Mating flights have been reported in midmorning, but also at dusk. Lachlania boanovae DA-SILVA & PEREIRA Lachlania boanovae DA-SILVA & PEREIRA 1993: 295 (female, nymph).

Taxonomy: Lachlania boanovae was described by DA-SILVA & PEREIRA (1993) based on one female imago and nymphs. The species appears similar to L. santosi based on color pattern but could be distinguished from it by vein IMP which originates at 1/2 the distance from the furcation of MP1 and MP2. The presence of spine-like posterolateral projections on the IX abdominal segment (Fig. 206D) separate L. boanovae from the other species of the genus (DASILVA & PEREIRA, 1993). The nymphs have apically black fore tarsi and a pair of papillae on the mesothoracic sternum. The nymphs are similar to L. iops (a species from Mexico), based on the shape of lateral projections of the thorax, rows of spines on the abdominal sterna and the relative size of the posterolateral projections on abdominal segment IX. Distribution and Biology: Brazil (Rio de Janeiro). The nymphs were collected in a small, pebbly- and sand-bottomed stream with moderate flow and patches of alloch-

A

IRs

R3

R1

C

IMP

B

D

Plate 206. Lachlania, imagos. Figs. 206A-D. L. dominguezi: 206A, fore wing; 206B, hind wing; 206C, male genitalia. L. boanovae, 206D: female abdominal sternum IX. (IRs = vein IRs). Figs. 206A-C modified from DOMÍNGUEZ et al., 1994; 206D from DA-SILVA & PEREIRA, 1993.

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tonous organic matter. Some nymphs were found in submerged roots of vegetation along the stream banks. A female imago (the holotype) was collected in April flying approximately 3 meters above the stream surface around 13:00 h. Lachlania cacautana (NEEDHAM) Alloydia cacautana NEEDHAM, 1932: 275 (male, female). Lachlania cacautana; DEMOULIN, 1952a: 3; HUBBARD, 1982: 269; PEREIRA, 1987: 5 (male, female).

Taxonomy: Lachlania cacautana was originally described in the genus Alloydia based on male and female adults from the upper Cacauta river in the high Andes of Colombia (NEEDHAM, 1932). DEMOULIN (1952) subsequently transferred the species to the genus Lachlania. The presence of several cross veins (approximately 20) between veins R1 and MA of the fore wings separates L. cacautana from the other South American species of the genus. The nymph remains unknown. Distribution and Biology: The species has only been collected in the Valle Papas, upper Cacauta river in the high Andes of Colombia. The adults were collected in March. According to NEEDHAM (1932), the adults flew approximately an inch above the water and followed perfectly the undulation of the swift riffles. They flew up and down-stream in copula in a manner similar to dragonflies, and oviposited by dipping the tip of the abdomen in the swift water during flight. Lachlania dominguezi PEREIRA Lachlania dominguezi PEREIRA, 1989a: 1 (male); PEREIRA, 1989b: 1 (nymph); HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 210; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 36 (male, nymph).

Taxonomy: Lachlania dominguezi was described by PEREIRA (1989a) based on male imagos from Argentina. The probable nymph, collected at the same time and in the same locality was described later (PEREIRA, 1989b). No discussion of its affinities was provided. According to the author, the adults can be distinguished from the other species of the genus by: 1) Number of cross veins between main longitudinal veins (Fig. 206A); and 2) position of IRs/R3 and origin of IMP (Fig. 206A) Distribution and Biology: Argentina (Salta). The adults were collected flying at dusk over a fast flowing and clear sandy-bottomed river with few scattered rocks coated with filamentous algae. Lachlania garciai (NAVÁS) Neophlebia garciai NAVÁS 1912a: 746 (female); ALBA-TERCEDOR & PETERS, 1985: 222. Noya garciai NAVÁS, 1912a: 746 (female). Lachlania garciai; ULMER, 1943: 38 (female); KOSS & EDMUNDS, 1970: 72; HUBBARD, 1982: 269; PEREIRA, 1987: 5 (female).

Taxonomy: Lachlania garciai was inadvertently described by NAVÁS (1912a) in Neophlebia, although he subsequently corrected the name to Noya in the same paper as the name Neophlebia was preoccupied; illustrations of the fore- and hind wings in the

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A

Plate 207. L. cf. dominguezi. Fig. 207A, nymphal habitus. Drawing by A. Dupuy.

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original description were correctly labeled Noya garciai. The species was described from a female imago and has several cross veins between veins R1 and MP2 of the fore wings similar to L. cacautana. Distribution: Colombia (Pasto, 2300 m). Lachlania pallipes (EATON) Spaniophlebia pallipes EATON, 1883-1888: 34 (male); COCKERELL, 1926: 189 Noya pallipes; ULMER, 1920c: 129; ULMER, 1920b: 20 (nymph); LESTAGE, 1924c: 38; LESTAGE, 1931a: 60; KIMMINS, 1960: 276. Lachlania pallipes; NEDHAM & MURPHY, 1924: 32; LESTAGE, 1930b: 434; ULMER, 1943: 39 (male); KOSS & EDMUNDS, 1970: 72; HUBBARD, 1982: 269; PEREIRA, 1987: 5 (male).

Taxonomy: EATON (1883) originally described L. pallipes in the genus Spaniophlebia based on a male imago from Ecuador. The nymph was first described by ULMER (1920c) (in the genus Noya) and was collected from Argentina. NEDHAM & MURPHY (1924) suggested that a Lachlania specimen from Guatemala was congeneric with the nymph from Argentina on which ULMER had based his description of Noya pallipes. The current descriptions of both the nymph and adults of L. pallipes are inadequate, which make it difficult to distinguish the species from its congeners. Distribution: Argentina and Ecuador. Lachlania radai (NAVÁS) Noya radai NAVÁS, 1926a: 102; LESTAGE, 1931b: 60; BERTHELEMY, 1965: 2. Lachlania radai; ULMER, 1943: 39 (female); KOSS & EDMUNDS, 1970: 72; HUBBARD, 1982: 269; PEREIRA, 1987: 5 (female).

Taxonomy: The species was originally described by NAVÁS (1926a) in the genus Noya based on a female imago from Peru. ULMER (1943) transferred the species to the genus Lachlania and provided more illustrations of the female. Since the species is known only from a female specimen and is inadequately described, its taxonomic status is dubious at best. Distribution: Peru. Lachlania santosi PEREIRA Lachlania santosi PEREIRA, 1987: 314 (female); PEREIRA, 1989: 326; DA-SILVA & PEREIRA, 1993 : 295.

Taxonomy: PEREIRA (1987) described Lachlania santosi based on female imagos from Brazil. According to her, Lachlania santosi can be distinguished from its congeners by: 1) Number of cross veins between veins R1 and MA of fore wings; and 2) point of basal attachment of vein IMP of the forewing. These characters must be used with caution. Distribution and Biology: Brazil (Rio de Janeiro). The adults were collected in April at 10:00 h flying over water at the small Santana river, about 95 km from the city of Rio de Janeiro. The species is currently known only from its type locality.

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Genus Oligoneuria PICTET Oligoneuria PICTET, 1843: 291; EATON, 1883-88: 29. (Type-species: Oligoneuria anomala PICTET, original designation).

Taxonomy: PICTET (1843) described the genus Oligoneuria based on a female imago from Brazil. The original description is inadequate to distinguish the genus with confidence. A study of the holotype enabled us to have a better understanding of the definition of the genus. Adult characteristics: 1) Fore wings with relatively few cross veins, concentrated mainly between R1 and R4+5 (Fig. 208A); 2) fore wings with four or more apparent longitudinal veins behind R1; 3) three caudal filaments; 4) vein IMP and many cross veins weakly indicated in MP field; and 5) female sternum IX apically cleft (Fig. 208B). Distribution: Brazil, Paraguay? Oligoneuria anomala PICTET Oligoneuria anomala PICTET, 1843: 291 (female); WALKER, 1853: 585; HAGEN, 1854: 318; HAGEN, 1855: 269 (female); HAGEN, 1861: 304; HAGEN, 1868: 373; EATON, 1871: 55; EATON, 1883-88: 30; LESTAGE, 1924c: 38; NEEDHAM & MURPHY, 1924: 29 (egg); LESTAGE, 1931b: 60; SPIETH, 1943: 12; ULMER, 1943: 37; DEMOULIN, 1955a: 23; PUTHZ, 1973: 91 (female, egg); HUBBARD, 1982: 269; MOL, 1986: 68.

Taxonomy: When PICTET (1843) established the genus Oligoneuria, the type species, O. anomala from Rio de Janeiro was inadequately described. ULMER (1921) expanded the description of the type and NEEDHAM & MURPHY (1924) subsequently redescribed the species based on another female imago in alcohol from an unspecified locality. PUTHZ (1973) also redescribed the species based on a female imago from Puerto Stroessner, Paraguay, which was presumably conspecific with O. anomala. One of us (MLP) had the opportunity to study the holotype (pinned specimen, Natural History Museum, Vienna) and additional characteristics of the species are herein included: 1) Thorax: pro-, mesoand methathorax light brown with no color pattern, except for two darker submedian pronotal bars; depressed postscutellum with posterolateral corners slightly elevated; 2) wings: longitudinal veins yellow-tan to pale tan; 3) abdomen: only coloration of abdominal segment II visible with dark brown median area, lighter area laterally with a dark loop close to lateral margin of tergum; most segments too shriveled to determine pattern, but coloration much darker than on thoracic segments; 4) female abdominal sternum IX with long posterolateral projections as illustrated by PICTET (1843) and PUTHZ (1973) (Fig. 208B); hyaline membrane at base of extension of sternum IX. Distribution: Brazil, Paraguay? Oligoneuriidae nymph (= Spaniophlebia nymph of DEMOULIN; Oligoneuria sp. of SPIETH 1943) SPIETH (1943) described an oligoneuriid nymph from Surinam as Oligoneuria sp. (anomala?) which according to him “may represent the nymph of Oligoneuria.” The

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nymph has a frontoclypeal region that is enormously expanded and flattened dorsoventrally forming a dome-shaped structure (Fig. 208D). DEMOULIN (1955a) transferred this nymph to Spaniophlebia and described another specimen which he believed to be the same species from the type locality of Oligoneurioides amazonicus. SPIETH ’s specimen is missing, and only a few mouthparts remain on slide which are insufficient for generic determination as the nymphs of neither Spaniophlebia, Oligoneuria, nor Oligoneurioides are known. Although illustrated (Fig. 208C), this specimen is not included in the generic keys.

C MA

A

R1 R3 IRs

CuP R4+5

CuA

A

IMP

MP2

MP1

B

C

Plate 208. Oligoneuria anomala. Figs. 208A-B, imago: 208A, fore wing; 208B, female abdominal sternum IX. 208C, Oligoneuriidae (Oligoneuria sp. of SPIETH) nymphal habitus. Figs. 208A drawn from the holotype by J. G. Peters; 208B modified from PUTHZ, 1973 and 208C modified from SPIETH, 1943.

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Genus Oligoneurioides DEMOULIN Oligoneurioides DEMOULIN, 1955a: 24. (Type-species: Oligoneurioides amazonicus DEMOULIN, original designation).

Taxonomy: The monotypic genus Oligoneurioides was established by DEMOULIN (1955a) based on male and female imagos from Rio Asaïhsal-Jutahy, Brazil, and material previously treated by EATON (1883) as the male of Oligoneuria anomala. Since then, no other taxonomic paper on the genus has been published. Nevertheless, the wing venation seems to be very characteristic. Adult Characteristics: 1) Fore wings with very few cross veins, mainly concentrated between veins R1 and IRs/R3 (Fig. 209A); 2) fore wings with four or more apparent longitudinal veins behind R1; 3) three caudal filaments; 4) vein IMP of fore wings absent in male (Fig. 209A) or sometimes with a very weak trace in female; and 5) genital forceps tubular, penis lobes apically tapered and hooked (Fig. 209C). Distribution: Brazil. Oligoneurioides amazonicus DEMOULIN Oligoneurioides amazonicus DEMOULIN, 1955a: 24 (male); HUBBARD, 1982a: 269. Oligoneuria anomala; EATON, 1883: 30 (male, female) non PICTET.

Taxonomy: DEMOULIN (1955a) provided the description of the species, based mainly on coloration. The shape of the male styliger plate and penes should also prove characteristic for the species. Distribution and Biology: Brazil (Rio Asaïhsal-Jutahy). The species is known only from adult specimens that were collected in February.

R1

C

R3 IRs

A

B Plate 209. Oligoneurioides amazonicus. Figs. 209A-C, imago: 209A, fore wing; 209B, hind wing; 209C, male genitalia, v.v. Figs. 209A-C drawn by J. G. Peters; 209B modified from DEMOULIN, 1955a.

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Genus Spaniophlebia EATON Spaniophlebia EATON, 1881: 191; EATON, 1883: 33; DEMOULIN, 1952a: 2; DOMÍNGUEZ, HUBBARD & PESCADOR, 1994: 37. (Type-species: Spaniophlebia trailiae EATON, original designation).

Taxonomy: EATON (1881) established the genus Spaniophlebia for the male imagos of S. trailiae from Rio Solimões, São Paulo, Brazil. Since then, two other species, S. assimilis and S. escomeli, have been described based on adults, the latter species known only from female imagos. The nymphs of Spaniophlebia are still not described. The descriptions of the Spaniophlebia species are inadequate, and one must exercise caution in identifying them. Adult Characteristics: 1) Fore wings with relatively few cross veins, concentrated between R1 and R4+5 (Fig. 210A); 2) fore wings with four or more apparent longitudinal veins behind vein R1; 3) vein IMP of fore wings well developed (Fig. 210A); 4) vein IRs/ R3 of fore wings basally attached near the base of R1; 5) two caudal filaments; and 6) genitalia as in Fig. 210C. Distribution: Brazil, Colombia and Peru. Spaniophlebia assimilis BANKS Spaniophlebia assimilis BANKS, 1913: 84 (male); LESTAGE, 1924c: 38; LESTAGE, 1931b: 60; DEMOULIN, 1955a: 26; HUBBARD, 1982b: 269.

Taxonomy: Spaniophlebia assimilis was described by BANKS (1913) based on a male imago from Camp 14, Rio Madeira, Brazil. According to BANKS, S. assimilis differs from S. trailiae in not having the cubitus vein forked, and an unmarked wing. However, his figure shows a forked Cu, so it is not clear to what he is referring. In his figure, the MP field is narrow and IMP appears free basally and attached to MP1 by a short cross vein. The wing membrane is hyaline with dark veins. Distribution: Brazil.

C R1 R3 IRs R4+5

A IMP

B Plate 210. Spaniophlebia trailiae. Figs. 210A-C adults: 210A, fore wing; 210B, hind wing; 210C, male genitalia, v.v. (IRs = vein IRs). Figs. 210A-C modified from EATON, 1883-1888.

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Spaniophlebia escomeli COCKERELL Spaniophlebia escomeli COCKERELL, 1926: 189 (female); HUBBARD, 1982a: 269.

Taxonomy: Spaniophlebia escomeli was described by COCKERELL (1926) based on a female imago from Arequipa, Peru. The description of the species is very brief. According to COCKERELL (1926), however, S. escomeli can be recognized by the greater number of cross veins (7) between the radial and medial veins of the fore wings, 4 cross veins in the next field and finally 2 small cross veins (wing not illustrated) and the dark markings of the wings. Distribution: Peru. Spaniophlebia trailiae EATON Spaniophlebia trailiae EATON, 1881: 191 (male); EATON, 1883: 33 (male); ESBEN-PETERSEN, 1912: 33; ULMER, 1920c: 129; COCKERELL, 1926: 189; NAVÁS, 1930c: 73 (male); LESTAGE, 1931b: 60; KIMMINS, 1960: 277; HUBBARD, 1982a: 269; HUBBARD, DOMÍNGUEZ & PESCADOR, 1992: 210. Spaniophlebia traili [sic]; BANKS, 1913: 85. Spaniophlebia trailliae [sic]; DALLAS, 1941: 131.

Taxonomy: Spaniophlebia trailiae was described from the Rio Solimões, Brazil. Later, a specimen matching its description was reported from Bompland, Misiones, Argentina (ESBEN-PETERSEN, 1912). NAVÁS (1930c) also reported the species but gave no locality, only mentioning that the specimen was sent to him by Dr. Angelina Suarez from Buenos Aires. Following EATON (1883), S. trailiae can be recognized by dark markings around the cross veins of the fore wings, the smoky gray coloration of the wing membrane, and cross veins as shown in Fig. 210A. Distribution: Argentina (Misiones and Buenos Aires).

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FAMILY ONISCIGASTRIDAE Taxonomy: This family includes three genera: Oniscigaster (from New Zealand), Tasmanophlebia (Australia) and Siphlonella (South America). Adult Characteristics: 1) Fore wings slender and triangular with numerous veins (Fig. 211A); 2) hind wings large (Fig. 211B); 3) hind legs with tarsal segment I (partly fused with tibiae) subequal or longer than tibiae; 4) genital forceps four-segmented; and 5) tarsal claws dissimilar, one blunt, the other acute. Nymphal Characteristics: 1) Head hypognathous; 2) abdominal tergum I with elevated hind margin; 3) gills on abdominal segments I-IV (in South America) or I-VII; 4) abdominal gill I operculate (in South America) or semioperculate; 5) median tubercles present on hind margin of abdominal terga II-IV or II-IX; 6) lateral margins of abdominal segments expanded and posteriorly pointed. Distribution: New Zealand, Australia, and Southern Argentina and Chile. Genus Siphlonella NEEDHAM & MURPHY Siphlonella NEEDHAM & MURPHY, 1924: 30. (Type-species: Siphlonella ventilans NEEDHAM & MURPHY, original designation).

Taxonomy: This genus is represented by two species, Siphlonella guttata very poorly known from female adults, and S. ventilans known from all the stages. Adult Characteristics (Fig. 220E): 1) Stigmatic area with anastomosed cross veins (Fig. 211A); 2) costal projection of hind wings rounded (Fig. 211B); 3) penes almost completely divided (Fig. 211C); 4) terminal filament vestigial.

A

B

C

Plate 211. Siphlonella ventilans male imago. Figs. 211A-C. 211A, fore wing; 211B, hind wing; 211C, genitalia, v.v.

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Nymphal Characteristics (Fig. 219A): 1) Gills on abdominal segments I-IV; 2) abdominal gill I operculate; 3) median tubercles on hind margin of abdominal terga II-IV (rarely II-V). Distribution: Southern Argentina and Chile. Biology: Nymphs of this species inhabit cold rivers, streams and lakes. They are very good swimmers that prefer the shallow margins and backwater zones, and are able to hide themselves under the sand. Mature nymphs generally crawl out of the water to emerge on rocks or sticks before molting to the subimago, although some were observed to molt on the water surface. The emergence of the subimago from the nymphal exuvia takes approximately 10 minutes, except those emerging at water surface that molted in a few seconds. The emergence period extended from morning to 18:00, and most subimagos then flew into the surrounding forest. The mating flight is still unknown. Siphlonella guttata (PICTET) Nomen Dubium Baetis guttata PICTET, 1843-1845: 187; FLOWERS & PETERS, 1981: 153. Ephemera-Baetis guttata; BLANCHARD, 1851: 106. Ecdyonurus guttatus; WALKER, 1853: 559; LESTAGE, 1931B: 42; HUBBARD, 1982a: 263. Heptagenia guttata; EATON, 1871: 156. Ecdyurus guttatus; EATON, 1883-1888: 301 (female); ULMER, 1904: 4.

Taxonomy: PICTET (1843) described Siphlonella guttata in the genus Baetis. This species cannot be identified from the description and figure in the literature, although EATON’s (1871) description of the body markings suggests that it is probably a Siphlonella (FLOWERS & PETERS, 1981). For these reasons we consider this species as a nomen dubium. Distribution: Chile. Siphlonella ventilans NEEDHAM & MURPHY Siphlonella ventilans NEEDHAM & MURPHY, 1924: 31 (nymph); ULMER, 1938: 97 (male, nymph); LESTAGE, 1931b: 49 (nymph); LESTAGE, 1935a: 97; HUBBARD, 1982a: 273. Siphlonella fazi NAVÁS, 1930d: 360 (male); ULMER, 1938: 107. Siphlonurus fazi; LESTAGE, 1935a: 96; PETERSEN & GAEDIKE, 1968: 960. Metamonius hollermayeri NAVÁS, 1936a: 138.

Taxonomy: Siphlonella ventilans is known from both nymphs and adults. Adult can be easily recognized because the anterior margin of fore wings has dark shading on cross veins of C, Sc and R1 sectors and stigmatic area (Fig. 211A). Distribution: Chile and Argentina (Patagonia).

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FAMILY POLYMITARCYIDAE Taxonomy: This widespread family is composed of three subfamilies: Polymitarcyinae (Ephoron, Holartic), Asthenopodinae (Povilla, Ethiopian-Oriental, and Asthenopus, Neotropical), and Campsurinae (Campsurus and Tortopus, Neotropical and Neartic). Ephoron umbrata HAGEN (1888) was described from Brazil, but here it is considered as a nomen dubium under the subheading “Incertae sedis”. Adult Characteristics: 1) Base of veins MP2 and CuA of fore wings strongly divergent from base of vein MP1 (Figs. 212A, 213A, 217A); 2) all legs of both sexes (except fore legs of male) reduced or vestigial (Figs. 213E, 217M); and 3) male forceps 1to 3-segmented (Figs. 212B, 213F, 217C). What may be interpreted as the basal segment of the forceps in Tortopus and Campsurus, is actually the reduced styliger plate; as it is generally strongly projected posteriorly, the name parastyli is used for this structure (MCCAFFERTY & BLOODGOOD, 1989; KLUGE, 2004). Nymphal Characteristics (Figs. 212I, 216A, 233D-E): 1) Labial and maxillary palpi 2-segmented; 2) legs modified to burrow (Fig. 3C); 3) abdominal gills arising dorsally; and 4) abdominal gills I vestigial. Distribution: Holarctic, Ethiopian-Oriental, and Neotropical. KEYS TO SOUTH AMERICA GENERA Imagos 1 – 2(1) –

Middle and hind legs reduced, broad and fin-shaped; tibiae and tarsi absent (Fig. 213E) .................................................................................................................... Campsurus Middle and hind legs not so reduced, all segments present (Fig. 217M) ............... 2 Fore wings with numerous cross veins connecting longitudinal and intercalary veins (Figs. 212A, C, E); veins near anal margin of hind wings not reticulated (Fig. 212F); forceps stout without projected parastyli (Figs. 212B, D, G) ...... Asthenopus Fore wings with numerous cross veins connecting longitudinal and intercalary veins except between veins CuA and A (Figs. 217A, D); veins near anal margin of hind wings reticulated (Figs. 217B, E); forceps long and slender, generally with parastyli strongly projected posteriorly (Figs. 217C, I) .................................. Tortopus

Nymphs 1 – 2(1)

–

Mandibular tusks short, broad, and robust (Figs. 3U, 212H-I) ................ Asthenopus Mandibular tusks relatively elongate and slender (Figs. 3T, 215N, 217N) ............ 2 Mandibular tusks with prominent basal or sub-basal tubercle on median margin (sometimes tubercle absent), from some to many apical crenulations, numerous setae on lateral margin of mandibles (Fig. 215N); abdominal gill I bifurcated (Fig. 215O) .................................................................................................................... Campsurus Mandibular tusks with single prominent subapical tubercle on median margin, few long setae on lateral margin of mandibles (Fig. 217N); abdominal gill I simple (Fig. 217O) ............................................................................................................... Tortopus

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PARA LOS

GÉNEROS

SUDAMERICANOS

Imagos 1 – 2(1)

–

Patas medias y posteriores reducidas, anchas y en forma de aleta; tibias y tarsos ausentes (Fig. 213E) ........................................................................................... Campsurus Patas medias y posteriores no tan reducidas, todos los segmentos presentes (Fig. 217M) .................................................................................................................................... 2 Alas anteriores con numerosas venas transversas conectando todas las venas longitudinales e intercalares (Figs. 212A, C, E); alas posteriores con venas cercanas al margen anal no reticuladas (Fig. 212F); fórceps robustos sin parastilo proyectado posteriormente (Figs. 212B, D, G) ............................................ Asthenopus Alas anteriores con numerosas venas transversas conectando todas las venas longitudinales e intercalares excepto entre las venas CuA y A (Figs. 217A, D); alas posteriores con venas cercanas al margen anal reticuladas (Figs. 217B, E); fórceps largos y delgados, generalmente con parastilo fuertemente proyectado (Figs. 217C, I) ..................................................................................................................... Tortopus

Ninfas 1 – 2(1)

–

Colmillos mandibulares cortos, anchos y robustos (Figs. 3U, 212H-I) ..... Asthenopus Colmillos mandibulares relativamente alargados y delgados (Figs. 3T, 215N, 217N) .... 2 Colmillos mandibulares con un prominente tubérculo basal o subbasal en el margen mediano (a veces puede estar ausente), y de algunas a muchas crenaciones apicales, numerosas setas en el margen lateral de las mandíbulas (Fig. 215N); branquia abdominal I bifurcada (Fig. 215O) ................................................ Campsurus Colmillos mandibulares con un único tubérculo subapical prominente en el margen mediano, pocas setas largas en el margen lateral de las mandíbulas (Fig. 217N); branquia abdominal I simple (Fig. 217O) ........................................... Tortopus SUBFAMILY ASTHENOPODINAE

Taxonomy: This subfamily is currently known from two extant genera: Asthenopus EATON (3 species, Neotropical) and Povilla NAVÁS (8 species, Ethiopian-Oriental). Also, one fossil genus was described (Asthenopodichnium) from traces doubtfully classified as Ephemeroptera (KLUGE, 2004). Adult Characteristics: 1) Pronotum short, ring-like, not broader posteriorly than in front; 2) fore legs of male about half as long or as long as fore wings; 3) cross veins of fore wings connected to margin along entire outer margin of wing (Figs. 212A, C, E); 4) styliger plate reduced to a pair of parastyli (segments at the base of forceps), and a small median lobe; 5) forceps 1-segmented and stout (Figs. 212B, D, G); and 6) penes cylindrical (Figs. 212B, D, G). Nymphal Characteristics: 1) Mandibular tusks short, broad, and robust, ending in 2 or 3 points (Fig. 212H). Distribution: Neotropical, Ethiopian-Oriental.

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A R2

559

B

A

B MC

MP1

CuP IMP

MP2

CuA

C

D E

F

H

G

I Plate 212. Asthenopus, imagos. Figs. 212A-I: A. curtus: 212A, male fore wing; 212B, male genitalia, v.v. (penes folded, left forceps omitted). A. gilliesi: 212C, male fore wing; 212D, male genitalia, v.v. A. picteti: 212E, male fore wing; 212F, male hind wing; 212G, male genitalia, v.v. Nymph, A. curtus: 212H, right mandible, d.v.; 212I, general aspect. Fig. 212B modified from KIMMINS, 1960; 212I from SATTLER, 1967.

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Genus Asthenopus EATON Asthenopus EATON, 1871: 59; LESTAGE 1922b: 142; TRAVER, 1950: 605; TRAVER, 1956b: 7; HUBBARD & DOMÍNGUEZ, 1988: 209; DOMÍNGUEZ, 1988a: 24. (Type-species: Palingenia curta HAGEN, original designation). Asthenopodes ULMER, 1924: 26; TRAVER, 1950: 611; TRAVER, 1956b: 1; HUBBARD, 1975: 111; HUBBARD & DOMÍNGUEZ, 1988: 209; DOMÍNGUEZ, 1988a: 24. (Type-species: Palingenia albicans PICTET, original designation).

Taxonomy: The genus Asthenopus was established by EATON (1871) for the species Palingenia curta HAGEN, 1861, and remained monotypic until 1988. ULMER (1924) established the genus Asthenopodes, also for a single species, Palingenia albicans PICTET. These genera were separated by ULMER (1924, 1932, 1942) by characters of legs and wing venation, characters that were completed by TRAVER in 1956. In 1988, DOMÍNGUEZ described a new species, Asthenopus gilliesi, with characters intermediate between these two Neotropical genera, placing in doubt the validity of their separation. In the same year, HUBBARD & DOMÍNGUEZ (1988) synonymized these genera, so that Asthenopus is presently represented by three species, of which only A. curtus is known from both nymphs and adults, the others are known only from adults. Adult Characteristics: 1) Forceps stout (Figs. 212B, D, G); 2) penes relatively simple, with triangular base and finger-like projection (Figs. 212B, D, G); and 3) numerous cross veins in cubito-anal area of fore wings (Figs. 212A, C, E). Nymphal Characteristics: 1) Short robust mandibular tusks (Figs. 3U, 212H). Distribution and Biology: Argentina, Colombia, French Guiana, Guyana, Uruguay, Paraguay, Brazil, Surinam, Venezuela, and Ecuador. The nymphs (Fig. 212I) burrow on relatively soft plant tissue (rotten wood, live Eichornia, Typha, etc.) and inhabit great rivers and lakes. KEY

ASTHENOPUS

TO

Imagos 1 – 2(1) –

CLAVE

Marginal intercalary veins of fore wings reaching posterior margin of wing and usually as long as or longer than distance between longitudinal veins (Figs. 212C, E) ........... 2 If the marginal intercalary veins of fore wings reach the posterior margin then they are usually shorter than distance between longitudinal veins (Fig. 212A) ...... ..................................................................................................................... Asthenopus curtus Fore legs of male 0.5 as long as fore wings; penes broad at base; penes thick, narrowing toward apex (Fig. 212D) .................................................... Asthenopus gilliesi Fore legs of male almost as long as fore wings; penes slender from base to apex (Fig. 212G) ................................................................................................ Asthenopus picteti PARA

ASTHENOPUS

Imagos 1

Venas transversas alcanzando el margen posterior de las alas anteriores y usualmente más largas que la distancia entre las venas longitudinales (Figs. 212C, E) ................ 2

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Si las venas transversas alcanzan el margen posterior de las alas anteriores entonces usualmente son más cortas que la distancia entre las venas longitudinales (Fig. 212A) ................................................................................................ Asthenopus curtus Patas anteriores del macho 0,5 tan largo como las alas anteriores; penes gruesos en la base, afinándose apicalmente (Fig. 212D) ............................... Asthenopus gilliesi Patas anteriores del macho casi tan largas como las alas anteriores; penes delgados desde la base hacia el ápice (Fig. 212G) ............................................. Asthenopus picteti

Asthenopus curtus (HAGEN) Palingenia albifilum var.; WALKER, 1853: 554. Palingenia curta HAGEN, 1861: 304. Campsurus curtus; EATON, 1868: 84 (male); EATON, 1883: 40 (male); ULMER, 1921: 240. Asthenopus curtus; EATON, 1871: 59 (male); ULMER, 1920c: 107; ULMER, 1921: 240; LESTAGE, 1922b: 142; LESTAGE, 1923: 124; LESTAGE, 1924c: 39; ULMER, 1942: 105 (male); SPIETH, 1943: 1; TRAVER, 1950: 606; TRAVER, 1956b: 7; KIMMINS, 1960: 312 (male); DEMOULIN, 1966a: 4; SATTLER, 1967: 104; BERNER, 1978: 103 (male, female); HUBBARD, 1982a: 270; DOMÍNGUEZ, 1988a: 24; HUBBARD & DOMÍNGUEZ, 1988: 207 (female); DOMÍNGUEZ, 1989a: 173 (male, female). Campsurus amazonicus HAGEN, 1888: 230 (male). Asthenopus amazonicus; ULMER, 1920c: 107; LESTAGE, 1923: 124; ULMER, 1942: 106 (male, female); TRAVER, 1950: 606 (female); TRAVER, 1956b: 7; DEMOULIN, 1966a: 4; SATTLER, 1967: 104; IRMLER, 1975: 368. Asthenopus amazonicus?; DEMOULIN, 1955a: 30 (male, female). Asthenopus sp.; TRAVER, 1950: 606; TRAVER, 1956b: 7; ROBACK, 1966: 131 (nymph).

Taxonomy: Asthenopus curtus is the only member of the genus known from males, females, and nymphs. Originally, this species was described by HAGEN (1861) as Palingenia curta and later described by the same author (1888) as Campsurus amazonicus. For a long time doubts existed about the validity of these two species (ULMER, 1942; TRAVER, 1950; DEMOULIN, 1955a; SATTLER, 1967), until finally BERNER (1978), based on abundant material reared from nymphs, clarified the error and established the synonymy of A. amazonicus with A. curtus. This species can be distinguished from the others in the genus by the following combination of characters: 1) Marginal intercalary veins of fore wings always shorter than distance between longitudinal veins (Fig. 212A); 2) vein IMP joined to vein MP1 in fore wings (Fig. 212A); 3) fore wings of males with 1-3 cross veins (females with 3-4) between Rs and MA basal to fork of Rs (Fig. 212A); and 4) penes (Fig. 212B) curved and thick at base, narrowing apically. The nymphs (Fig. 212I) of A. curtus are the only known for this genus, in spite of extensive material collected in this subfamily. Distribution and Biology: Argentina (Corrientes, Formosa), Guyana, Paraguay, Brazil, Surinam, Venezuela, Ecuador. The nymphs of Asthenopus are miners in plant tissue such as Eichornia and Typha, or in submerged timber. The tunnels are U-shaped and covered by pieces of wood joined by adhesive secretions of the Malpighian tubules. Apparently, feeding is by means of a funnel formed by a system of setae that capture food particles.

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Asthenopus gilliesi DOMÍNGUEZ Asthenopus gilliesi DOMÍNGUEZ, 1988a: 21 (male); HUBBARD & DOMÍNGUEZ, 1988: 207 (male).

Taxonomy: Asthenopus gilliesi is known only from male imagos, which can be distinguished from other species of the genus by the following combination of characters: 1) Marginal intercalary veins of fore wings generally longer than distance between longitudinal veins (Fig. 212C); 2) vein IMP not joined to vein MP1 in fore wings (Fig. 212C); 3) fore wings of males with one cross vein between Rs and MA basal to fork of Rs (Fig. 212C); 4) penes (Fig. 212D) curved and thick at base, narrowing apically. This species shows characters intermediate between those considered generic for Asthenopus and Asthenopodes, which led HUBBARD & DOMÍNGUEZ (1988) to synonymize both genera. Distribution: Uruguay (Artigas). Asthenopus picteti (HUBBARD) Palingenia albicans; PICTET, 1843: 149 (misidentification); WALKER, 1853: 548 (female) [partim]; HAGEN, 1861: 304. Campsurus albicans; EATON, 1871: 58 (male). Asthenopus albicans; ULMER, 1920c: 107; ULMER, 1921: 239; LESTAGE, 1924c: 39. Asthenopodes albicans; ULMER, 1924: 26; TRAVER, 1956b: 1 (male). Asthenopodes sp.; TRAVER, 1950: 611 (male); TRAVER, 1956b: 5 (female); TRAVER, 1956b: 6 (male). Asthenopodes picteti; HUBBARD, 1975: 111; HUBBARD, 1982a: 270; DOMÍNGUEZ, 1988a: 24 (male). Asthenopus picteti; HUBBARD & DOMÍNGUEZ, 1988: 207 (male).

Taxonomy: ULMER (1924) established the genus Asthenopodes, designating as typespecies Palingenia albicans PICTET which at that time was in the genus Asthenopus. However, the specimen described by PICTET (1843) was a misidentified specimen and not the species Palingenia albicans PERCHERON (1838) which is considered to belong to the genus Campsurus. For this reason, HUBBARD (1975) gave ULMER’s species the name Asthenopodes picteti. This species, known only from adults, can be distinguished by the following combination of characters: 1) Marginal intercalary veins of fore wings generally longer than distance between longitudinal veins (Fig. 212E); 2) vein IMP not joined at base to vein MP1 in fore wings (Fig. 212E); 3) fore wings of males with one cross vein between Rs and MA basal to fork of Rs (Fig. 212E); and 4) penes (Fig. 212G) slender and curved from base. Distribution: Uruguay, Brazil, and Guyana. SUBFAMILY CAMPSURINAE Taxonomy: This subfamily is composed by two specious genera: Campsurus and Tortopus. Adult Characteristics: 1) Pronotum about as wide as long, much broader posteriorly than in front; 2) fore legs of male about half as long as fore wings; 3) few cross veins in cubito-anal area of fore wings (Figs. 213A, C, 217A, D); 4) styliger plate reduced to a pair of parastyli (segments at base of forceps), small median lobe absent; 5) forceps 1segmented and thin (Figs. 213F-215K, 217C, J, K); 6) penes showing many varied forms;

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and 7) females of some species with a complementary structure to male parastyli, consisting of an excavated groove in VIII female sternum. Nymphal Characteristics: 1) Mandibular tusks relatively elongate and slender (Figs. 215N, 217N). Distribution: From Canada to Argentina. This subfamily reaches the most Southern and Northern distribution of the family, inhabiting not only in the tropics (where it thrives) but also in temperate areas. Genus Campsurus EATON Campsurus EATON, 1868b: 131; EATON, 1871: 56; TRAVER, 1947b: 371; TRAVER, 1950: 593. (Type-species: Palingenia latipennis WALKER, original designation).

Taxonomy: Campsurus, with 42 described species, constitutes the most specious genus from the Neotropics, but not the best known. Many species have been described from only one sex, but due to the marked sexual dimorphism, it is very difficult to associate the complementary sex without rearing or collecting them together in nuptial swarms (rarely observed in this group). There is not much agreement among authors on characters that can be used to discriminate species, and thus descriptions are highly heterogeneous. TRAVER (1947b) discussed the key and the characters given by NEEDHAM & MURPHY (1924) concluding that the division of species groups proposed by these authors was not well supported. At this time, any attempt to key all named species of the genus is futile. The taxonomy of the group is rather confused for several reasons: 1) many species are described only from females; 2) many descriptions are poor and lack illustrations; and 3) some types are lost. In the present section we give a brief description of the known although the only sure way to determine species of Campsurus is by comparison with the types. We strongly recommend that descriptions of new species in this genus be avoided until a major revision of the group is done. Nevertheless, we present below a key for Campsurus species groups, based on characteristics of male genitalia. These groups were delimited through the study of material collected throughout South America, from the study of type material and published illustrations. We recognize three groups of male genitalia that should not be regarded as formal (i.e. monophyletic) groups until a cladistic hypothesis for these taxa is proposed. These groups are (see characteristics in the key below): 1) segnis group, composed by: C. cuyuniensis, C. duplicatus, C. essequibo, C. indivisus, C. jorgenseni, C. latipennis, C. litaninensis, C. lucidus, C. pedicellarius, C. segnis, C. scutellaris and C. ulmeri. 2) albifilum group: C. albifilum, C. argentinus, C. brasiliensis, C. burmeisteri, C. evanidus, and C. major. Probably also C. juradinus and C. pallidus belong to this group, but original illustrations are too poor to be sure. 3) notatus group: C. assimilis, C. mahunkai, C. melanocephalus, C. meyeri, C. notatus, C. truncatus, and C. violaceus. Probably also C. longicauda belongs to this group. Other species not-assignable to a species-group: 1) Males known but descriptions and drawings inadequate (also generally the type material is lost): C. albicans, C. dallasi, C. holmbergii, C. nappii, C. paranensis, C. pfeifferi, C. quadridentatus, C. wappaei, and C. zikani. 2) Only known from female: C. claudus, C. corumbanus, C. dorsalis, C. mutilus, and C. striatus.

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Adult Characteristics: 1) Anal margin of hind wings not reticulated (Figs. 213B, D); 2) middle and hind legs blade-like (Fig. 213E), tibiae and tarsi absent; 2) forceps 1segmented, parastyli present (seeming a basal segment of forceps, Figs. 213F-M) and projected posteriorly on the inner or the outer corner, or not projected; 3) penes relatively complex, with 2 or more prominent projections of various forms (Figs. 213F-M). Nymphal Characteristics: 1) Mandibular tusks with prominent basal or sub-basal tubercle on median margin, and from a few to many apical crenulations (Fig. 215N); 2) numerous setae on outer-lateral margin of mandibles (Fig. 215N); 3) abdominal gill I bilobed (Fig. 215O). Distribution: From Argentina to Texas (USA). Biology: Nymphs burrow U-shaped tunnels in the bottom of lentic habitats (large rivers, lakes, ponds, swamps and dams). The flight activity of one species from Santa Fé (Argentina), probably C. violaceus, was observed in April 1998 (high water season). Around 18:30 h some male subimagos began a slow, hovering flight, at 3 m above riverbank. After a few minutes these males molted the subimaginal exuviae in flight, changing almost immediately to a much more rapid flight. Male imagos congregated in swarms of 50-100 individuals. Swarms occupied a spherical space of 1 m diameter, which maintained its form in spite of the wind. The entire swarm moved slowly but constantly up and down from 3 to 6-7 m. Each male performed a rapid horizontal zigzag (not vertically as is common for the order) of about 40 cm of amplitude. Female adults flew above the swarm and were grabbed by the males from behind. The flight activity lasted 30-40 minutes, ending around 19 h. Nevertheless, adults of both sexes continued arriving in large numbers to the lights until 21:30 h. KEY

TO

SOUTH AMERICAN

SPECIES GROUPS

Male imagos 1 – 2(1) –

CLAVE

Penes relatively simple, each lobe with one finger-like projection on a membranous subtriangular base (Figs. 214P-Q); parastyli posteriorly projected on inner (medial) corner (Figs. 214P-Q) ........................................................ notatus group Penes not as above, generally more complex; parastyli projected posteriorly on outer corner (Figs. 213N, 215G), or without projections (Figs. 213F, S) ............. 2 Parastyli stout (wide and short), without long projections paralleling the forceps (Figs. 213F, S) .............................................................................................. albifilum group Parastyli slender and posteriorly projected on outer corner paralleling the forceps or curving dorsally around penes (Figs. 213N, 215G) ..................................... segnis group PARA LOS GRUPOS DE ESPECIES SUDAMERICANOS

Imagos macho 1 –

Penes relativamente simples, cada lóbulo con una proyección digitiforme ubicada sobre una base membranosa subtriangular (Figs. 214P-Q); parastyli con una proyección posterior en el lado interno (medial) (Figs. 214P-Q) .......... grupo notatus Penes no como en el dilema anterior; parastyli proyectado posteriormente en el borde externo (Figs. 213N, 215G) o sin proyección (Figs. 213F, S) ....................... 2

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Plate 213. Campsurus, imagos. Figs. 213A-U: Campsurus cf. violaceus: 213A, male fore wing; 213B, male hind wing; 213C, female fore wing; 213D, female hind wing; 213E, male hind leg. Male genitalia (v.v. unless indicated otherwise): 213F, C. albifilum; 213G, C. argentinus, l.v.; 213H, same, v.v.; 213I, C. assimilis; 213J, same, l.v.; 213K, C. brasiliensis; 213L, C. burmeisteri, l.v.; 213M, same, v.v.; 213N, C. cuyuniensis; 213O, C. dallasi; 213P, C. duplicatus; 213Q, C. essequibo; 213R, same, detail of penes; 213ST, C. evanidus; 213U, C. indivisus. (PS = parastyli; FP = finger-like projections of penes; MB = membranous bases of penes). Fig. 213F modified from KIMMINS, 1960; 213G-H, L-M, U from ULMER, 1942; 213I-K from TRAVER, 1944; 213N, Q-S from TRAVER, 1947; 213O from NAVÁS, 1927a; 213P from SPIETH, 1943; and 213T from NEEDHAM & MURPHY, 1924.

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Parastilo robusto (ancho y corto), sin proyecciones notables paralelas al segmento del fórceps (Figs. 213F, S) ....................................................................... grupo albifilum Parastilo más delgado y proyectado posteriormente en el borde externo o curvándose dorsalmente alrededor de los penes (Figs. 213N, 215G) ............................ grupo segnis

Campsurus albicans (PERCHERON) Ephemera albicans PERCHERON in GUERIN & PERCHERON, 1838. Palingenia albicans; BURMEISTER 1839: 803; WALKER, 1853: 548 (female) [partim]. Campsurus albicans; EATON, 1883: 39 (male); LESTAGE, 1923: 120; LESTAGE, 1924c: 39; HUBBARD, 1982a: 270.

Taxonomy: This species was originally described in the genus Ephemera from one male imago, only indicating “Brazil” as the type locality. After being placed in Palingenia by BURMEISTER (1839), it was moved to Campsurus by EATON (1883). TRAVER (1947b) proposed to “maintain it just as a name”. This species is very poorly known and there are doubts as to its identity, but the following characteristics can be mentioned: 1) Wingspan 30 mm; 2) fore legs grayish violet; 3) wings whitish tinged toward base with violet gray, main veins violet gray; 4) yellowish species. Distribution: Brazil. Campsurus albifilum (WALKER) Palingenia albifilum WALKER, 1853: 554; HAGEN, 1861: 304. Campsurus albifilum; EATON, 1871: 57; EATON, 1883: 39 (male); LESTAGE, 1923: 120; NEEDHAM & MURPHY, 1924: 15; NAVÁS, 1926c: 110; TRAVER, 1947b: 383; KIMMINS, 1960: 312 (male); HUBBARD, 1982a: 270.

Taxonomy: This species, known only from male imagos, can be characterized as follows: 1) Fore wing length, 13 mm; 2) fore femora and fore tibiae sepia-brown, tarsi sepia-gray; 3) veins C, Sc and R1 of fore wings light brown ochreous at base, distally light sepia gray; 4) parastyli robust, apex curved outward (Fig. 213F); 5) each lobe of penes with one large and one smaller lobe (Fig. 213F). Distribution: Brazil (Pará, Nova Friburgo); French Guiana (Maroni); Argentina (Santa Fé). Campsurus argentinus ESBEN-PETERSEN Campsurus argentinus ESBEN-PETERSEN, 1912: 334 (male); ULMER, 1920c: 106; LESTAGE, 1923: 120; NEEDHAM & MURPHY, 1924: 16; ULMER, 1942: 117 (male); TRAVER, 1947b: 379; TRAVER, 1950: 593; HUBBARD, 1982a: 270.

Taxonomy: This species described from only one male imago, can be characterized as follows: 1) Fore wing length, 13 mm; 2) dorsum of head dark; 3) fore legs dark grayish violet; 4) wings slightly tinged with violet, fore wing veins Sc and R1 dark grayish violet at basal half, remaining veins yellowish gray; 5) parastyli robust, outer apical margin slightly projected posteriorly (Fig. 213H); 6) each lobe of penes with one large and one smaller lobe (Figs. 213G-H). Distribution: Argentina (Alto Paraná).

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Campsurus assimilis TRAVER Campsurus assimilis TRAVER, 1944: 39 (male, female); HUBBARD, 1982a: 270.

Taxonomy: This species described from imagos of both sexes can be characterized as follows: 1) Fore wing length: male, 15 mm; female, 21 mm; 2) dorsum of head purplish black, without reddish-purple marks on lateral ocelli; 3) fore femora light purplish brown, tibiae deep purplish gray; 4) fore wings with veins C, Sc and R1 purplish, stigmatic area grayish purple; 5) parastyli robust, inner apical margin slightly projected posteriorly (Fig. 213I); 6) lobes of penes finger-like and curved downward (Fig. 213J); 7) whitish species. Distribution: Brazil (Rio Grande do Sul). Campsurus brasiliensis TRAVER Campsurus brasiliensis TRAVER, 1944: 37 (male); HUBBARD, 1982a: 270.

Taxonomy: This species, known from male imagos, can be characterized by: 1) Fore wing length 15-16 mm; 2) area between eyes and median ocellus light reddish brown, with a V-shaped mark between the eyes, and with small purplish red spot on upper surface of each ocellus; 3) fore femora streaked with brown dorsally, tibiae tinged with light purplish brown; 4) fore wings veins C, Sc and R1 purplish, except near apex, remaining veins light yellowish; 5) male genitalia as in Fig. 213K; 6) whitish species. Distribution: Brazil (Rio Grande do Sul). Campsurus burmeisteri ULMER Campsurus burmeisteri ULMER, 1942: 122 (male); HUBBARD, 1982a: 270.

Taxonomy: This species was erected from a single male (ULMER, 1942), previously described as the male of Campsurus dorsalis (BURMEISTER) by ULMER (1921). TRAVER (1947b) suggested that this species may be a Tortopus (see discussion under C. dorsalis), which seems acceptable when looking ULMER’s drawings of the male genitalia (Figs. 213L-M). As nothing is said about the legs (fin-like in Campsurus) in the original description, study of type material is needed to define its generic assignment. Following the description given by ULMER (1942) we can extract one possible character to aid in the identification of the species (besides genitalia in Figs. 213L-M): fore wing veins Sc and R1 fused together before reaching the apex of the wing. Distribution: Brazil. Campsurus claudus NEEDHAM & MURPHY Campsurus claudus NEEDHAM & MURPHY, 1924: 23 (female); TRAVER, 1947b: 383; HUBBARD, 1982a: 270.

Taxonomy: This species known from two female adults, can be characterized by: 1) Fore wing length, 16 mm; 2) dorsum of head completely blackish; 3) fore legs blackish; 4) main veins tinged with purplish brown. Distribution: Brazil (Minas Gerais).

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Plate 214. Campsurus, imagos. Figs. 214A-S, male genitalia (v.v., unless counter-indicated): 214A, C. jorgenseni, l.v.; 214B, same, v.v.; 214C, C. juradinus; 214D, C. latipennis, penes and left forceps omitted; 214E, same, detail of penes; 214F, same, detail of penes and forceps base; 214G, C. litaninensis; 214H, C. longicauda; 214I-J, C. lucidus; 214K, C. mahunkai, apex of forceps omitted; 214L, C. major, penes folded; 214M, same, penes visible; 214N, C. melanocephalus; 214O, C. meyeri; 214P, C. notatus, forceps omitted; 214Q, same, penes folded back; 214R, C. pallidus, penes folded; 214S, same, penes visible. (FP = finger-like projections of penes; MB = membranous bases of penes; PS = parastyli). Figs. 214A-B modified from ULMER, 1942; 214C from NAVÁS, 1930; 214D-E from KIMMINS, 1960; 214F from EATON, 1883-1888; 214G from SPIETH, 1943; 214H from NAVÁS, 1931b; 214J, M, P-Q from TRAVER, 1947b; 214K from PUTHZ, 1973; 214L, R, S from NEEDHAM & MURPHY, 1924; 214N from PEREIRA & DA-SILVA, 1991; and 214O from NAVÁS, 1934a.

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Campsurus corumbanus NEEDHAM & MURPHY Campsurus corumbanus NEEDHAM & MURPHY, 1924: 21 (female); TRAVER, 1944: 43; TRAVER, 1947b: 382 (female); HUBBARD, 1982a: 270.

Taxonomy: This species was described only from female adults, showing great variation in size. In the original description, fore wing length varies from 14 mm to 19 mm, but TRAVER (1947b) remarked that fore wings of the holotype are just 10 mm long. For this reason TRAVER doubted that all the specimens belong to the same species. Campsurus corumbanus can be characterized as follows: 1) Dorsum of head blackish between eyes and median ocellus, lighter posteriorly; 2) fore legs purple; 3) fore wing veins C, Sc, and R1 (and cross veins between them) tinged with purplish. Distribution: Brazil (Corumbá, Pirapora). Campsurus cuyuniensis TRAVER Campsurus cuyuniensis TRAVER, 1947b: 387 (male, female); HUBBARD, 1982a: 270.

Taxonomy: This small species, described from imagos of both sexes, can be characterized as follows: 1) Fore wing length, 5.5 mm in male, 6-7 mm in female; 2) head reddish-brown on hind margin and with V-shaped area on vertex, blackish line between eyes; 3) fore tibia smoky brown, tarsi yellowish; 4) wings slightly tinged with yellowish, fore wings with veins Sc and R1 purplish brown on basal half, rest yellowish; 5) parastyli long, well developed, outer apical margin strongly projected posteriorly (Fig. 213N); 6) each lobe of penes with three finger-like processes (Fig. 213N); 6) yellowish species. Distribution: Guyana (Cuyuni river). Campsurus dallasi NAVÁS Campsurus dallasi NAVÁS, 1927a: 28 (male); TRAVER, 1947b: 371; HUBBARD, 1982a: 270.

Taxonomy: This species, very imperfectly known, is very similar to C. segnis according to NAVÁS (1927a). TRAVER (1947b) pointed out that the genitalia of the two species are different. NAVÁS drawings of the genitalia of C. dallasi are inadequate; nevertheless this species could be characterized by: 1) Fore wing length, 12.3 mm; 2) head ochraceous; 3) fore legs dark; 4) fore wing veins Sc and R1 purple; 5) genitalia poorly known from NAVÁS’ drawing (Fig. 213O). Distribution: Argentina. Campsurus dorsalis (BURMEISTER) Palingenia dorsalis BURMEISTER, 1839: 803; WALKER, 1853: 549; HAGEN, 1861: 304. Asthenopus dorsalis; EATON, 1871: 59 (female). Tortopus dorsalis; TRAVER, 1947b: 372. Campsurus dorsalis; EATON, 1883: 41 (female); HAGEN, 1888: 228; BANKS, 1913: 84 (male); NAVÁS, 1920d: 35; NAVÁS, 1920b: 414; NAVÁS, 1920e: 53; ULMER, 1920c: 106; ULMER, 1920b: 20; ULMER, 1921: 231 (male); LESTAGE, 1922b: 142; LESTAGE, 1923: 121; LESTAGE, 1924c: 39; ULMER, 1942: 119 (female); TRAVER, 1947b: 371; TRAVER, 1950: 593; HUBBARD, 1982a: 270.

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Taxonomy: This species was described from females that EATON (1883) placed in Campsurus. BANKS (1913) figured the genitalia from what he considered the male of this species, but without any certainty about its actual identity. ULMER (1921) described a male, also as C. dorsalis, which led TRAVER (1947b) to move this species to Tortopus. When ULMER (1942) erected a new species for this male (C. burmeisteri), TRAVER (1950) reconsidered this species as a Campsurus, stating that C. burmeisteri was the one that should be a Tortopus (but never transferred it formally). As a result, C. dorsalis is only known from females, that could be characterized as follows: 1) Wingspan 38 mm; 2) head blackish; 3) fore legs blackish gray; 4) membrane of wings almost hyaline, costal region of fore wings dark violet, remaining veins slightly darkened, hind wing veins whitish. Distribution: Brazil (Nova Friburgo, São Paulo); Paraguay (Asunción). Campsurus duplicatus SPIETH Campsurus duplicatus SPIETH, 1943: 7 (male); TRAVER, 1947b: 383; HUBBARD, 1982a: 270.

Taxonomy: This species, known from male adults, can be characterized by: 1) Fore wing length, 7-8 mm; 2) head piceous; 3) fore legs tinged with piceous; 4) fore wing veins S, Sc and R1, and hind wing veins C and Sc, tinged with piceous basally, lightening toward the apex, remaining veins translucent; 5) parastyli slender, outer apical margin projected posteriorly (Fig. 213P); 6) penes U-shaped, each lobe finger-like (Fig. 213P). Distribution: Brazil (Rio Negro), Guyana (Bartica). Campsurus essequibo TRAVER Campsurus essequibo TRAVER, 1947b: 388; HUBBARD, 1982a: 270.

Taxonomy: This species is known from male adults and can be characterized as follows: 1) Fore wing length, 6 mm; 2) head dark reddish brown; 3) fore legs blackish brown, tarsi lighter; 4) fore wing veins C, Sc and R1 purplish brown on basal 2/3, rest yellowish; 5) parastyli long, well developed, outer apical margin strongly projected posteriorly (Fig. 213Q); 6) each lobe of penes with three finger-like processes (Fig. 213R); 6) reddish-brown species. This species shows close relationships with C. cuyuniensis. Distribution: Guyana (Cuyuni river). Campsurus evanidus NEEDHAM & MURPHY Campsurus evanidus NEEDHAM & MURPHY, 1924: 18 (male); TRAVER, 1947b: 379 (male); HUBBARD, 1982a: 270.

Taxonomy: This species known from adults of both sexes can be characterized by: 1) Fore wing length of male, 8-10 mm; female, 10-12 mm; 2) head yellowish except around eyes and ocelli, blackish; 3) fore legs yellowish, slightly tinged with purplish; 4) parastyli robust, outer apical margin directed outward (Fig. 213S); 5) each lobe of penes with two finger-like processes (Figs. 213S-T); 6) light yellowish species. No mention is made in the original description regarding the coloration of the wing veins, but we have studied the holotype. Veins C, Sc, and R1 are purplish brown basally and whitish yellow distally, with the remaining veins whitish. Distribution: Brazil (Minas Gerais).

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Plate 215. Campsurus. Figs. 215A-M, male genitalia (v.v., unless counter-indicated): 215A, C. paranensis; 215B, C. pedicellarius; 215C, C. pfeifferi; 215D, C. quadridentatus, with details; 215E-F, C. scutellaris; 215G, C. segnis; 215H, C. truncatus; 215I-J, C. ulmeri; 215K-L, C. violaceus; 215M, C. zikani. Figs. 215N-O, nymph: 215N, left mandible, d.v.; 215O, abdominal gill I. Fig. 215A modified from NAVÁS, 1932; 215B from SPIETH, 1943; 215C from NAVÁS, 1931; 215D from EATON, 1883; 215E, G, I-L from TRAVER, 1947b; 215F from NEEDHAM & MURPHY, 1924; 215H from PUTHZ, 1973; and 215M from NAVÁS, 1934a.

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Campsurus indivisus ULMER Campsurus indivisus ULMER, 1942: 16 (male, female); HUBBARD, 1982a: 271.

Taxonomy: This species, known from male and female adults, can be characterized as follows: 1) Head brownish yellow; 2) fore wing length, 7.5 mm; 3) basal half of fore wing veins C, Sc and R1 shaded with grayish violet, remaining veins not pigmented; and 4) genitalia as in Fig. 213U. Distribution: Brazil (Santa Rita). Campsurus jorgenseni ESBEN-PETERSEN Campsurus jorgenseni ESBEN-PETERSEN, 1912: 333 (male, female); ULMER, 1920c: 116; LESTAGE, 1923: 122; ULMER, 1942: 114 (male, female); TRAVER, 1950: 593; HUBBARD, 1982a: 271. Campsurus jorgensini [sic]; NEEDHAM & MURPHY, 1924: 16.

Taxonomy: This species, known from male and female adults, can be characterized as follows: 1) Length of fore wing: male, 10 mm; female, 11 mm; 2) head dark anteriorly, posteriorly whitish yellow with a pair of grayish purple marks; 3) fore legs whitish tinged with grayish purple; 4) wings translucent, fore wing veins C, Sc and R1 grayish purple on basal half, remaining veins translucent; 5) parastyli with outer apical margin projected posteriorly (Figs. 214A-B); 6) forceps shorter than penes (Fig. 214B); 7) each lobe of penes with two finger-like processes (Figs. 214A-B). The male genitalia of this species is very similar to that of C. scutellaris, treated below. Distribution: Argentina (Alto Paraná). Campsurus juradinus NAVÁS Campsurus juradinus NAVÁS, 1930e: 129 (male); TRAVER, 1947b: 371; HUBBARD, 1982a: 271.

Taxonomy: Original description and drawings are poor, but this species could be characterized by: 1) Fore wing length, 10 mm; 2) head dorsally ochraceous, posteriorly blackish; 3) fore legs whitish yellow, tibia dark-streaked; 3) wings hyaline with costal and subcostal areas tinged with purple basally; 5) penes divided in two lobes (Fig. 214C). The typus is not present where NAVÁS said he deposited it (Museo Nacional de Buenos Aires). Distribution: Argentina (Corrientes). Campsurus latipennis (WALKER) Palingenia latipennis WALKER, 1853: 554; HAGEN, 1861: 304. Campsurus latipennis; EATON, 1871: 57 (male, female); EATON, 1873: 388; EATON, 1883-1888: 39 (male, female); BANKS, 1913: 84 (male); LESTAGE, 1923: 122; NEEDHAM & MURPHY, 1924: 15; TRAVER, 1947b: 383 (male); KIMMINS, 1960: 313 (male); HUBBARD, 1982a: 271.

Taxonomy: Campsurus latipennis, the type species of the genus, was described by WALKER, redescribed by EATON (1871, 1883), and treated by several other authors (BANKS, 1913; TRAVER, 1947b) who had doubts about its identity. For this reason we extract the following characterization from EATON, who revised the types: 1) Fore wing

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length, 8 mm; 2) fore legs sepia brown, claws whitish; 3) basal part of fore wing veins C, Sc and R1 sepia gray; 4) parastyli (first forceps “segment”) with outer apical margin projected posteriorly (Figs. 214D, F); 5) penes complex, each lobe with three finger-like processes (Figs. 214E-F). TRAVER (1950) suggested a close relationship of this species with C. cuyuniensis and C. essequibo. Distribution: Brazil (Pará, Santarém). Campsurus litaninensis SPIETH Campsurus litaninensis SPIETH, 1943: 5 (male, female); HUBBARD, 1982a: 271.

Taxonomy: This species, known from male and female imagos, can be characterized as follows: 1) Fore wing length: male, 9 mm; female, 9-10 mm; 2) head blackish; 3) fore legs blackish translucent, lightening toward apex; 4) fore wing veins C, Sc and R1 basally blackish, lightening distally; 5) parastyli with outer apical margin projected posteriorly (Fig. 214G); 6) styliger plate strongly projected medially (Fig. 214G); 7) each lobe of penes simple, finger-like (Fig. 214G); 8) yellowish species. Distribution: Surinam (Litani river). Campsurus longicauda NAVÁS Campsurus longicauda NAVÁS, 1931a: 455 (male); TRAVER, 1944: 43; TRAVER, 1947b: 371; HUBBARD, 1982a: 271. Compsurus longicauda; PETERSEN & GAEDIKE, 1968: 961.

Taxonomy: This species, known from one male adult, could be characterized as follows: 1) Fore wing length 14.5 mm; 2) head dark blackish; 3) fore wings hyaline, veins C, Sc and R1 purplish; 4) genitalia poorly known, penes apparently V-shaped (Fig. 214H). Distribution: Brazil (Ypiranga). Campsurus lucidus NEEDHAM & MURPHY Campsurus lucidus NEEDHAM & MURPHY, 1924: 16 (male); ULMER, 1942: 122 (male); TRAVER, 1947b: 376 (male); HUBBARD, 1982a: 271.

Taxonomy: This species was originally described from a single male (NEEDHAM & MURPHY, 1924), but TRAVER (1947b) added one additional male from the same date and locality of the holotype (Peru, Putumayo), and a few other male imagos from British Guiana. This author suggested that C. lucidus and C. pedicellarius should be synonymized. This species can be characterized by: 1) Fore wing length, 7 mm; 2) head blackish except line on posterior part; 3) fore legs pale, tinged with purplish brown on apex of tibia and body of tarsal segments; 4) basal part of fore wings and of veins C, Sc and R1 tinged with purple; 5) parastyli with outer apical margin projected posteriorly (Figs. 214I-J); 6) each lobe of penes with small apical rounded lobe and two small processes (Fig. 214J). Distribution: Brazil (Santa Catarina); Colombia; Guyana (Bartica); Peru (Putumayo river).

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Plate 216. Campsurus sp. Fig. 216A: nymphal habitus. Drawing by A. Dupuy.

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Campsurus mahunkai PUTHZ Campsurus mahunkai PUTHZ, 1973: 94 (male, female, egg); HUBBARD, 1982a: 271.

Taxonomy: This species, known from adults of both sexes, can be characterized as follows: 1) Fore wing length of male 10-12 mm; 2) head blackish with a Y-shaped blackish mark; 3) fore legs dark brown, tinged with purple; 4) fore wing whitish translucent, with costal and subcostal areas tinged with purple, veins of anterior half purplish brown, but whitish yellow on posterior half; 5) parastyli robust, inner and outer apical margins projected posteriorly (inner more pronounced) (Fig. 214K); 6) each lobe of penes fingerlike and curved downward (Fig. 214K); 7) yellowish brown species. Distribution: Bolivia (La Paz). Campsurus major NEEDHAM & MURPHY Campsurus major NEEDHAM & MURPHY, 1924: 17 (male); TRAVER, 1944: 36; TRAVER, 1947b: 378 (male); HUBBARD, 1982a: 271.

Taxonomy: This species was described by NEEDHAM & MURPHY from a single male, but TRAVER (1947b) redrew the genitalia and corrected the wing size using material from the same locality and date, although this material was not listed by the original authors. Apparently the holotype is in bad condition and the 20 mm length of the wings given originally is a mistake. Male adults of this species can be characterized as follows: 1) Fore wing length, 16 mm; 2) head yellowish; 3) fore legs yellowish with light shading on tibia; 4) fore wings light-colored, except costal area tinged with purplish gray; 5) parastyli robust, outer apical margin slightly projected posteriorly (Figs. 214L-M); 6) each lobe of penes with a large and a small rounded lobe (Fig. 214M); 6) light yellowish species. Distribution: Argentina (Buenos Aires). Campsurus melanocephalus PEREIRA & DA-SILVA Campsurus melanocephalus PEREIRA & DA-SILVA, 1991: 322

Taxonomy: This species, known from adults of both sexes and nymphs, can be characterized by: 1) Fore wing length: male, 13-15 mm; female, 20-23 mm; 2) head blackish; 3) fore legs with longitudinal brownish stripes; 4) fore wings vein R1 brownish, remaining veins translucent; 5) parastyli robust, outer and inner apical margin projected posteriorly (projection on inner margin more pronounced) (Fig. 214N); 6) each lobe of penes single, finger-like (Fig. 214N); 6) light brownish species. Distribution: Brazil (Rio de Janeiro). Biology: PEREIRA & DA-SILVA (1991) reported huge swarms for this species occurring during most of the year. Males performed the nuptial flight just after sundown. Eggs take 10-30 days to hatch in laboratory. Nymphs were found to dig tunnels in fine shifting sediment, at shallow depositional zones in a large lowland river.

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Campsurus meyeri NAVÁS Campsurus meyeri NAVÁS, 1934: 24 (male); TRAVER, 1947b: 371; HUBBARD, 1982a: 271.

Taxonomy: This species was described from a single male that is related to C. notatus according to NAVÁS (1934). TRAVER (1947b) could not corroborate this statement from the poor original drawing. Campsurus meyeri could be characterized as follows: 1) Fore wing length, 12 mm; 2) head dark; 3) fore wing hyaline, with costal area slightly tinged with yellowish pink, darkened at base of C and along Sc and R1; 4) parastyli robust, inner apical margin projected posteriorly (Fig. 214O); 5) each lobe of penes with a finger-like lobe and a triangular lobe (Fig. 214O). Distribution: Argentina (Chaco). Campsurus mutilus NEEDHAM & MURPHY Campsurus mutilus NEEDHAM & MURPHY, 1924: 22 (female); TRAVER, 1947b: 383 (female); HUBBARD, 1982a: 271.

Taxonomy: This species known only from female adults, can be characterized by: 1) Fore wing length, 15-16 mm; 2) head dorsally dark blackish purple; 3) fore legs purplish; 4) costal area basally dark; 5) yellowish species. Distribution: Brazil (from Rio Solimões to mouth of Rio Içá). Campsurus notatus NEEDHAM & MURPHY Campsurus notatus NEEDHAM & MURPHY, 1924: 20 (male); NAVÁS, 1926c: 109; TRAVER, 1944: 39; TRAVER, 1947b: 382 (male); DEMOULIN, 1955a: 30 (male, female); IRMLER, 1975: 348; HUBBARD, 1982a: 271.

Taxonomy: This species was described from male imagos (holotype and three other males, treated as paratypes by TRAVER, 1947b). In 1955a, DEMOULIN described females that show coloration similar to the males but slightly darker. TRAVER (1947b) remarked that from the four male adults types: two were large (fore wing length, 10-11 mm) and two small (8 mm). Also she commented on the similarity between the male genitalia of this species and C. violaceus, but still considering them as distinct. This species can be characterized by: 1) Fore wing length, 8-11 mm; 2) head dorsally blackish, with a brownish line crossing the occiput; 3) hind legs purplish brown; 4) fore wings translucent, tinged with purplish along veins C, Sc and R1; 5) parastyli robust, inner apical margin strongly projected posteriorly (Figs. 214P-Q); 6) each lobe of penes simple, finger-like (Figs. 214P-Q); 6) purplish-brown designs along the entire body. Distribution and Biology: Brazil (Paraguay river, between Puerto Esperança and Corumbá); Argentina (Santiago del Estero); Paraguay. LEAL & DE ASSIS ESTEVES (2000) studied the life cycle and production of this species in an Amazonian lake (Lake Batata, Para, Brazil) impacted by bauxite tailings effluents, and found that it was adapted to the conditions imposed by the bauxite tailings. The nymphs (not described formally) had a 105-day lifespan, with individuals between 2 mm and 17 mm in length. The estimated secondary production (dry weight) was 2.5 gm-²y-1. These authors, in another paper (LEAL et al., 2003) reported that the nymphs are important bioturbators (digging vertical tunnels almost 10 cm long) greatly influencing the ammonium and oxygen movement in the water-substrate interface.

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Campsurus pallidus NEEDHAM & MURPHY Campsurus pallidus NEEDHAM & MURPHY, 1924: 20 (male); TRAVER, 1947b: 380 (male); HUBBARD, 1982a: 271.

Taxonomy: This species is known from a single male and can be characterized by: 1) Fore wing length, 12 mm; 2) head light yellow, slightly tinged with purplish brown on dorsum; 3) fore legs faint purplish; 4) fore wing slightly tinged with purplish on basal costal half; 5) parastyli robust (Figs. 214R-S); 6) each lobe of penes with two finger-like processes (Fig. 214S). TRAVER (1947b) reported that the penes are no longer visible, probably due to movement of the mounted genitalia after the NEEDHAM & MURPHY drawings were made; 6) pale yellow species. Distribution: Argentina-Paraguay border (Alto Paraná). Campsurus paranensis NAVÁS Campsurus paranensis NAVÁS, 1932a: 111 (male); HUBBARD, 1982a: 271.

Taxonomy: This species is known from a single male and can be characterized as follows: 1) Fore wing length, 12 mm; 2) head blackish brown; 3) fore legs reddish brown; 4) fore wings hyaline, costal base purplish brown, stigmatic area whitish, Sc and R1 light purple; 5) yellowish brown species. Male genitalia were not described and the illustration is too poor to aid in its characterization (Fig. 215A). Distribution: Argentina (Santa Fé). Campsurus paraquarius NAVÁS Campsurus paraguarius [lapsus] NAVÁS, 1920e: 53 (male, female); LESTAGE, 1923: 122; TRAVER, 1947b: 371; HUBBARD, 1982a: 271. Campsurus paraquarius; NAVÁS, 1924a: 359.

Taxonomy: LESTAGE (1923) doubted the generic assignment of this species, known from one male and one female adults. According to LESTAGE, the anal area of the fore wings and the pronotal structure, were more typical of the genus Asthenopus. When TRAVER (1947b) revised the genus Campsurus, she proposed that this species be maintained as a name until more was known. This species could be characterized by: 1) Fore wing length of male, 6.0 mm, of female, 4.6 mm; 2) head blackish; 3) fore legs whitish, femora tinged with purplish brown; 4) fore wings hyaline with veins tinged with purplish brown; 5) whitish yellow species. The male genitalia were not illustrated, and the description is poor. Distribution: Paraguay (Paraguay river). Campsurus pedicellarius SPIETH Campsurus pedicellarius SPIETH, 1943: 7 (male, female); TRAVER, 1947b: 386; HUBBARD, 1982a: 271.

Taxonomy: This species is known from imagos of both sexes. According to SPIETH (1943) the wide and flattened penes, and the styliger plate, were unique characters, but TRAVER (1947b) stated that it was very similar to or synonymous with C. lucidus. She

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considered that the only difference in the male genitalia of both species (a profound incision in the styliger plate, between the base of the forceps) could be an artifact. This species can be characterized by: 1) Fore wing length of male, 8 mm, of female, 16 mm; 2) head pale, area between ocelli piceous; 3) fore legs shaded with piceous; 4) fore wing venation pale, except for Sc and basal part of C and R1; 5) parastyli with outer apical margin projected posteriorly (Fig. 215B); 6) each lobe of penes with a large and a small rounded lobe (Fig. 215B); 7) styliger plate with a deep median incision (Fig. 215B). Distribution: Guyana, Surinam. Campsurus pfeifferi NAVÁS Compsurus [sic] pfeifferi NAVÁS, 1931b: 457 (male); PETERSEN & GAEDIKE, 1968: 961. Campsurus pfeifferi; TRAVER, 1947b: 371; HUBBARD, 1982a: 271.

Taxonomy: This species, known only from male imagos, can be characterized by: 1) Fore wing length, 12.5 mm; 2) head yellowish brown; 3) fore legs yellowish brown, tibia and tarsus grayish; 4) fore wings hyaline, Sc purplish, C and R1 lighter; 5) male genitalia poorly illustrated (Fig. 215C); 6) yellowish brown species, abdomen lighter. Distribution: Argentina (Chaco). Campsurus quadridentatus EATON Campsurus quadridentatus EATON, 1871: 58 (male); EATON, 1883: 40 (male); ULMER, 1920c: 106; ULMER, 1921: 240; LESTAGE, 1923: 123; TRAVER, 1947b: 372; TRAVER, 1950: 593; KIMMINS, 1960: 314; HUBBARD, 1982a: 271.

Taxonomy: This species, briefly described and known only from male adults, could be characterized by: 1) Fore wing length, 13 mm; 2) legs yellowish-brown; 3) fore wings whitish-translucent, veins Sc and R1 blackish gray; 4) male genitalia poorly illustrated, but EATON (1883: 40) described it as follows: “styliger plate angularly excised behind on both sides of a large deep central subcircular sinus; adjacent to these excisions, tooth-like projections are present” (Fig. 215D); 5) light yellowish brown species. Distribution: Argentina (Santiago del Estero); Brazil (Santarém); Peru. Campsurus scutellaris NEEDHAM & MURPHY Campsurus scutellaris NEEDHAM & MURPHY, 1924: 19 (male); TRAVER, 1947b: 380 (male); HUBBARD, 1982a: 271.

Taxonomy: This species was described from a single male adult, that can be characterized by: 1) Fore wing length, 7 mm; 2) head blackish between eyes and ocelli, but purplish posteriorly; 3) fore legs yellowish with purplish mark on coxa and trochanter; 4) wings whitish translucent, tinged slightly on costal area; 5) parastyli with outer apical margin projected posteriorly (Figs. 215E-F); 6) forceps short; 7) each lobe of penes with two finger-like processes (Figs. 215E-F); 6) light brown species. Male genitalia of this species are very similar to that of C. jorgenseni. Distribution: Argentina (Misiones).

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Campsurus segnis NEEDHAM & MURPHY Campsurus segnis NEEDHAM & MURPHY, 1924: 19 (male, female); MORGAN, 1929: 61 (male, female); SPIETH, 1943: 5; TRAVER, 1947b: 380 (male); HUBBARD, 1982a: 271.

Taxonomy: This species was described from imagos of both sexes from British Guiana and Brazil. Later, MORGAN (1929) redescribed the species with material from British Guiana, but from larger individuals (body length, 10 mm) than those described by NEEDHAM & MURPHY. TRAVER (1947b) studied this species and discussed its unique male genitalia. Nevertheless, she recommended care with the assignment of individuals to C. segnis, because C. ulmeri has similar genitalia. She separated both species by the smaller size and darker coloration of C. segnis. Campsurus segnis can be characterized by: 1) Fore wing length: male, 7 mm, female, 8 mm; 2) head reddish brown; 3) fore legs pale, tarsi blackish; 4) basal portion of veins C, Sc and R1 brownish; 5) parastyli with a long outer projection on apical margin (Fig. 215G); 6) each lobe of penes with two long finger-like processes (Fig. 215G); 7) brownish species, abdomen reddish brown. Females are slightly darker than males. Distribution: Brazil, British Guiana, and Surinam. Campsurus striatus NEEDHAM & MURPHY Campsurus striatus NEEDHAM & MURPHY, 1924: 22 (female); TRAVER, 1947b: 383; HUBBARD, 1982a: 271.

Taxonomy: This species is known from a single female adult that can be characterized as follows: 1) Fore wing length, 13 mm; 2) head dorsally blackish; 3) fore legs reddish; 4) fore wings translucent with main veins reddish; 5) reddish brown species, with strong blackish marks present on abdomen, specially on posterior segments, giving an annulated appearance. Distribution: Brazil. Campsurus truncatus ULMER Campsurus truncatus ULMER, 1920a: 2 (male); LESTAGE, 1923: 123; PUTHZ, 1973: 94 (male); HUBBARD, 1982a: 271.

Taxonomy: This species, known only from male imagos, can be characterized by: 1) Fore wing length, 13-14 mm; 2) head dorsally reddish brown, blackish between ocelli; 3) fore wings veins light brownish-gray, except veins C, Sc, and R1 dark brownish violet; 4) parastyli robust, outer and inner apical margin projected posteriorly (inner projected more pronouncedly) (Fig. 215H); 5) each lobe of penes single, finger-like (Fig. 215H). Distribution: Brazil (Espirito Santo), Bolivia (Santa Cruz, Sara prov.). Campsurus ulmeri TRAVER Campsurus sp. “Dr. Ulmer’s unnamed species from Brazil”: TRAVER, 1947:376, 381. Campsurus ulmeri TRAVER, 1950: 594; HUBBARD, 1982a: 271.

Taxonomy: This species, known from male imagos, can be characterized by: 1) Fore wing length, 11 mm; 2) head strongly shaded with black; 3) fore legs yellowish, with

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blackish lines at base of femur and tibia, femoro-tibial joint blackish, grayish tarsi; 4) membrane of wings hyaline, with C, Sc and R1 purplish gray, remaining veins hyaline; 5) parastyli with a long outer projection on apical margin (Figs. 215I-J); 6) each lobe of penes with two long finger-like processes (Figs. 215I-J); 7) reddish brown species, abdomen whitish yellow. In addition to the males, TRAVER indicated the presence of 5 small and 5 larger females. She supposed that the large females corresponded to this species, but stated that possibly NEEDHAM & MURPHY had described them in another species. TRAVER commented on the similarity between C. ulmeri and C. segnis; nevertheless she distinguished C. ulmeri by its larger size and paler coloration. Distribution: Brazil (Santa Catarina). Campsurus violaceus NEEDHAM & MURPHY Campsurus violaceus NEEDHAM & MURPHY, 1924: 18 (male, female); TRAVER, 1947b: 379 (male); HUBBARD, 1982a: 271.

Taxonomy: This species is known from adults of both sexes. TRAVER (1947b) redrew the male genitalia, and corrected the wing length given in the original description. Also she designated as paratypes the rest of the material used by NEEDHAM & MURPHY. Adults of this species can be characterized by: 1) Fore wing length: male, 11 mm (10 mm in original description), female, 13 mm (originally 14 mm); 2) head blackish, tinged with violet near margins; 3) fore legs light purple; 4) fore wings with costal area strongly shaded with purplish; 5) parastyli stout, with apical outer margin strongly projected (Figs. 215K-L); 6) each lobe of penes single, finger-like (Figs. 215K-L); 7) purplish brown species, abdomen reddish brown. Distribution and Biology: Argentina, Brazil (Paraná). This species has been used for studies on nymphal density (MELO et al., 1993) and seasonal dynamics and secondary production (TAKEDA & GRZYBKOWSKA, 1997). Campsurus zikani NAVÁS Campsurus zikani NAVÁS, 1934a: 25; TRAVER, 1947b: 371; HUBBARD, 1982a: 271.

Taxonomy: This species, insufficiently known from male and female adults, could be characterized by: 1) Fore wing length: male, 14 mm, female, 20.5 mm; 2) head blackish; 3) membrane of wings slightly yellowish, veins Sc and R1 purplish red; 4) yellowish species, darker on head and thorax. The illustration of male genitalia is too poor to be useful (Fig. 215M). Distribution: Brazil (Rio de Janeiro). Campsurus holmbergii (WEYENBERGH) Nomen Dubium Ephemera holmbergii WEYENBERGH, 1883: 160. Ephemera holmbergi; HAGEN, 1888: 230. Campsurus holmbergi; NAVÁS, 1915a: 119; NAVÁS, 1919: 80; LESTAGE, 1923: 122; NAVÁS, 1924a: 359; NEEDHAM & MURPHY, 1924: 15; TRAVER, 1947b: 371; BERTHÉLEMY, 1965: 2. Campsurus holmbergii; EATON, 1883-1888: 303; ULMER, 1920c: 106; LESTAGE, 1924c: 58; HUBBARD, 1982a: 270.

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Taxonomy: TRAVER (1947b) stated that this species should be maintained only as a name. We prefer to treat this species as a nomen dubium because the type material is lost and identification of the species by means of the description is not possible. Distribution: Argentina (Santiago del Estero, Buenos Aires). Campsurus nappii (WEYENBERGH) Nomen Dubium Palingenia nappii WEYENBERGH, 1883: 162. Campsurus nappii; EATON, 1887: 304; HAGEN, 1888: 230; ULMER, 1920c: 106; LESTAGE, 1923: 122; LESTAGE, 1924c: 58; TRAVER, 1947b: 371; HUBBARD, 1982a: 271.

Taxonomy: TRAVER (1947b) stated that the species should be maintained only as a name. We here propose to treat it as a nomen dubium because the type material is lost and the descriptions are useless for identification. Distribution: Argentina (Santiago del Estero). Campsurus wappaei (WEYENBERGH) Nomen Dubium Ephemera wappaei WEYENBERGH, 1883: 159. Campsurus wappaei; EATON, 1883-1888: 303 (male, female); HAGEN, 1888: 228 (female); ULMER, 1920c: 106; LESTAGE, 1923: 123; LESTAGE, 1924c: 58; TRAVER, 1944: 43; TRAVER, 1947b: 371; HUBBARD, 1982a: 271.

Taxonomy: According to TRAVER (1947b) this species “should be maintained only as a name”. We here propose to treat it as a nomen dubium because the type material is lost and the descriptions useless to identification. Distribution: Argentina (Córdoba). Genus Tortopus NEEDHAM & MURPHY Tortopus NEEDHAM & MURPHY, 1924: 23; ULMER, 1933: 197; TRAVER, 1950: 596; MCCAFFERTY, 1975: 489. (Type-species: Tortopus igaranus NEEDHAM & MURPHY, original designation).

Taxonomy: The genus Tortopus was established by NEEDHAM & MURPHY based on a single species known from females. Presently, it contains eleven species, of which eight are South American. Tortopus exhibits a marked sexual dimorphism and only a very few of the species are known from both sexes. As yet, no South American nymphs have been described. Adult Characteristics: 1) Presence of reticulated veins in anal margin of hind wings (Figs. 217B, E); 2) middle and hind legs with all segments present, although atrophied and non-functional (Fig. 217M); 3) small basal segment (not totally divided) present between forceps and parastyli (Figs. 217C, I, K-L); 4) penes relatively simple, U or V-shaped (Figs. 217C, I-K); and 5) females with complementary structure to male parastyli, consisting of an excavated groove in female sternum VIII (also present in some species of Campsurus). Nymphal Characteristics (Figs. 233D-E): 1) Mandibular tusks with single prominent subdistal tubercle on median margin (arrow in Fig. 217N); 2) few long setae on outer

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lateral margin of mandibles (Fig. 217N); 3) abdominal gill I simple (Fig. 217O). These nymphs are very similar to those of Campsurus, so care must be applied when using keys. Distribution: From Argentina to Canada. Biology: Nymphs of this genus burrow U-shaped tunnels (Fig. 233C) in exposed clay banks of rivers and streams. Nymphs are filter feeders of organic particulate matter. Adults reared from nymphs collected in NW Argentina emerged in laboratory during March and April. When ready to emerge, nymphs swim to the surface where the cuticle is shed and the subimago take flight immediately. Subimaginal skin from the dorsal and the ventral sides of wings are shed independently, the extremely thin cuticle disintegrates in small to large fragments, never impeding the flight. The thicker cuticle of the body is shed also in the air, the exuvia is pushed backwards (apparently by abdominal undulations) until reaching the apex of caudal filaments, from where it falls. KEYS

TO

Males 1 – 2(1) –

SOUTH AMERICAN TORTOPUS

43

Penes long and slender, widely separated from base; parastyli well developed (Figs. 217K-L) ................................................................................................ Tortopus unguiculatus Penes relatively stout, fused at base; parastyli short or very reduced (Figs. 217C, I-J) ................................................................................................................................................ 2 Parastyli short but visible (Fig. 217C); fore wing veins C, Sc and R1 yellowish at base, lighter toward apex, remaining veins whitish ............................... Tortopus zottai Parastyli greatly reduced (Figs. 217I-J); basal part of veins C, Sc, and R1 weakly purple ............................................................................................................. Tortopus harrisi

Females 1 – 2(1) – 3(1) – 4(3)

43

Small species, length of fore wings 9 to 12 mm ......................................................... 2 Large species, length of fore wings 17 to 21 mm ...................................................... 3 Wings with veins purple; abdomen dark brown, with pale median line .................. ...................................................................................................................... Tortopus igaranus Wings with veins C and Sc violet at base, clearer near apex, remaining veins whitish; abdomen light yellowish, tinted with blackish violet laterally ..................... ................................................................................................................ Tortopus unguiculatus Wing veins brown; abdominal terga whitish shaded with brownish gray, color pattern as in Fig. 217G .................................................................... Tortopus obscuripennis Wings with some veins yellowish gray or yellowish white; abdominal color pattern not as above ........................................................................................................................ 4 Eggs filling the abdomen orangish; legs II and III brownish yellow; abdominal color pattern as in Fig. 217F .............................................................. Tortopus bruchianus Eggs filling the abdomen yellowish; legs II and III whitish; abdominal color pattern as in Fig. 217H ................................................................................ Tortopus sarae

The species T. parishi (BANKS) is not included because of lack of data.

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CLAVES

PARA LOS

Machos 1 – 2(1) –

TORTOPUS

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SUDAMERICANOS

44

Penes largos y delgados, ampliamente separados desde la base; parastyli bien desarrollados (Figs. 217K-L) ........................................................... Tortopus unguiculatus Penes relativamente más robustos, unidos en la base; parastyli cortos o muy reducidos (Figs. 217C, I-J) ................................................................................................ 2 Parastyli cortos pero notorios (Fig. 217C); alas anteriores con las venas C, Sc y R1 amarillentas en la base, aclarándose hacia el ápice, restantes venas blanquecinas . .......................................................................................................................... Tortopus zottai Parastyli muy reducidos (Figs. 217I-J); parte basal de las venas C, Sc y R1 púrpura suave ............................................................................................................... Tortopus harrisi

Hembras 1 – 2(1) – 3(1) – 4(3) –

Especies pequeñas, alas anteriores entre 9 y 12 mm de longitud ............................ 2 Especies grandes, alas anteriores entre 17 y 21 mm de longitud ............................ 3 Alas con todas las venas purpúreas; abdomen castaño-oscuro con la línea medial más pálida .................................................................................................. Tortopus igaranus Alas con las venas C y Sc violáceas en la base, aclarándose hacia el ápice, restantes venas blanquecinas; abdomen amarillento claro, teñido con violeta negruzco lateralmente ......................................................................................... Tortopus unguiculatus Alas con todas las venas castañas; tergitos abdominales blanquecinos, sombreados de castaño-grisáceo con un patrón de coloración como el de la Fig. 217G ........... ............................................................................................................... Tortopus obscuripennis Alas con parte de las venas amarillo-grisáceas o amarillo-blanquecinas; patrón de coloración abdominal no como el arriba mencionado (Figs. 217F, H) .................. 4 Huevos que llenan el abdomen de color anaranjado; patas II y III castañoanaranjadas; patrón de coloración abdominal como en la Fig. 217F ....................... .................................................................................................................. Tortopus bruchianus Huevos que llenan el abdomen de color amarillento; patas II y III blanquecinas; patrón de coloración abdominal como en la Fig. 217H ....................... Tortopus sarae

Tortopus bruchianus (NAVÁS) Campsurus bruchianus NAVÁS, 1926c: 110 (female); TRAVER, 1944: 43; HUBBARD, 1982a: 270; DOMÍNGUEZ, 1989b: 273. Tortopus bruchianus; DOMÍNGUEZ, 1987b: 9 (female).

Taxonomy: Tortopus bruchianus (NAVÁS) is known only from female adults. It was described by NAVÁS (1926c) based on a series of females from which DOMÍNGUEZ (1987b), when transferring the species to Tortopus, designated a lectotype. The females of this species can be separated from those known from other species of Tortopus by the following combination of characters: 1) Wing veins C, Sc, and R1 brownish violet, clearer near apex, 44

No se incluye la especie T. parishi (BANKS) debido a los pocos datos disponibles.

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C A B

PS

D E F

G

I

PS

H

J

K

L

PS

N

PS

M

O

Plate 217. Tortopus, imagos. Figs. 217A-O. T. zottai: 217A, male fore wing; 217B, male hind wing; 217C, male genitalia, v.v. T. obscuripennis: 217D, female fore wing; 217E, female hind wing; 217G, female abdominal color pattern. T. bruchianus: 217F, female abdominal color pattern. T. sarae: 217H, female abdominal color pattern. T. harrisi: 217I, detail of penes and forceps bases; 217J, male genitalia, v.v. T. unguiculatus: 217K, male genitalia, v.v.; 217L, same, l.v. Tortopus sp.: 217M, male imaginal hind leg. Nymph, Tortopus sp.: 217N, left mandible, d.v.; 217O, abdominal gill I. (PS = parastyli). Figs. 217A-C, F modified from DOMÍNGUEZ, 1985; 217D-E, G-H from DOMÍNGUEZ, 1987b; 217I-J from TRAVER, 1950; 217K-L from ULMER, 1920a.

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remaining longitudinal and cross veins yellowish white; 2) length of fore wings from 18.5 to 19 mm; 3) abdominal terga II-VII brownish orange, VIII-IX brownish black; color pattern as in Fig. 217F; 4) eggs filling abdomen orangish; and 5) legs II and III brownish orange. Distribution: Argentina (Córdoba, Salta). Tortopus harrisi TRAVER Tortopus harrisi TRAVER, 1950: 604 (male); HUBBARD, 1982a: 271; DOMÍNGUEZ, 1985: 69.

Taxonomy: Tortopus harrisi TRAVER is known only from male imagos which can be separated from those of the other species of Tortopus by the following combination of characters: 1) Veins C, Sc, and R1 very weakly shaded with purple in basal 1/3, remaining veins whitish; 2) fore wing length, 9 mm; 3) abdomen yellowish, with basal and medial portions of terga smoothly shaded with brownish gray, principally forming parallel lines at sides of median area lighter, and small black stigmatic spots; 4) legs II and III yellowish; and 5) genitalia as in Figs. 217I-J). Distribution: Brazil (Mato Grosso). Tortopus igaranus NEEDHAM & MURPHY Tortopus igaranus NEEDHAM & MURPHY, 1924: 24 (female); ULMER, 1942: 111; TRAVER, 1950: 600 (female); HUBBARD, 1982a: 271; DOMÍNGUEZ, 1985: 69.

Taxonomy: Tortopus igaranus, the type-species of the genus, is known only from females. It can be separated from the other species of the genus by the following combination of characters: 1) Veins purple; 2) length of fore wings 9.5-12.5 mm; 3) abdomen dark brown, with lighter median line; and 4) legs II and III yellowish. Distribution: Peru (Igará-Paraná and Putumayo rivers). Tortopus obscuripennis DOMÍNGUEZ Tortopus obscuripennis DOMÍNGUEZ, 1985: 71 (female).

Taxonomy: Tortopus obscuripennis is known from female adults and can be separated from the other species of the genus by the following combination of characters: 1) Wings (Figs. 217D-E) with brownish veins; 2) length of fore wings 19.5-20.5 mm; 3) abdominal terga whitish, shaded with brownish gray in a color pattern as in Fig. 217G; 4) eggs filling abdomen yellowish; and 5) legs II and III yellowish. Distribution: Argentina (Salta). Tortopus parishi (BANKS) Campsurus parishi BANKS, 1918: 10 (male); LESTAGE, 1923: 123. Tortopus parishi; ULMER, 1942: 110; TRAVER, 1947b: 372; TRAVER, 1950: 600; HUBBARD, 1982a: 271; DOMÍNGUEZ, 1985: 69.

Taxonomy: Tortopus parishi, known only from males, was described by BANKS (1918) in the genus Campsurus and transferred to Tortopus by ULMER (1942). Practically all that is known of this species is a deficient drawing of the genitalia, its size, and something of the

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coloration. The illustration of the genitalia does not show the basal appendages (parastyli) characteristic of the genus that could be due to its small size or because they are absent. It can be characterized by the following combination of characters: 1) Length of fore wings 10-12 mm; 2) costal veins grayish, with remaining veins whitish; 3) abdomen light, blackish at end; 4) according to BANKS (1918) figure, penes more or less straight as in T. unguiculatus (Fig. 217K). Distribution: Ecuador (Durar). Tortopus sarae DOMÍNGUEZ Tortopus sarae DOMÍNGUEZ, 1985: 71 (female).

Taxonomy: Tortopus sarae, known only from female adults, can be separated from the other species of the genus by the following combination of characters: 1) Veins C, Sc, and R1 brownish black at base, lighter towards apex, remaining veins yellowish gray; 2) length of fore wings 17.5-18.0 mm; 3) abdominal terga whitish and widely shaded with black, color pattern as in Fig. 217H; 4) eggs filling abdomen yellowish; and 5) legs II and III whitish. Distribution: Argentina (Jujuy). Tortopus unguiculatus (ULMER) Campsurus unguiculatus ULMER, 1920a: 4 (male); ULMER, 1920c: 107; LESTAGE, 1923: 123. Tortopus unguiculatus; ULMER, 1932: 208; ULMER, 1942: 109 (male, female); TRAVER, 1950: 600 (male, female); HUBBARD, 1982a: 271; DOMÍNGUEZ, 1985: 69.

Taxonomy: Tortopus unguiculatus is the only South American species known from both sexes. ULMER (1920a) first described the male and later (1942) added the description of the female. This species can be characterized by the following combination of characters: 1) Veins C and Sc violet at base, lighter toward apex, remaining veins whitish; 2) length of fore wings: males, 9-11 mm; females, 11-12 mm; 3) abdominal terga light yellowish, with sides tinted with blackish violet, and median line clear; 4) legs II and III yellowish; 5) eggs filling abdomen yellowish; and 6) genitalia as in Figs. 217K-L. Distribution: Colombia, Central America. Tortopus zottai (NAVÁS) Campsurus zottai NAVÁS, 1920c: 11 (male); LESTAGE, 1923: 124; NAVÁS, 1926c: 110 (male, female); TRAVER, 1947b: 371; BERTHÉLEMY, 1965: 2; HUBBARD, 1982a: 271; ALBA-TERCEDOR & PETERS, 1985: 221. Tortopus zottai; DOMÍNGUEZ, 1987b: 11 (male).

Taxonomy: Tortopus zottai was described by NAVÁS (1920c) in Campsurus, including, theoretically, males and females. In 1923, LESTAGE made reference to an abnormal venational character for the genus and emphasized the lack of information about the male genitalia. NAVÁS (1926c) included a figure of the subgenital plate, but not of the penes nor the basal projections of the forceps, diagnostic of the genus. DOMÍNGUEZ (1987b) studied the material, consisting of 18 specimens, two of which had the label “Typus”, but found that

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all were males, and transferred the species to Tortopus. This species can be separated from the other species of the genus by the following combination of characters: 1) Wing veins (Figs. 217A-B) C, Sc, and R1 yellowish at base, lighter near apex, remaining veins whitish; 2) length of fore wings 12.0-12.5 mm; 3) abdominal terga brownish yellow, tergum I strongly shaded with brown, terga II to IX progressively darker toward back, tergum X yellowish, and median line lighter; 4) legs II and III brownish yellow; and 5) genitalia as in Fig. 217C. Distribution: Argentina (Buenos Aires). INCERTAE SEDIS The species listed here are of uncertain placement and both must be considered nomina dubia. Ephoron umbratum (HAGEN, 1888) Nomen Dubium Palingenia atrostoma (WEBER, 1801) Nomen Dubium

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Plate 218. Figs. 218A-C, progressive eye division between dorsal and ventral portions in Leptophlebiidae males: 218A, Thraulodes sp.; 218B, Miroculis (Ommaethus) sp.; 218C, Miroculis (Atroari) sp. (DP = Eye Dorsal Portion; VP = Eye Ventral Portion; O = Ocellus; A = Antenna).

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Plate 219. Siphlonella ventilans (Oniscigastridae). Figs. 219A-B. 219A, nymph, dorsal view; 219B, habitat (Laguna Chica de Icalma, Southern Chile), arrow showing pole used by nymphs to crawl out of the water. The nymphs were found hidden in the sand of the beaches of the lake. Photo 219A by S. Elliott; 219B by E. Domínguez.

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Plate 220. Siphlonella ventilans (Oniscigastridae). Figs. 220A-E, subimago emergency sequence (whole process lasted 10´). 220A, nymph crawling out of the water on a pole (see Fig. 219B); 220B-E, different stages of the emergence of the male subimago from the nymphal skin. Photos by E. Domínguez.

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E Plate 221. Figs. 221A-E, Different eggs with chorion structures. 221A, Campylocia sp. (Euthyplociidae); 221B, Coryphorus aquilus (Coryphoridae); 221C, Tricorythopsis minimus (Leptohyphidae); 221D, Tricorythodes popayanicus (Leptohyphidae); 221E, Melanemerella brasiliana (Melanemerellidae). (PC = Polar cap; Ma = Micropilar area; Af = Adhesive filament). Fig. 221E modified from MOLINERI & DOMÍNGUEZ, 2003.

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Plate 222. Figs. 222A-F, different Leptophlebiidae eggs with chorion structures. 222A, Farrodes yungaensis; 222B, Hermanella grandis; 222C, Leentvaaria palpalis; 222D, Nousia delicata; 222E, Ulmeritus carbonelli; 222F, same, detail of base of Kct. (Kct = Knob-terminated coiled thread; M = Micropyle; F = Filament). Fig. 222E, modified from DOMÍNGUEZ & CUEZZO, 2002.

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Plate 223. Figs. 223A-D, Traverella (Zonda) calingastensis, egg with chorion structures. 223A, general aspect; 223B, detail of Kct, untriggered; 223C-D, same, with filament uncoiled in different degrees.

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Plate 224. Figs. 224A-B, Chaquihua penai (Ameletopsidae, Ameletopsinae). 224A, male subimago; 224B, female subimago. Photos by E. Domínguez.

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Plate 225. Figs. 225A-B, Ameletopsidae, male imagos. 225A, Chaquihua penai (Ameletopsinae); 225B, Chiloporter eatoni (Chiloporterinae). Photos by E. Domínguez.

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Plate 226. Figs. 226A-B, Chiloporter eatoni (Ameletopsidae, Chiloporterinae) female. 226A, subimago; 226B, imago. Photos by E. Domínguez.

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C Plate 227. Figs. 227A-C, nymphs. 227A, Chiloporter eatoni (Ameletopsidae, Chiloporterinae); 227B, Chaquihua penai (Ameletopsidae, Ameletopsinae); 227C, Murphyella needhami (Coloburiscidae). Photos 227B-C by E. Domínguez.

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Plate 228. Figs. 228A-C, Baetidae. 228A, Varipes minutus, male nymph; 228B, Harpagobaetis gulosus, nymph, detail of carnivorous mouthparts; 228C, Callibaetis radiatus, female imago.

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Plate 229. Figs. 229A-B, Leptohyphidae. 229A, Tricorythopsis gibbus, nymph; 229B, Leptohyphes sp., male imago. Photos by E. Domínguez.

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Plate 230. Figs. 230A-B, Penaphlebia flavidula (Leptophlebiidae). 230A, nymph; 230B, male imago. Photo 230A by S. Elliott; 230B by G. Cuezzo.

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B Plate 231. Figs. 231A-B. Leptophlebiidae. 231A, Tikuna atramentum, nymph; 231B, Thraulodes liminaris, female subimago. Photo 231A by R.W. Flowers; 231B by G. Cuezzo.

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Plate 232. Figs. 232A-B. Leptophlebiidae, male imagos. 232A, Hagenulopsis sp.; 232B, Miroculis sp. male imago. Photo 232A by G. Cuezzo; Photo 232B by E. Domínguez.

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Plate 233. Figs. 233A-E. 233A-B, Mayobaetis sp.: 233A, general aspect of habitat; 233B, nymph on vertical wall. 233C-D, Tortopus sp. nymph: 233C, detail of tunnel; 233D, dorsal view; 233E, lateral view; Photos 233A-C by C. Molineri; Photos 233D-E by E. Domínguez.

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10 GUIDE TO CHANGES IN SCIENTIFIC NAMES This is a list of the names that have been synonymized. The first names are the old names, and the second names are the valid ones. Acerpenna robacki = Americabaetis robacki Adenophlebia colombiae = Thraulodes colombiae Allenhyphes asperlus = Allenhyphes? asperulus Allenhyphes edmundsi = Traverhyphes (Mocoihyphes) edmundsi Allenhyphes minimus = Tricorythopsis minimus Allenhyphes nanus = Traverhyphes (Byrsahyphes) nanus Allenhyphes rallus = Yaurina ralla Allenhyphes tinctus = Tricorythopsis artigas Allenhyphes viriosus = Tricorythopsis minimus Alloydia cacautana = Lachlania cacautana Ameletoides fuegiensis = Metamonius anceps Ameletopsis eatoni = Chiloporter eatoni Ameletus fuegiensis = Metamonius anceps Asioplax nicholsae = Tricorythodes nicholsae Asthenopodes albicans = Asthenopus picteti Asthenopodes picteti = Asthenopus picteti Asthenopus albicans = Asthenopus picteti Asthenopus amazonicus = Asthenopus curtus Asthenopus dorsalis = Campsurus dorsalis Atalonella grandis = Nousia grandis Atalonella maculata = Nousia maculata Atalonella minor = Nousia minor Atalonella ophis = Nousia delicata Atalophlebia anastasii = Penaphlebia anastasii Atalophlebia anastomosis = Hapsiphlebia anastomosis Atalophlebia athanasii = Penaphlebia athanasii Atalophlebia axilata = Massartella brieni

Atalophlebia brieni = Massartella brieni Atalophlebia chilensis (partim) = Penaphlebia chilensis Atalophlebia chilensis (partim) = Meridialaris patagonica Atalophlebia discolor = Penaphlebia discolor Atalophlebia fenestrata = Penaphlebia fenestrata Atalophlebia fulvipes (partim) = Meridialaris lestagei Atalophlebia fulvipes (partim) = Penaphlebia fulvipes Atalophlebia hyalina = Penaphlebia hyalina Atalophlebia patagonica = Meridialaris patagonica Atalophlebia rofucensis = Penaphlebia rofucensis Atalophlebia sepia = Penaphlebia fulvipes Atalophlebia valdiviae = Penaphlebia valdiviae Atalophlebia vinosa = Penaphlebia vinosa Atalophlebioides haarupi = Ulmeritoides haarupi Atalophlebioides flavopedes = Ulmeritoides flavopedes Atalophlebioides lestagei = Meridialaris lestagei Atalophlebioides sao-paulense = Ulmeritus saopaulensis Atolophlebia rofucensis = Penaphlebia rofucensis Baetis abundans = Callibaetis abundans Baetis albivitta = Hexagenia (Pseudeatonica) albivitta Baetis alcyoneus = Camelobaetidius alcyoneus Baetis angolinus = Andesiops angolinus Baetis aymara = Cloeodes aymara Baetis coveloae = Camelobaetidius coveloae Baetis dryops = Camelobaetidius dryops Baetis ellenae = Mayobabetis ellenae Baetis gloriosus = Callibaetis fasciatus

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Baetis guttata = Siphlonella guttata Baetis nocturnus = Cloeodes nocturnus Baetis oldendorffi = Americabaetis oldendorffi Baetis opacus = Callibaetis zonalis Baetis peruvianus = Andesiops peruvianus Baetis stelzneri = Cloeodes stelzneri Baetis tantillus = Camelobaetidius tantillus Baetis virellus = Callibaetis zonalis Baetis weiseri = Andesiops peruvianus Baetodes spinifer = Baetodes spiniferum Bruchella nigra = Leptohyphes eximius Caenis ludiera = Caenis ludicra Calliarcys laetus = Thraulodes laetus Callibaetis alegre = Callibaetis willineri Callibaetis amoenus = Callibaetis zonalis Callibaetis apertus = Callibaetis zonalis Callibaetis apicatus =Callibaetis guttatus Callibaetis bruchius =Callibaetis guttatus Callibaetis depressus = Callibaetis zonalis Callibaetis fasciatus = Callibaetis sellacki Callibaetis gloriosus = Callibaetis fasciatus Callibaetis jaffueli = Callibaetis jocosus Callibaetis jocosa = Callibaetis jocosus Callibaetis lineatus = Callibaetis sellacki Callibaetis rimatus = Callibaetis jocosus Callibaetis sobrius = Callibaetis zonalis Callibaetis spegazzinus = Callibaetis jocosus Callibaetis stictogaster = Callibaetis jocosus Callibaetis trifasciatus (partim) = Callibaetis fasciatus Callibaetis trifasciatus (partim) = Callibaetis sellacki Callibaetis vitreus = Callibaetis zonalis Callibaetis zonatus = Callibaetis guttatus Campsurus albicans (partim) = Asthenopus picteti Campsurus amazonicus = Asthenopus curtus Campsurus bruchianus = Tortopus bruchianus Campsurus curtus = Asthenopus curtus Campsurus jorgensini = Campsurus jorgenseni Campsurus paraguarius = Campsurus paraquarius Campsurus parishi = Tortopus parishi Campsurus unguiculatus = Tortopus unguiculatus Campsurus zottai = Tortopus zottai Campylocia ampla = Campylocia anceps Campylocia burmeisteri = Campylocia anceps Chiloporter penai = Chiloporter eatoni

Choroterpes bilineata = Tikuna bilineata Choroterpes (Choroterpes) bilineata = Tikuna bilineata Choroterpes emersoni = Miroculitus emersoni Cloe fasciata = Callibaetis fasciatus Cloe lorentzii = Callibaetis sellacki Cloe sellacki = Callibaetis sellacki Cloe siewertii = Thraulus siewertii Cloe stelzneri = Cloeodes stelzneri Cloe vogleri = Thraulus vogleri Cloeodes (Notobaetis) penai = Cloeodes penai Cloeon brunneum = Americabaetis peterseni Cloeon gonzalezi = Callibaetis gonzalezi Cloeon jorgenseni = Americabaetis jorgenseni Cloeon oldendorffii = Americabaetis oldendorffi Cloeon peterseni = Americabaetis peterseni Cloeon vitripennis = Thraulodes vitripennis Dactylobaetis anubis = Camelobaetidius anubis Dactylobaetis cayumba = Camelobaetidius cayumba Dactylobaetis penai = Camelobaetidius penai Dactylobaetis phaedrus = Camelobaetidius phaedrus Deceptiviosa angolina = Andesiops angolinus Deceptiviosa ardua = Andesiops ardua Deceptiviosa torrens = Andesiops torrens Deleatidium biobionicum = Meridialaris biobionica Deleatidium chiloeense = Meridialaris chiloeensis Deleatidium diguillinum = Meridialaris diguillina Deleatidium haarupi = Ulmeritoides haarupi Deleatidium illapeli = Meridialaris laminata Deleatidium laminatum = Meridialaris laminata Deleatidium patagonicum = Meridialaris patagonica Deleatidium penai = Meridialaris chiloeensis Deleatidium vittatum = Ulmeritoides haarupi Dictyosiphlon molinai = Murhpyella needhami Ecdyonurus guttatus = Siphlonella guttata Ecdyurus guttatus = Siphlonella guttata Ephemera albicans = Campsurus albicans Ephemera-Baetis guttata = Siphlonella guttata Ephemera colombiae = Thraulodes colombiae Ephemera (Cloe) vitripennis = Thraulodes vitripennis Ephemera holmbergii = Campsurus holmbergii Ephemera wappaei = Campsurus wappaei Eurycaenis tenella = Brachycercus tenella Euthyplocia anceps = Campylocia anceps

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Euthyplocia bullocki = Chaquihua bullocki Euthyplocia burmeisteri = Campylocia anceps Euthyplocia guntheri = Campylocia anceps Euthyplocia intercalata = Campylocia anceps Hagenulopsis minutus = Hagenulopsis minuta Haplohyphes furtiva = Haplohyphes baritu Haplohyphes furtivus = Haplohyphes baritu Heptagenia guttata = Siphlonella guttata Heptagenia? mollinai = Murphyella needhami Hermanella velma = Hermanella thelma Hermanella incertans = Hermanellopsis incertans Hermanella (Hermanellopsis) incertans = Hermanellopsis incertans Hexagenia albivitta = Hexagenia (Pseudeatonica) albivitta Hexagenia benedicta = Hexagenia (Pseudeatonica) albivitta Hexagenia continua = Hexagenia (Pseudeatonica) albivitta Hexagenia dominans = Hexagenia (Pseudeatonica) albivitta Homothraulus convexus = Paramaka convexa Homothraulus maculatus = Farrodes maculatus Leptohyphes albus = Leptohyphes tacajalo Leptohyphes asperulus = Allenhyphes? asperulus Leptohyphes bruchi = Leptohyphes eximius Leptohyphes comatus = Leptohyphes maculatus Leptohyphes echinatus = Leptohyphes setosus Leptohyphes edmundsi = Traverhyphes (Mocoihyphes) edmundsi Leptohyphes flinti = Allenhyphes flinti Leptohyphes gibbus = Tricorythopsis gibbus Leptohyphes hirsutus = Leptohyphes maculatus Leptohyphes indicator = Traverhyphes (Traverhyphes) indicator Leptohyphes minimus = Tricorythopsis minimus Leptohyphes myllonotus = Leptohyphes maculatus Leptohyphes mithras = Haplohyphes mithras Leptohyphes nanus = Traverhyphes (Byrsahyphes) nanus Leptohyphes nicholsae = Tricorythodes nicholsae Leptohyphes niger = Leptohyphes eximius Leptohyphes nigra = Leptohyphes eximius Leptohyphes pereirae = Leptohyphes plaumanni

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Leptohyphes rallus = Yaurina ralla Leptohyphes spinosus = Allenhyphes? spinosus Leptohyphes tinctus = Tricorythopsis artigas Leptohyphes undulatus = Tricorythopsis undulatus Leptohyphes viriosus = Tricorythopsis minimus Leptohyphodes australis = Tricorythodes australis Leptophlebia colombiae = Thraulodes colombiae Longinella guntheri = Campylocia anceps Massartella fruhstorferri = Massartella brieni Meridialaris chiloeense = Meridialaris chiloeensis Meridialaris illapeli = Meridialaris laminata Meridialaris penai = Meridialaris chiloeensis Metamonius fuegiensis = Metamonius anceps Metamonius hollermayeri = Siphlonella ventilans Metamonius needhami = Murhpyella needhami Murphyella molinai = Murhpyella needhami Neophlebia garciai = Lachlania garciai Notobaetis penai = Cloeodes penai Nousia ophis = Nousia delicata Noya garciai = Lachlania garciai Noya pallipes = Lachlania pallipes Noya radai = Lachlania radai Oligoneuria anomala = Oligoneurioides amazonicus Ordella albata = Caenis albata Ordella ludicra = Caenis ludicra Ordella nemoralis = Caenis nemoralis Oxycypha oldendorffii = Americabaetis oldendorffi Palingenia albicans (partim) = Asthenopus picteti Palingenia albicans (partim) = Campsurus albicans Palingenia albifilum (partim) = Asthenopus curtus Palingenia albifilum (partim) = Campsurus albifilum Palingenia continua = Hexagenia (Pseudeatonica) albivitta Palingenia curta = Asthenopus curtus Palingenia dorsalis = Campsurus dorsalis Palingenia dorsigera = Hexagenia (Pseudeatonica) albivitta Palingenia hecuba = Euthyplocia hecuba Palingenia latipennis = Campsurus latipennis Palingenia nappii = Campsurus nappii Penaphlebia sepia = Penaphlebia fulvipes Perissophlebia flinti = Perissophlebiodes flinti Potamanthus inanis = Leptohyphodes inanis Potamanthus siewertii = Thraulus siewertii

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Potamanthus vogleri = Thraulus vogleri Pseudeatonica mexicana = Hexagenia (Pseudeatonica) mexicana Pseudocloeon albinerve = Americabaetis albinervis Pseudocloeon anduzei = Cloeodes anduzei Pseudocloeon arawak = Baetodes arawak Pseudocloeon binocularis = Cloeodes binocularis Pseudocloeon bridarolli = Americabaetis bridarolli Pseudocloeon bruchi = Americabaetis bruchi Pseudocloeon brunneum = Americabaetis peterseni Pseudocloeon jorgenseni = Americabaetis jorgenseni Pseudocloeon oldendorffi = Americabaetis oldendorffi Pseudocloeon peterseni = Americabaetis peterseni Pseudocloeon turbinops = Cloeodes turbinops Pseudocloeon venezuelensis = Cloeodes venezuelensis Pseudocloeon weiseri = Americabaetis weiseri Simothraulopsis surinamensis = Simothraulopsis demerara Siphlonella fazi = Siphlonella ventilans Siphlonurus fazi = Siphlonella ventilans Siphlurus? anceps = Metamonius anceps Spaniophlebia pallipes = Lachlania pallipes Spaniophlebia traili = Spaniophlebia trailiae Thraulodes flavopedes = Ulmeritoides flavopedes Thraulodes laeta = Thraulodes laetus Thraulodes loetus = Thraulodes laetus Thraulodes plicatus = Thraulodes telegraphicus Thraulodes trijuncta = Thraulodes trijunctus Thraulus bomplandi = Thraulodes bomplandi Thraulus bradleyi = Traverella (Traverella) bradleyi Thraulus caribbianus = Farrodes caribbianus Thraulus colombiae = Thraulodes colombiae Thraulus convexus = Paramaka convexa Thraulus costalis = Homothraulus costalis Thraulus demerara = Simothraulopsis demerara Thraulus ehrhardti = Needhamella ehrhardti

Thraulus haarupi = Ulmeritoides haarupi Thraulus laetus = Thraulodes laetus Thraulus larensis = Homothraulus larensis Thraulus lorensis = Homothraulus larensis Thraulus maculatus = Farrodes maculatus Thraulus maculipennis = Hermanella maculipennis Thraulus montium = Traverella (Traverella) montium Thraulus misionensis = Homothraulus misionensis Thraulus roundsi = Farrodes roundsi Thraulus trijunctus = Thraulodes trijunctus Thraulus valdemari = Traverella (Traverella) valdemari Thraulus vitripennis = Thraulodes vitripennis Tortopus dorsalis = Campsurus dorsalis Traverella ehrhardti = Needhamella ehrhardti Traverella maculipennis = Hermanella maculipennis Tricorythopsis artigas (partim) = Tricorythopsis minimus Tricorythopsis fictilis = Tricorythopsis artigas Tricorythopsis petersorum = Tricorythopsis undulatus Tricorythus australis = Tricorythodes australis Ulmeritoides fidalgoi = Ulmeritoides haarupi Ulmeritus adustus = Ulmeritoides uruguayensis Ulmeritus (Pseudulmeritus) flavopedes = Ulmeritoides flavopedes Ulmeritus luteotinctus = Ulmeritoides luteotinctus Ulmeritus (Ulmeritoides) luteotinctus = Ulmeritoides luteotinctus Ulmeritus (Ulmeritoides) uruguayensis = Ulmeritoides uruguayensis Ulmeritus (Ulmeritus) haarupi = Ulmeritoides haarupi Ulmeritus patagiatus = Ulmeritoides patagiatus Ulmeritus sao-paulense = Ulmeritus saopaulensis

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SALLES F.F., LUGO-ORTIZ, C.R. & E.R. DA-SILVA (2004b): Descrição da fêmea adulta de Americabaetis titthion (Ephemeroptera: Baetidae). - Acta Zoológica Mexicana 20 (1): 23-26. SALLES F.F., PEREIRA, S.M. & J.E. SERRAO (2005): Redescription of Camelobaetidius leentvari Demoulin, 1966 from Suriname and Brazil (Ephemeroptera, Baetidae). - Ephemera 5 (2): 69-75. SALLES, F.F., LUGO-ORTIZ, C.R., DA-SILVA, E.R. & C.N. FRANCISCHETTI (2003a): Novo gênero e espécie de Baetidae (Insecta, Ephemeroptera) do Brasil. - Arquivos do Museu Nacional (Rio de Janeiro) 61 (1): 23-30. SATTLER, W. (1967): Über die Lebensweise, insbesondere das Bauverhalten, neotropischer Einstagsfliegen-Larven (Ephemeroptera: Polymitarcyidae). - Beiträge zur Neotropischen Fauna 5: 89-110. SAVAGE, H.M. (1982): A curious new genus and species of Atalophlebiinae (Ephemeroptera: Leptophlebiidae) from the southern coastal mountains of Brazil. - Studies on Neotropical Fauna and Environment 17: 209-217. SAVAGE, H.M. (1983a): Perissophlebiodes, a replacement name for Perissophlebia SAVAGE nec TILLYARD (Ephemeroptera: Leptophlebiidae). - Entomological News 94: 204. SAVAGE, H.M. (1983b): Wing evolution within Miroculis and related genera (Ephemeroptera: Leptophlebiidae) from northern South America. - Zeitschrift für zoologische Systematik und Evolutionsforschung 21: 124-142. SAVAGE, H.M. (1986): Systematics of the Terpides lineage from the Neotropics: Definition of the Terpides lineage, methods, and revision of Fittkaulus SAVAGE & PETERS. - Spixiana 9: 255-270. SAVAGE, H.M. (1987a): Two new species of Miroculis from Cerro de la Neblina, Venezuela with new distribution records for Miroculis fittkaui and Microphlebia surinamensis (Ephemeroptera: Leptophlebiidae). - Aquatic Insects 9: 97-108. SAVAGE, H.M. (1987b): Biogeographic classification of the Neotropical Leptophlebiidae (Ephemeroptera) based upon geological centers of ancestral origin and ecology. - Studies on Neotropical Fauna and Environment 22: 199-222. SAVAGE, H.M., & E. DOMÍNGUEZ (1992): A new genus of Atalophlebiinae (Ephemeroptera, Leptophlebiidae) from northern South America. - Aquatic Insects 14: 243-248. SAVAGE, H.M. & W.L. PETERS (1978): Fittkaulus maculatus a new genus and species from northern Brazil (Leptophlebiidae: Ephemeroptera). - Acta Amazonica 8: 293-298. SAVAGE, H.M. & W.L. PETERS (1983): Systematics of Miroculis and related genera from northern South America. - Transactions of the American Entomological Society 108: 491-600. SAVAGE, H.M., FLOWERS, R.W. & W. PORRAS (2005): Rediscovery of Choroterpes atramentum in Costa Rica, type species of Tikuna new genus (Ephemeroptera: Leptophlebiidae: Atalophlebiinae), and its role in the “Great American Interchange”. - Zootaxa 932:1-14. SOLDÁN, T. (1986): A revision of the Caenidae with ocellar tubercles in the nymphal stage (Ephemeroptera). - Acta Universitatis Carolinae, Biologica 1982-1984: 289-362. SPIETH, H.T. (1933): The phylogeny of some mayfly genera. - Journal of New York Entomological Society 41: 327-390. SPIETH, H.T. (1941): Taxonomic studies on the Ephemeroptera. II. The genus Hexagenia. American Midland Naturalist 26: 233-280. SPIETH, H.T. (1943): Taxonomic studies on the Ephemeroptera. III. Some interesting Ephemerids from Surinam and other Neotropical localities. - American Museum Novitates 1244: 1-13. STEPHENS, J.F. (1835): Illustrations of British Entomology, Mandibulata. - Vol. 6. Baldwin, London.

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SUTER, P.J. (1986): The Ephemeroptera (mayflies) of South Australia. - Records of the South Australian Museum 19: 339-397. SWEENEY, B.W., JACKSON, J.K. & D.H. FUNK (1995): Semivoltinism, seasonal emergence, and adult size variation in a tropical stream mayfly (Euthyplocia hecuba). - Journal of the North American Benthological Society 14 (1): 131-146. TAKEDA, M.S. & M. GRZYBKOWSKA (1997): Seasonal dynamics and production of Campsurus violaceous nymphs (Ephemeroptera, Polymitarcyidae) in the Baía river, upper Paraná river floodplain, Brazil. - Hydrobiologia: 356: 149-155. THEW, T.B. (1958): Dodd’s types of two species of Callibaetis (Ephemeroptera). - Entomological News 69: 132. THEW, T.B. (1960a): Revision of the genera of the family Caenidae (Ephemeroptera). - Transactions of the American Entomological Society 86: 187-205. THEW, T.B. (1960b): Taxonomic studies on some Neotropical Leptophlebiid mayflies (Ephemeroptera: Leptophlebiidae). - Pan-Pacific Entomologist 36: 119-132. THOMAS, A., DOMINIQUE, Y. & V. HOREAU (2001a): Les Éphémères de la Guyane Française. 7. Redescription de Camelobaetidius cayumba (TRAVER & EDMUNDS, 1968) (Ephemeroptera, Baetidae). - Ephemera 3 (2): 117-122. THOMAS, A., PERU, N. & V. HOREAU (2001b): Les Éphémères de la Guyane Française. 8. Description de Camelobaetidius matilei n. sp. et clé de détermination des espèces guyanaises de ce genre (Ephemeroptera, Baetidae). - Ephemera 3 (2): 123-133. THOMAS, A. & N. PERU (2002): Les Éphémères de la Guyane Francaise. 6. Description de Waltzoyphius roberti n. sp. (Ephemeroptera, Baetidae). - Bulletin de la Société d´Histoire Naturelle de Toulouse 138: 15-20. THOMAS, A., BOUTONNET, J., PERU, N. & V. HOREAU (2004): Les Éphémères de la Guyane Française. 9. Descriptions d’Hydrosmilodon gilliesae n. sp. et d’ H. mikei n. sp. (Ephemeroptera, Leptophlebiidae). - Ephemera 4: 65-80. THOMAS, A., ORTH, K., HOREAU, V. & Y. DOMINIQUE (2000): Les Éphémères de la Guyane Française. 3. Bessierus doloris n. gen., n. sp. (Ephemeroptera: Leptophlebiidae). - Ephemera 2: 49-57. TRAVER, J.R. (1935): Systematic portion in NEEDHAM, TRAVER, & HSU (1935): 239-739. TRAVER, J.R. (1938): Mayflies of Puerto Rico. - Journal of Agriculture of the University of Puerto Rico 22: 5-42, pl. 1-3. TRAVER, J.R. (1943): New Venezuelan mayflies. - Boletin de Entomologia Venezolana 2: 79-98. TRAVER, J.R. (1944): Notes on Brazilian mayflies. - Boletim do Museu Nacional, Nova Série, Zoología (Rio de Janeiro) 22: 2-53. TRAVER, J.R. (1946): Notes on Neotropical mayflies. Part I. Family Baetidae, subfamily Leptophlebiinae. - Revista de Entomologia 17: 418-436. TRAVER, J.R. (1947a): Notes on Neotropical mayflies. Part II. Family Baetidae, subfamily Leptophlebiinae. - Revista de Entomologia 18: 149-160. TRAVER, J.R. (1947b): Notes on Neotropical mayflies. Part III. Family Ephemeridae. - Revista de Entomologia 18: 370-395. TRAVER, J.R. (1950): Notes on Neotropical mayflies. Part. IV. Family Ephemeridae (continued). Revista de Entomologia 21: 593-614. TRAVER, J.R. (1956a): A new genus of Neotropical mayflies (Ephemeroptera, Leptophlebiidae). Proceedings of the Entomological Society of Washington 58: 1-13.

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TRAVER, J.R. (1956b): The genus Asthenopodes (Ephemeroptera). - Comunicaciones Zoológicas del Museo de Historia Natural de Montevideo 4 (75): 1-10, pl. 1-2. TRAVER, J.R. (1958a): The subfamily Leptohyphinae (Ephemeroptera: Tricorythidae). Part I. Annals of the Entomological Society of America 51: 491-503. TRAVER, J.R. (1958b): Some Mexican and Costa Rican mayflies. - Bulletin of the Brooklyn Entomological Society 53 (4): 81-89. TRAVER, J.R. (1959a): The subfamily Leptohyphinae. Part II: Five new species of Tricorythodes (Ephemeroptera, Tricorythidae). - Proceedings of the Entomological Society of Washington 61: 121-131. TRAVER, J.R. (1959b): Uruguayan mayflies. Family Leptophlebiidae: Part I. - Revista de la Sociedad Uruguaya de Entomologia 3: 1-13, pl. 1-3. TRAVER, J.R. (1960a): Uruguayan mayflies. Family Leptophlebiidae: Part II. - Revista de la Sociedad Uruguaya de Entomologia 4: 19-28. TRAVER, J.R. (1960b): Uruguayan mayflies. Family Leptophlebiidae: Part III. - Revista de la Sociedad Uruguaya de Entomologia 4: 73-85, pl. 1. TRAVER, J.R. (1963): Uruguayan mayflies. Family Leptophlebiidae: Part IV. - Revista de la Sociedad Uruguaya de Entomologia 5: 25-31. TRAVER, J.R. (1964): A new species of Thraulodes from Uruguay (Ephemeroptera: Family Leptophlebiidae). - Revista de la Sociedad Uruguaya de Entomologia 6: 33-37. TRAVER, J.R. (1971): Four new species of Neotropical Baetis (Ephemeroptera: Baetidae). Proceedings of the Entomological Society of Washington 73: 58-63. TRAVER, J.R. & G.F. EDMUNDS JR. (1967): A revision of the genus Thraulodes (Ephemeroptera: Leptophlebiidae). - Miscellaneous Publications of the Entomological Society of America 5: 349-395. TRAVER, J.R. & G.F. EDMUNDS JR. (1968): A revision of the Baetidae with spatulate-clawed nymphs (Ephemeroptera). - Pacific Insects 10: 629-677. TSHERNOVA, O.A. (1962): Order Ephemeroptera. Mayflies - In: ROHDENDORF, B.B. (ed.). Osnovy Paleontologi: 55-64. Vol. 9. Izdat. Akad. Nauk, Moscow (in Russian). TSUI, P.T.P. & W.L. PETERS (1972): The comparative morphology of the thorax of selected genera of the Leptophlebiidae (Ephemeroptera). - Journal of Zoology 168: 309-367. TSUI, P.T.P. & W.L. PETERS (1975): The comparative morphology and phylogeny of certain Gondwanian Leptophlebiidae based on the thorax, tentorium, and abdominal terga (Ephemeroptera). - Transactions of the American Entomological Society 101: 505-595. ULMER, G. (1904): Ephemeriden. - Hamburger Magalhaensische Sammelreise 1904: 1-8, 1 pl. ULMER, G. (1920a): Neue Ephemeropteren. - Archiv für Naturgeschichte 85 (A): 1-80. ULMER, G. (1920b): Über die Nymphen einiger exotischer Ephemeropteren. - Festschrift für Zschokke 25: 1-25. ULMER, G. (1920c): Übersicht über die Gattungen der Ephemeropteren, nebst Bemerkungen über einzelne Arten. - Stettiner Entomologische Zeitung 81: 97-144. ULMER, G. (1921): Über einige Ephemeropteren-Typen älterer Autoren. - Archiv für Naturgeschichte 87: 229-267. ULMER, G. (1924): Einige alte und neue Ephemeropteren. - Konowia 3: 23-37. ULMER, G. (1932): Bemerkungen über die seit 1920 neu aufgestellten Gattungen der Ephemeropteren. - Stettiner Entomologische Zeitung 93: 204-219.

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ULMER, G. (1933): Aquatic insects of China. Article VI. Revised key to the genera of Ephemeroptera. - Peking Natural History Bulletin 7: 195-218, pl. 1-2. ULMER, G. (1938): Chilenische Ephemeropteren, hauptsächlich aus dem Deutschen Entomologischen Institut, Berlin-Dahlem. - Arbeiten über morphologische und taxonomische Entomologie aus Berlin-Dahlem 5: 85-108. ULMER, G. (1942): Alte und neue Eintagsfliegen (Ephemeropteren) aus Süd- und Mittelamerika. Stettiner Entomologische Zeitung 103: 98-128. ULMER, G. (1943): Alte und neue Eintagsfliegen (Ephemeropteren) aus Süd- und Mittelamerika. Stettiner Entomologische Zeitung 104: 14-46. WAIS, I. (1985): Stratégies adaptatives aux eaux courantes des invertébrés du bassin du fleuve Negro, Patagonie, Argentina. - Verhandlungen der internationalen Vereinigung für theoretische und angewandte Limnologie 22: 2167-2172. WAIS, I. & A.A. BONETTO (1988): Analysis of the allochthonous organic matter and associated macroinvertebrates in some streams of Patagonia (Argentina). - Verhandlungen der internationalen Vereinigung für theoretische und angewandte Limnologie 23: 1455-1459. WALKER, F. (1853): Ephemerinae. - List of the specimens of neuropterous insects in the collection of the British Museum, Part III (Termitidae-Ephemeridae): 533-585. WALKER, F. (1860): Characters of undescribed Neuroptera. - Transactions of the Entomological Society of London 5: 176-199. WALSH, B.D. (1863): Observations on certain N. A. Neuroptera, by H. HAGEN, M. D., of Koenigsberg, Prussia; translated from the original French MS, and published by permission of the author, with notes and descriptions of about twenty new N. A. species of Pseudoneuroptera. - Proceedings of the Entomological Society of Philadelphia 2: 167-272. WALTZ, R.D. & W.P. MCCAFFERTY (1985): Moribaetis: A new genus of Neotropical Baetidae (Ephemeroptera). - Proceedings of the Entomological Society of Washington 87: 239-251. WALTZ, R.D. & W.P. MCCAFFERTY (1986): Apobaetis etowah TRAVER, a new combination in Nearctic Baetidae (Ephemeroptera). - Proceedings of the Entomological Society of Washington 88: 191. WALTZ, R.D. & W.P. MCCAFFERTY (1987a): Generic revision of Cloeodes and description of two new genera (Ephemeroptera: Baetidae). - Proceedings of the Entomological Society of Washington 89: 177-184. WALTZ, R.D. & W.P. MCCAFFERTY (1987b): Systematics of Pseudocloeon, Acentrella, Baetiella, and Liebebiella, new genus (Ephemeroptera: Baetidae). - Journal of the New York Entomological Society 95: 553-568. WALTZ, R.D. & W.P. MCCAFFERTY (1987c): Revision of the genus Cloeodes TRAVER (Ephemeroptera: Baetidae). - Annals of the Entomological Society of America 80: 191-207. WALTZ, R.D. & W.P. MCCAFFERTY (1987d): New genera of Baetidae for some Nearctic species previously included in Baetis LEACH (Ephemeroptera). - Annals of the Entomological Society of America 80: 667-670. WANG, T.-Q. & W.P. MCCAFFERTY (1996): Redescription and reclassification of the South American mayfly Melanemerella brasiliana (Ephemeroptera: Leptophlebiidae). - Entomological News 107 (2): 99-103. WANG, T.-Q., SITES, R.W. & W.P. MCCAFFERTY (1998): Two new species of Leptohyphes (Ephemeroptera: Leptohyphidae) from Ecuador. - Florida Entomologist 81: 68-75. WEBER, F. (1801): Observationes entomologicae continentes novorum quae condidit generum characteres et nuper detectarum specierum descriptiones: 99-100. - Bibliopol. Academici Novi, Kiliae (Kiel).

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WESTWOOD, J.O. (1840): An introduction to the modern classification of insects. - Volume 2. Longman, Orme, Brown & Green, London. WEYENBERGH, H. (1883): Bijdrage tot de Kennis der zuid-amerikaansche Ephemeriden. - Tijdschrift voor Entomologie 26: 159-174, pl. 10 figs. 1-6. WIERSEMA, N.A. & W.P. MCCAFFERTY (2000): Generic Revision of the North and Central American Leptohyphidae (Ephemeroptera: Pannota). - Transactions of the American Entomological Society 126 (3-4): 337-371. WIERSEMA, N.A. & W.P. MCCAFFERTY (2003): Ableptemetes: a new genus of Tricorythodinae (Ephemeroptera: Leptohyphidae) from Mexico and Central America. - Entomological News 114: 37-40. WILLIAMSON, H. (1802): On the Ephoron leukon, usually called the white fly of the Passaick River. Transactions of the American Philosophical Society 5: 71-73. ZUÑIGA, M DEL C., MOLINERI, C. & E. DOMÍNGUEZ (2004): El Orden Ephemeroptera (Insecta) en Colombia. - In: FERNANDEZ, F., ANDRADE, G. & G. AMAT (eds): Insectos de Colombia: 17-42. Vol. 3. Universidad Nacional de Colombia, Bogota. ZUÑIGA, M. DEL C., ROJAS, A.M. & S. MOSQUERA (1997): Biological aspects of Ephemeroptera in rivers of southwestern Colombia (South America). - In: LANDOLT, P. & M. SARTORI (eds.): Ephemeroptera and Plecoptera: Biology-Ecology-Systematics: 216-268. Mauron + Tinguely & Lachat, Fribourg.

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12 SUBJECT INDEX Ableptemetes, 244 Acerpenna, 66 Adebrotus, 60, 64 amazonicus, 64, 65 Afroplocia, 234 Allenhyphes, 244, 245, 246, 247, 248, 250, 252, 294, 295 asperulus, 250, 252, 253, 254 flinti, 250, 251, 252, 253 spinosus, 250, 252, 253, 254 Ameletoides, 536 Ameletopsidae, 28, 29, 45, 47, 49, 51, 52, 46, 597, 598 Ameletopsinae, 52, 53, 597 Ameletopsis, 52, 54, 55 Americabaetis, 58, 60, 61, 63, 66, 67, 69, 70, 71, 72 albinervis, 73 alphus, 66, 67, 68, 69 bridarolli, 69 bruchi, 73 jorgenseni, 69 labiosus, 67, 69, 70, 71 longetron, 67, 69, 70, 71 maxifolium, 67, 69, 70, 71 oldendorffii, 72 peterseni, 72 robacki, 67, 69, 71, 72 titthion, 67, 71, 72 weiseri, 73 Andesiops, 58, 60, 61, 63, 74, 75, 76 angolinus, 76 *

Figures in bold

*

ardua, 76, 77, 78 peruvianus, 74, 76, 78, 79 torrens, 74, 76, 77, 78, 79 Apobaetis, 58, 61, 62, 80, 81, 82, 83 fiuzai, 80, 83 indeprensus, 80, 82 signifer, 80, 83, 84 Archethraulodes, 333, 340, 347, 348 spatulus, 347, 348 Asioplax, 296, 298, 310 Askola, 332, 348, 349, 351 froehlichi, 348, 349, 350, 351 Asthenopodes, 560, 562 picteti, 562 Asthenopodichnium, 558 Asthenopodinae, 557, 558 Asthenopus, 557, 558, 559, 560, 561, 562, 577 curtus, 559, 560, 561 gilliesi, 559, 560, 561, 562 picteti, 559, 560, 561, 562 Atalonella, 453 fusca, 453 ophis, 453, 459 Atalophlebia, 387, 465 brieni, 412, 417 chilensis, 465, 472 Atalophlebiinae, 331 Atalophlebioides, 422 lestagei, 431 Atopophlebia, 335, 337, 343, 345, 350, 352, 353 flowersi, 352, 353, 354 fortunensis, 350, 352, 353, 354, 355

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obrienorum, 352, 353, 354 yarinacocha, 352, 353, 355 Aturbina, 57, 60, 61, 63, 84, 85, 87 beatrixae, 84, 85, 86, 87 georgei, 84, 85, 86, 87 Baetidae, 11, 20, 24, 28, 29, 43, 46, 57, 194, 331, 487, 536, 601 Baetis, 66, 104, 106, 148, 156, 165, 556 Baetiscidae, 18 Baetodes, 20, 58, 86, 88, 89, 91, 96, 99, 101, 103, 161, 172 andamagensis, 90, 92, 93, 96 arawak, 89, 91, 93 chilloni, 90, 92, 95, 96 cochunaensis, 91, 93, 95, 96 copiosus, 91, 93, 94, 95 gibbus, 90, 92, 96 huaico, 88, 89, 97, 99 itatiayanus, 89, 91, 97 levis, 90, 92, 97 pehuenche, 90, 92, 98, 99 peniculus, 90, 92, 98, 99 proiectus, 90, 92, 98, 101 pseudogibbus, 90, 92, 98, 101 rutilus, 91, 93, 100, 101 sancticatarinae, 89, 91, 100 serratus, 86, 91, 93, 100 solus, 91, 93, 102, 103 spinae, 90, 92, 102, 103 spinifer, 89, 91, 102 traverae, 91, 93, 102, 103 uruguai, 88, 89, 90, 91, 92, 103, 104 yuracare, 90, 92, 103, 104 Bernerius, 59, 62, 104 incus, 104, 105, 106 Bessierus, 339, 347, 356 doloris, 356, 357 Brachycercus, 194, 195, 196, 197, 223 harrisella, 196 tenella, 196, 197 Brasilocaenis, 194, 195, 196, 197, 198, 199, 200, 202 intermedia, 197, 198, 199, 200 irmleri, 197, 198, 199, 200, 201, 202

mendesi, 197, 198, 199, 201 puthzi, 197, 198, 199, 200, 201 renata, 197, 198, 199, 200, 201 septentrionalis, 197, 198, 199, 202 Caenidae, 18, 20, 28, 33, 44, 45, 46, 49, 50, 194 Caenini, 194 Caenis, 46, 194, 195, 196, 197, 202, 203, 204, 206, 209, 212, 214, 219 albata, 221 argentina, 203, 205, 206, 208, 210, 214, 217, 218, 219 axillata, 221 burmeisteri, 203, 204, 207, 211 candelata, 203, 205, 207, 211 chamie, 203, 204, 205, 206, 208, 210, 211, 219 chilensis, 221 cigana, 203, 203, 204, 205, 206, 208, 213 cuniana, 203, 204, 206, 207, 208, 213, 214, 219 dominguezi, 205, 207, 209, 215 fittkaui, 202, 205, 206, 207, 209, 210, 211, 214, 216, 219 gonseri, 205, 207, 209, 216 grimi, 205, 208, 209, 216 ludicra, 205, 206, 208, 209, 210, 214, 217, 219 macrura, 202 nemoralis, 217 nigella, 221 panamensis, 211 pflugfelderi, 205, 206, 208, 209, 210, 211, 218 219 plaumanni, 205, 206, 207, 210, 212, 214, 218, 219 pseudamica, 204, 207, 210, 212 quatipuruica, 206, 208, 212, 220 reissi, 202, 206, 208, 212, 220 sigillata, 206, 208, 212, 220 Callibaetis, 11, 24, 57, 106, 107, 108, 110, 111, 114 abundans, 118 camposi, 107, 108, 110, 112 dominguezi, 107, 109, 110, 112 fasciatus, 107, 109, 110, 113, 116 gonzalezi, 107, 108, 110, 113 gregarius, 108, 110, 113 guttatus, 106, 109, 110, 111, 112, 113, 114, 115 jocosus, 109, 111, 112, 115 nigrivenosus, 108, 110, 115

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pictus, 106 pollens, 108, 110, 111, 116 radiatus, 24, 106, 107, 109, 110, 112, 114, 116 sellacki, 24, 109, 110, 111, 116 viviparus, 109, 110, 111, 117 willineri, 24, 106, 109, 111, 112, 117 zonalis, 109, 111, 112, 117 Camelobaetidius, 20, 58, 62, 118, 119, 120, 124, 125, 127, 129, 130, 132, 134, 136, 138, 139, 140 alcyoneus, 120, 124, 127 anubis, 119, 120, 122, 123, 124, 126, 127 apis, 120, 125, 126, 128 billi, 120, 124, 125, 126, 128 cayumba, 124, 127, 128, 130 coveloae, 120, 124, 129 dryops, 120, 124, 129 edmundsi, 124, 127, 129, 130 huarpe, 122, 127, 129, 130 ipaye, 120, 124, 131, 132 janae, 122, 126, 131, 132 leentvaari, 118, 120, 124, 131 mantis, 120, 124, 133 mathuriae, 122, 126, 133, 134 matilei, 122, 127, 133, 134 ortizi, 122, 127, 134, 135 patricki, 122, 126, 135, 136 penai, 119, 120, 135, 136 phaedrus, 119, 120, 137, 138 serapis, 122, 126, 137 suapi, 119, 120, 122, 124, 127, 137, 138 tantillus, 120, 124, 139 tuberosus, 120, 126, 139, 140 yacutinga, 122, 126, 139, 140 Campsurinae, 41, 46, 557, 562 Campsurus, 11, 20, 502, 506, 557, 558, 562, 563, 565, 568, 571, 574, 581,582 albicans, 562, 563, 566 albifilum, 563, 565, 566 argentinus, 563, 565, 566 assimilis, 563, 565, 567 brasiliensis, 563, 565, 567 burmeisteri, 563, 565, 567, 570 claudus, 563, 567 corumbanus, 563, 569

cuyuniensis, 563, 565, 569, 570, 573 dallasi, 563, 565, 569 dorsalis, 563, 567, 569, 570 duplicatus, 563, 565, 570 essequibo, 563, 565, 570, 573 evanidus, 563, 565, 570 holmbergii, 563, 580 indivisus, 563, 565, 572 jorgenseni, 563, 568, 572, 578 juradinus, 563, 568, 572 latipennis, 563, 568, 572 litaninensis, 563, 568, 573 longicauda, 563, 568, 573 lucidus, 563, 568, 573, 577 mahunkai, 563, 568, 575 major, 563, 568, 575 melanocephalus, 563, 568, 575 meyeri, 563, 568, 576 mutilus, 563, 576 nappii, 563, 581 notatus, 563, 568, 576 pallidus, 563, 568, 577 paranensis, 563, 571, 577 paraquarius, 577 pedicellarius, 563, 571, 573, 577 pfeifferi, 563, 571, 578 quadridentatus, 563, 571, 578 scutellaris, 563, 571, 572, 578 segnis, 563, 569, 571, 579, 580 striatus, 563, 579 truncatus, 563, 571, 579 ulmeri, 563, 571, 579, 580 violaceus, 563, 564, 565, 571, 576, 580 wappaei, 563, 581 zikani, 563, 571, 580 Campylocia, 234, 235, 236, 237, 594 ampla, 235, 238 anceps, 235, 236, 237, 238 bocainensis, 236, 237, 238, 239 dochmia, 235, 236, 237, 239 Cercobrachys, 194, 195, 222, 223 colombianus, 222, 223 etowah, 222 peruanicus, 222, 223

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Chane, 58, 59, 61, 62, 141 baure, 141, 142, 143 Chaquihua, 46, 52, 53, 54 bullocki, 54 penai, 53, 54, 597, 598, 600 Chiloporter, 52, 54, 55 eatoni, 4, 54, 55, 56, 598, 599, 600 Chiloporterinae, 52, 54, 598, 599, 600 Chironomidae, 422 Choroterpes, 335, 337, 343, 345, 357, 511 (Choroterpes) bilineata, 511 albiannulata, 357 atramentum, 511 emersoni, 383, 386, 449, 450 picteti, 357 sp., 357 Chromarcyinae, 538 Cloeodes, 58, 59, 61, 62, 143, 144, 145, 146, 149, 152, 153 anduzei, 144, 145, 146, 148 auwe, 144, 146, 148, 152 aymara, 144, 145, 146, 148 binocularis, 145, 148 hydation, 144, 145, 146, 149, 152 irvingi, 144, 146, 149 jaragua, 146, 149, 152 maculipes, 143 nocturnus, 149 penai, 144, 145, 146, 150, 151 redactus, 144, 146, 151 stelzneri, 151 turbinops, 145, 151 venezuelensis, 144, 145, 146, 153 Coloburiscidae, 20, 28, 44, 45, 46, 49, 50, 224, 600 Coloburiscoides, 224 Coloburiscus, 224, 225 Colocrinae, 538 Colocrurinae, 538 Colocrus, 538 Coryphoridae, 44, 46, 47, 49, 51, 227, 594 Coryphorus, 20, 44, 46, 47, 48, 51, 228, 594 aquilus, 227, 228, 229, 594 Cryptonympha, 64, 153, 154, 155

copiosa, 153, 154 dasilvai, 153, 154, 155 Dactylobaetis, 118 Dactylophlebia, 337, 345, 358 carnulenta, 358, 359, 411 Deceptiviosa, 74, 76 Deleatidium, 421, 422, 429, 431 biobionicum, 426 chiloeense, 426 diguillinum, 428 illapeli, 429 laminatum, 429 penai, 426 vittatum, 524 Demoulinellus, 335, 338, 343, 347, 359, 361 coloratus, 359, 360, 361, 362 Dictyosiphlon. See Murphyella Eatonica, 230 Eatonigenia, 230 Ecuaphlebia, 335, 338, 343, 346, 362, 363 rumignaui, 362, 363, 364 sp. A, 363, 364, 365, 366 Epeorus, 27 Ephemera, 230, 510, 566 limbata, 230 Ephemerellidae, 533 Ephemeridae, 43, 45, 46, 48, 50, 230 Ephoron, 46, 557 umbrata, 557 umbratum, 315 Epiphrades, 296, 298, 307, 308 Euthyplocia, 46, 234, 235, 238, 239, 240, 242, 243 haenschi, 239, 240, 241 hecuba, 239, 240, 241 inaccessibile, 243 Euthyplociidae, 28, 43, 45, 46, 48, 50, 234, 594 Euthyplociinae, 234, 235 Exeuthyplocia, 234 Exeuthyplociinae, 234 Fallceon, 58, 60, 61, 63, 64, 156 inops, 156 murphyae, 156 yaro, 156

ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America

Farrodes, 333, 338, 341, 346, 366, 367, 370, 371, 374, 400, 483 bimaculatus, 367, 370 caribbianus, 367, 368, 369, 370, 372, 374 carioca, 368, 369, 370, 372, 373 cururensis, 379 hyalinus, 366, 373 iguazuanus, 368, 369, 372, 373 longispinus, 368, 369, 370, 373 maculatus, 367, 368, 369, 370, 372, 375 ochraceous, 369, 370, 371, 375 pakitza, 369, 370, 371, 375 roundsi, 367, 370, 371, 376 savagei, 369, 370, 371, 374, 376 tepui, 369, 371, 372, 376 xingu, 369, 371, 372, 377 yungaensis, 369, 371, 372, 374, 377, 595 Fittkaulus, 332, 336, 339, 343, 377, 378, 379, 380, 381, 511 cuiabae, 378, 379, 380 cururuensis, 378, 380 maculatus, 377, 378, 379,380, 381 Fittkauneuria, 538, 539, 540, 541 adusta, 540, 541, 542 carina, 540, 541, 542 Gonserellus, 338, 347, 381, 382, atopus, 381, 382, 383 Guajirolus, 58, 59, 61, 62, 156,157, 158 ektrapeloglossa, 156, 157, 158 nanus, 156 queremba, 156, 157, 158, 159 Hagenulopsis, 332, 339, 356, 383, 384, 385 diptera, 383, 384, 385 minuta, 383, 384, 385, 386 sp., 605 Hagenulus, 334, 342, 386, 387 caligatus, 386 marshali, 386, 387 sp., 387 Haplohyphes, 245, 246, 248, 249, 255, 256, 257, 261 aquilonius, 255, 256, 257, 258, 261 baritu, 257, 259 dominguezi, 255, 256, 257, 259 huallaga, 255, 256, 257, 260

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mithras, 255, 256, 257, 258, 260 yanahuicsa, 255, 256, 257, 258, 260, 261 Hapsiphlebia, 332, 337, 340, 345, 387, 388, 389 anastomosis, 387, 388, 389 Harpagobaetis, 59, 62, 159 gulosus, 159, 160, 601 Heptagenia, 224 Heptageniidae, 27, 28 Hermanella, 334, 336, 342, 344, 390, 391, 392, 393, 396, 404, 409, 450, 463, 513, 514 (Hermanellopsis), 390, 398 (Hermanellopsis) incertans, 390, 400 (s.s.) thelma, 390 costalis, 395 froehlichi, 391, 392, 393, 394, 395 grandis, 391, 392, 393, 395, 595 guttata, 391, 392, 393, 396 incertans, 390, 398, 400 maculipennis, 391, 392, 393, 396, 397 thelma, 390, 391, 392, 393, 396, 397 Hermanellopsis, 334, 342, 366, 390, 398, 399 arsia, 398, 399 incertans, 398, 399, 400 Hexagenia, 46, 230 (Hexagenia), 230 (Pseudeatonica), 43, 45, 48, 50, 230, 231 (Pseudeatonica) albivitta, 231, 232, 233 (Pseudeatonica) callineura, 231, 232 (Pseudeatonica) mexicana, 231, 233 Homoeoneuria, 538, 542, 543, 544 (Homoeoneuria) cahabensis, 544 (Notochora), 538, 539, 540, 543 (Notochora) fittkaui, 543, 544 salviniae, 542 Homoleptohyphes, 296, 298 Homothraulus, 333, 338, 341, 346, 366, 367, 375, 400, 401, 402, 403, 463, 483, 486 larensis, 400, 402 lucretiae, 400, 401, 402 misionensis, 400, 401, 402, 403 Hydrosmilodon, 334, 336, 341, 344, 404, 405, 406, 513 gilliesae, 404, 405, 406 mikei, 404, 405, 406, 407 saltensis, 404, 405, 406, 407

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Hylister, 333, 336, 341, 344, 404, 407 plaumanni, 407, 408, 409 Ichthybotus, 230 Iguaira, 59, 62, 161 poranga, 161, 162 Isonychiidae, 28 Lachlania, 20, 46, 538, 539, 540, 545, 546 boanovae, 545, 546 cacautana, 545, 547, 549 dencyanna, 545 dominguezi, 546, 547, 548 garciai, 545, 547 pallipes, 549 radai, 545, 549 santosi, 545, 549 Leentvaaria, 336, 344, 409, 410 palpalis, 409, 411, 595 Leptohyphes, 244, 245, 246, 248, 249, 250, 253, 260, 262, 263, 264, 266, 271, 274, 298, 325, 602 carinus, 265, 267, 269, 274 cornutus, 263, 264, 265, 266, 269, 274 ecuador, 265, 267, 269, 271, 274, 277 eximius, 262, 263, 264, 266, 268, 270, 271, 274 illiesi, 266, 267, 270, 271, 274 invictus, 265, 267, 272, 274 jodiannae, 265, 267, 271, 272, 274 liniti, 265, 267, 272 maculatus, 263, 265, 266, 268, 271, 272, 273, 274 mollipes, 263, 264, 266, 273 nigripunctum, 264, 266, 273 peterseni, 263, 265, 266, 273, 276 petersi, 263, 264, 265, 266, 267, 271, 274, 275 plaumanni, 263, 265, 266, 267, 273, 274, 275 populus, 265, 267, 275 setosus, 263, 264, 265, 266, 267, 271, 274, 276 tacajalo, 265, 267, 276 tuberculatus, 265, 267, 271, 274, 276, 277 Leptohyphidae, 18, 20, 28, 44, 46, 47, 51, 194, 227, 244, 247, 253, 316, 325, 594, 602 Leptohyphinae, 244 Leptohyphodes, 20, 244, 245, 246, 248, 249, 277 inanis, 277, 278, 279, 280 Leptophlebia, 331, 431

Leptophlebiidae, 6, 20, 28, 44, 47, 49, 51, 57, 331, 358, 367, 487, 533, 591, 595, 603, 604, 605 Leptophlebiini, 331 Litobrancha, 230 Longinella, 235 Lumahyphes, 245, 247, 248, 250, 279, 281, 282, 284 guacra, 279, 281, 282, 283, 284 pijcha, 279, 281, 282, 283, 284 yagua, 279, 281, 282, 283, 284, 285 Machadorythinae, 227 Machadorythus, 227 Magallanella, 336, 344, 411 flinti, 411, 412 Massartella, 20, 333, 337, 340, 345, 412, 413, 414, 415, 416 alegrettae, 413, 414, 415, 417 brieni, 412, 413, 414, 415, 416, 418 devani, 415, 416, 417, 418 sp., 418 venezuelensis,418, 413, 414, 415, 416, 417, 418 Massartellopsis, 333, 338, 340, 346, 418, 422 irarrazavali, 418, 419, 420, 421 Mayobaetis, 58, 59, 61, 63, 100, 161, 163, 164 ellenae, 161, 163, 164 sp., 163, 164, 606 Melanemerella, 20, 44, 47, 49, 51, 533 brasiliana, 533, 534, 535, 594 Melanemerellidae, 44, 47, 49, 51, 533, 594 Melanemerellinae, 533 Meridialaris, 334, 338, 342, 346, 419, 421, 422, 424, 425, 430, 431, 432 biobionica, 423, 424, 425, 426 chiloeensis, 423, 424, 425, 426, 428, 429, 430 diguillina, 423, 424, 425, 426, 428, 429, 430 inflata, 423, 424, 425, 429 laminata, 423, 424, 425, 426, 427, 429, 430 lestagei, 422, 431 patagonica, 422, 431 spina, 423, 426, 430, 431 tintinnabula, 423, 424, 425, 426, 430 Mesoplocia, 234, 235, 242, 243 intermedia, 242, 243 Metamonius, 45, 46, 47, 51, 224, 536 anceps, 536, 537

ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America

Microphlebia, 334, 337, 342, 345, 432, 433, 434, 435 pallida, 433, 434, 435 surinamensis, 433, 434, 435 Mirawara, 52 Miroculis, 334, 337, 342, 345, 436, 437, 439, 441, 443, 446, 448 (Atroari), 438, 440, 442 (Atroari) amazonicus, 437, 438, 439, 440, 441, 442, 443 (Atroari) colombiensis, 439, 442, 443, 446, 447 (Atroari) duckensis, 438, 439, 440, 441, 442, 443, 446 (Atroari) sp., 591 (Miroculis), 436, 439, 444 (Miroculis) bicoloratus, 438, 440, 441, 445 (Miroculis) brasiliaensis, 439, 442, 445, 446 (Miroculis) fittkaui, 437, 438, 439, 440, 441, 442, 445, 446 (Miroculis) marauiae, 437, 438, 439, 440, 441, 442, 445, 446 (Miroculis) nebulosus, 438, 439, 440, 441, 442, 447 (Miroculis) rossi, 437, 438, 440, 441, 447 (Ommaethus), 438, 440, 447 (Ommaethus) froehlichi, 438, 440, 441, 448 (Ommaethus) mourei, 437, 438, 439, 440, 441, 442, 444, 446, 447, 448 (Ommaethus) sp., 591 (Yaruma), 448 (Yaruma) sp., 441 (Yaruma) wandae, 438, 439, 448, 449 sp., 6, 605 Miroculitus, 334, 342, 449 emersoni, 449, 450 Moribaetis, 57, 59, 61, 63, 161, 165 aneto, 165 comes, 165 socius, 165 Murphyella, 20, 44, 45, 46, 49, 50, 224 needhami, 224, 225, 226, 600 Nanomis, 60, 64, 165 galera, 165, 166, 167 Needhamella, 334, 336, 342, 344, 390, 404, 450, 451, 513 ehrhardti, 450, 451, 452

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Neophlebia, 545, 547 Nesameletidae, 28, 45, 46, 47, 49, 51, 536 Nesameletus, 536 Nothofagus, 536 Nousia, 333, 338, 340, 347, 453, 454, 455, 460 bella, 454, 455, 456, 458, 460 crena, 454, 455, 456, 458, 459, 460 delicata, 453, 454, 455, 456, 457, 458, 459, 460, 595 grandis, 454, 455, 456, 461 maculata, 454, 455, 456, 458, 460, 461, 462 minor, 454, 455, 456, 460, 462 Noya, 545, 547, 549 garciai, 547, 549 pallipes, 549 Oligoneuria, 538, 539, 550, 551 anomala, 550, 551, 552 Oligoneuriidae, 20, 28, 43, 46, 47, 51, 224, 538, 550, 551 Oligoneurinae, 538 Oligoneurioides, 538, 539, 551, 552 amazonicus, 551, 552 Oniscigaster, 555 Oniscigastridae, 24, 28, 44, 45, 46, 49, 50, 555, 592, 593 Oxycypha, 72 Palingenia, 11, 499, 566 albicans, 560, 562, 566 atrostoma, 587 curta, 560, 561 Palingeniidae, 11, 23 Paracloeodes, 58, 59, 61, 62, 167, 168, 169, 171 abditus, 167 binodulus, 168, 169, 170 eurybranchus, 167, 168, 169, 170 ibicui, 168, 170, 171 leptobranchus, 168, 170, 171, 172 minutus, 167 Paramaka, 334, 336, 341, 344, 463 convexa, 463, 464 Penaphlebia, 332, 338, 340, 346, 465, 466, 467, 468, 473 (Megalophlebia), 465, 467, 468, 469, 470, 471

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(Megalophlebia) vinosa, 466, 467, 468, 469, 471, 473 (Penaphlebia), 465, 467, 468, 469, 471 (Penaphlebia) barriai, 466, 467, 468, 469, 472, 473 (Penaphlebia) chilensis, 466, 467, 468, 469, 470, 472, 473, 474 (Penaphlebia) exigua, 466, 467, 468, 469, 473, 474 (Penaphlebia) flavidula, 466, 467, 468, 469, 473, 474, 475, 603 (Penaphlebia) fulvipes, 466, 467, 468, 469, 470, 475 anastasii, 476 athanasii, 476 discolor, 476 fenestrata, 476 hyalina, 476 rofucensis, 477 valdiviae, 477 Perissophlebia, 477 Perissophlebiodes, 332, 339, 347, 356, 477 flinti, 478, 479 Polymitarcyidae, 11, 20, 28, 43, 45, 46, 48, 50, 557 Polymitarcyinae, 46, 557 Polyplocia, 234, 235 Povilla, 557, 558 Prebaetodes, 59, 63, 172 sitesi, 172, 173, 174 Proboscidoplocia, 234 Promirara, 52 Prosopistomatidae, 30 Rhigotopus, 333, 337, 340, 345, 479 andinensis, 479, 480, 481 Rivudiva, 60, 63, 174, 176 minantenna, 174, 176 trichobasis, 174, 175, 176 Secochela, 335, 338, 343, 346, 481 illiesi, 481, 482, 483 Simothraulopsis, 333, 338, 341, 346, 367, 400, 483 demerara, 483, 484, 485, 486 surinamensis, 483, 486 Siphlonella, 20, 44, 45, 46, 49, 50, 555, 556 guttata, 555, 556 ventilans, 24, 555, 556, 592, 593

Siphlurus anceps, 536 Spaniophlebia, 538, 539, 540, 549, 550, 551, 553 assimilis, 553 escomeli, 553, 554 trailiae, 553, 554 Spiritiops, 59, 62, 177 silvudus, 177, 178 Symbiocladius, 422, 461 wygodzinski, 419, 428, 431 Tasmanophlebia, 555 Terpides, 332, 335, 339, 343, 486, 487, 488, 511 guyanensis, 486, 487 jessiae, 487, 488 sp., 488 Thraulodes, 335, 338, 343, 346, 432, 489, 490, 491, 493, 497, 500, 505, 507, 510 bolivianus, 490, 492, 493, 495, 496, 505 bomplandi, 490, 492, 495, 498, 501 cochunaensis, 490, 492, 493, 495, 496, 498, 505 colombiae, 490, 491, 493, 499 consortis, 490, 491, 493, 494, 496, 498, 499, 503, 504, 505 daidaleus, 492, 493, 495, 496, 497, 498, 501 flinti, 492, 495, 497, 498, 501 furficulus, 492, 495, 497, 498, 501 guanare, 492, 495, 497, 502, 506 itatiajanus, 491, 493, 494, 496, 497, 502 laetus, 489, 493, 495, 497, 502, 503, 508 limbatus, 504 liminaris, 490, 491, 493, 494, 496, 497, 504, 505, 604 marreroi, 492, 494, 497, 502, 504 mucuy, 492, 494, 497, 506 osiris, 492, 494, 497, 506 papilionis, 491, 494, 497, 506 paysandensis, 493, 496, 500, 507 regulus, 491, 493, 500, 507 schlingeri, 491, 494, 500, 507 sp., 591 subfasciatus, 508 telegraphicus, 491, 494, 500, 508 traverae, 493, 496, 500, 508 trijunctus, 500, 509

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ulmeri, 492, 495, 500, 509 venezuelana, 491, 494, 500, 509 vitripennis, 510 Thraulus, 451, 463, 483, 503, 510 bellus, 510 caribbianus, 372 convexus, 463 demerara, 483, 486 laetus, 489, 502 maculipennis, 397, 451, 513 misionensis, 375, 400 primanus, 404 siewertii, 510 vogleri, 510 Tikuna, 332, 336, 340, 343, 511 atramentum, 512, 604 bilineata, 511 Tomedontus, 59, 63, 177 primus, 177, 179, 180 Tortopus, 46, 557, 558, 562, 567, 570, 581, 582, 583, 584, 606 bruchianus, 582, 583, 584 harrisi, 582, 583, 584, 585 igaranus, 581, 582, 583, 585 obscuripennis, 582, 583, 584, 585 parishi, 585 sarae, 582, 583, 584, 586 sp., 584, 606 unguiculatus, 582, 583, 584, 586 zottai, 582, 583, 584, 586 Traverella, 336, 344, 390, 397, 400, 404, 513, 514, 515, 518 (Traverella), 514, 516 (Traverella) albertana, 515, 516 (Traverella) bradleyi, 514, 515, 516 (Traverella) calingastensis, 513, 514, 515, 517, 518, 519, 596 (Traverella) montium, 514 515, 516 (Traverella) valdemari, 513, 514, 515, 518 (Zonda), 334, 341, 513, 514, 519 (Zonda) calingastensis, 513, 514, 515, 517, 518, 519, 596 Traverhyphes, 246, 247, 248, 250, 285, 286, 287, 288, 293, 297

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(Byrsahyphes), 293 (Byrsahyphes) nanus, 250, 286, 288, 289, 290, 293, 294, 295, 297 (Byrsahyphes) yuqui, 286, 288, 289, 294 (Mocoihyphes), 294 (Mocoihyphes) edmundsi, 250, 286, 288, 289, 290, 293, 295, 296, 297 (Mocoihyphes) yuati, 286, 288, 289, 290, 293, 295, 296, 297 (Traverhyphes), 290 (Traverhyphes) chiquitano, 286, 287, 288, 290 (Traverhyphes) indicator, 286, 287, 288, 290, 291, 292, 293, 295, 296, 297 (Traverhyphes) pirai, 285, 286, 287, 288, 292 Tricoryhyphes, 296, 298, 307, 312 Tricorythidae, 227, 244, 533 Tricorythodes, 46, 204, 244, 245, 246, 247, 249, 277, 296, 298, 299, 300, 302, 308, 309 arequita, 299, 300, 301, 302, 305, 306, 309, 310 australis, 245, 247, 298, 299, 300, 302, 307 barbus, 301, 305, 306, 307, 309, 312 bullus, 299, 300, 301, 302, 304, 305, 307, 308, 312 cristatus, 301, 304, 305, 307, 308 hiemalis, 299, 300, 302, 305, 306, 308, 309, 310 lichyi, 310 mirca, 299, 300, 301, 302, 305, 306, 310 nicholsae, 301, 304, 310, 312 ocellus, 298, 299, 300, 301, 302, 305, 306, 312 popayanicus, 298, 299, 300, 301, 303, 304, 309, 310, 312, 314, 594 quizeri, 299, 300, 302, 306, 309, 313 santarita, 299, 300, 301, 302, 304, 305, 309, 311, 312, 313, 314, 325 sp., 298, 502, 506 yura, 299, 300, 301, 304, 309, 314 zunigae, 299, 300, 301, 302, 304, 305, 309, 314 Tricorythodinae, 228, 244 Tricorythopsis, 244, 245, 246, 247, 250, 315, 316, 317, 318, 319, 322, 324 artigas, 315, 316, 317, 318, 319, 320, 321, 323, 324 chiriguano, 316, 317, 318, 319, 320, 321, 322, 324

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gibbus, 316, 317, 318, 319, 320, 321, 322, 323, 324, 602 minimus, 250, 316, 317, 318, 319, 320, 321, 322, 324, 594 sigillatus, 316, 317, 318, 323 undulatus, 316, 317, 318, 319, 320, 323 volsellus, 316, 317, 318, 323 yacutinga, 315, 316, 317, 319, 320, 324 Tupiara, 58, 60, 62, 64, 180 ibirapitanga, 180, 181, 182 Ulmeritoides, 335, 337, 342, 345, 519, 520, 521, 522, 525, 528, 532 flavopedes, 519, 521, 522, 524 guanacaste, 520 haarupi, 521, 522, 524, 526 huitoto, 521, 522, 526 luteotinctus, 520, 526 misionensis, 520, 521, 522, 523, 525, 526 oepa, 527 patagiatus, 527 spinulipenis, 520, 521, 522, 527, 528 tifferae, 520, 525, 526 uruguayensis, 519, 521, 522, 527, 528

Ulmeritus, 335, 337, 342, 345, 400, 504, 508, 519, 524, 527, 528, 529, 530, 531, 532 balteatus, 528, 529, 530, 531 carbonelli, 528, 529, 530, 531, 532, 595 saopaulensis, 529, 531, 532 Vacupernius, 244, 246, 247, 248, 249, 325 Varipes, 58, 60, 61, 63, 182, 183, 184, 187 cajuato, 184, 185, 186 helenae, 182, 184, 186, 187 lasiobrachius, 182, 184, 186, 187 minutus, 184, 187, 188, 601 singuil, 183, 184, 188, 189 Waltzoyphius, 60, 64, 188, 190, 191 fasciatus, 188, 190, 191 roberti, 189, 190, 191, 192 sp., 191 Yaurina, 245, 247, 248, 250, 252, 325, 326, 327, 330 mota, 325, 326, 327, 330 ralla, 250, 325, 329 yapa, 325, 326, 327, 329 yuta, 325, 326, 327, 328, 329 Zelusia, 60, 63, 192 principalis, 192, 193

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13 ADDENDUM

*

FAMILY BAETIDAE Camelobaetidius hamadae SALLES & SERRÃO, 2005 Camelobaetidius francischettii SALLES et al., 2005 Camelobaetidius lassance SALLES & SERRÃO, 2005 Camelobaetidius maranhensis SALLES & SERRÃO, 2005 FAMILY LEPTOHYPHIDAE Macunahyphes australis (BANKS, 1913) in DIAS et al., 2005 (=Tricorythodes australis) Tricorythopsis araponga DIAS & SALLES, 2005 Tricorythopsis baptistai DIAS & SALLES, 2005 Tricorythopsis pseudogibbus DIAS & SALLES, 2005 FAMILY LEPTOPHLEBIIDAE Lisetta ernsti THOMAS et al., 2005 Paramaka antonii SARTORI, 2005 Thraulodes basimaculatus GIORDANO & DOMÍNGUEZ, 2005

REFERENCES DIAS, L.G. & F.F. SALLES (2005): Three new species of Tricorythopsis (Ephemeroptera: Leptohyphidae) from southeastern Brazil. - Aquatic Insects 27: 235-241. DIAS L.G., SALLES, F.F. & C. MOLINERI (2005): Macunahyphes: A new genus for Tricorythodes australis (Ephemeroptera: Leptohyphidae). - Annales Limnologie 41: 195-201. GIORDANO, J.A. & E. DOMÍNGUEZ (2005): Thraulodes basimaculatus sp. n., a new species of mayfly from Bolivia: (Ephemeroptera: Leptophlebiidae: Atalophlebiinae). - Zootaxa 1040: 45-48. SALLES, F.F., ANDRADE, M.B. & E.R. DA-SILVA. (2005): Camelobaetidius francischettii: a new species of Baetidae (Ephemeroptera) from Brazil. - Zootaxa 1027: 46-53 SALLES, F.F. & J.E. SERRÃO (2005): The genus Camelobaetidius DEMOULIN (Ephemeroptera: Baetidae) in Brazil: new species, new records, and key for the identification of the species. - Annales Limnologie 41: 267-279.

*

New species published after book manuscript had been completed.

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SARTORI, M. (2005): A new species of the genus Paramaka SAVAGE & DOMÍNGUEZ, 1992 with some comments on related genera (Ephemeroptera, Leptophlebiidae, Atalophlebiinae). - Studies on Neotropical Fauna and Environment 40: 237-245. THOMAS, A., DOMINIQUE, Y. & K. ORTH (2005): Les Ephémères de la Guyane Française. 10. Description de Lisetta ernsti n. gen., n. sp. [Ephemeroptera, Leptophlebiidae]. - Ephemera 2003 (2005) 5: 77-84. Publications in press with new taxa from South America DIAS L.G. & F.F. SALLES (in press): A new species of Tricorythodes (Ephemeroptera: Leptohyphidae) from Minas Gerais, southeastern Brazil. - Neotropical Entomology 35: 56-58. MOLINERI, C. & M.C. ZÚÑIGA (in press): New species of Leptohyphidae (Insecta: Ephemeroptera) from Colombia with evidence of reproductive time segregation. - Studies on Neotropical Fauna & Environment 41: 139-151. SALLES, F.F. & C. MOLINERI (in press): Amanauhyphes saguassu: a new genus and species of Leptohyphidae (Ephemeroptera: Ephemerelloidea) from Northern Brazil. - Aquatic Insects 28: 1-12.