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ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 1 Aquatic Biodiversity in Latin America Biodiversidad Acuática en América Latina Volume 2 Ephemeroptera of South America Volumen 2 Ephemeroptera de América del Sur 2 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 3 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 4 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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) © 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. 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 Pensoft Publishers Geo Milev Str. 13a, Sofia 1111, Bulgaria pensoft@mbox.infotel.bg www.pensoft.net Printed in Bulgaria, June 2006 AQUATIC BIODIVERSITY IN LATIN AMERICA 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 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/Editores de la Serie: Joachim Adis, Jorge R. Arias, Guillermo Rueda-Delgado & Karl Matthias Wantzen 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. ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 5 To Bill and Jan Peters whose constant support and encouragement inspired us to write this book. 6 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.victoria- saxe.com) © 2006 CyanMagentaYellowBlack Adis Vol.2 Page 6 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 7 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: 978-954-642- 258-3, 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: 978-954-642- 259-0, 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 8 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. ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 9 Authors: PROF. DR. EDUARDO DOMÍNGUEZ CONICET-INSUE Facultad de Ciencias Naturales Universidad Nacional de Tucumán Miguel Lillo 205 4.000 Tucumán Argentina e-mail: mayfly@unt.edu.ar DR. CARLOS MOLINERI CONICET-INSUE Facultad de Ciencias Naturales Universidad Nacional de Tucumán Miguel Lillo 205 4.000 Tucumán Argentina e-mail: cmolineri@csnat.unt.edu.ar PROF. DR. MANUEL L. PESCADOR Laboratory of Aquatic Entomology Florida A&M University Tallahassee, Florida 32307-4100, USA e-mail:manuel.pescador@famu.edu PROF. DR. MICHAEL D. HUBBARD Laboratory of Aquatic Entomology Florida A&M University Tallahassee, Florida, 32307-4100 USA e-mail: michael.hubbard@famu.edu. ADDRESSES DR. CAROLINA NIETO CONICET-INSUE Facultad de Ciencias Naturales Universidad Nacional de Tucumán Miguel Lillo 205 4.000 Tucumán Argentina e-mail: carolinanieto@csnat.unt.edu.ar Series Editors: PROF. DR. JOACHIM ADIS Max-Planck-Institute for Limnology Tropical Ecology Working Group Postfach 165 24302 Plön, Germany e-mail: adis@mpil-ploen.mpg.de DR. JORGE R. ARIAS 5870 Colfax Avenue Alexandria, Virginia 22311, USA e-mail: jaria2@fairfaxcounty.gov 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: guillermo.ruedadelgado@utadeo.edu.co 10intercalaries generally attached to longitudinal veins (Figs. 123D, 175C); penes usually prominent and visible externally (Figs. 88D, 123H) ............................................................................................ 6 44 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 6 (5) Hind wings usually absent (Figs. 72A, 80A, 113A-C); if present (Figs. 90A-F, 93A- F), 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, 202B- C); 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 one- segmented (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 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 45 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 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 2(1) 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 3(2) 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 4(3) 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 5(2) 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 6(5) 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 7(6) 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-shapedfigure (Fig. 3I); gills III-VI with fringed margins and single lamella (Fig. 3M) ...................................... Caenidae 46 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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). A B C D E F G I J K H L M N O P Q R S T U ES T PP C OG T T T OGOG 1 2 3 4 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 47 – 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 48 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur CLAVES PARA LAS FAMILIAS SUDAMERICANAS Adultos 1 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 longitudi- nales presentes detrás de R1 (Figs. 8A, 72A, 80A, 113A, 123D, 212A) ........................ 2 2 (1) 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 3 (2) 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 4 (3) 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) 5 (2) 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 6 (5) 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, relativamentepequeña 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 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 49 7 (6) 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 8 (7) 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 9 (6) 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 10 (9) 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 11 (9) Vena CuP del ala anterior fuertemente recurvada (Fig. 78A) .................................... ............................................................................................. Coloburiscidae.....Murphyella – Vena CuP del ala anterior recta o levemente recurvada (Fig. 4A) ........................ 12 12 (11) 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 l3 (12) 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 50 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur Ninfas 1 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 2(1) 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. 3E- G, T-U) ................................................................................................................................. 3 – Branquias abdominales variables, márgenes de las branquias con o sin flecos; colmillos mandibulares ausentes ..................................................................................... 5 3(2) 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 4(3) 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 5(2) 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 6(5) 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 7(6) Branquias abdominales en los segmentos I-VI, las branquias del segmento I reducidas, filiformes (casi sobre coxas posteriores) (Fig. 75A); branquias opercula- res del segmento II grandes subcuadrangulares, con dos costillas unidas formando unafigura 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 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 51 8(5) 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 com- puestas 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 I- VII (Fig. 203D); ángulos posterolaterales de los segmentos abdominales expandi- dos 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 simultá- neamente; sin cámara branquial .......................................................... Leptohyphidae 52 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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 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 CLAVES PARA LOS GÉNEROS SUDAMERICANOS Adultos 1 Filamento terminal mucho más corto que los cercos; fórceps genitales tri- segmentados (Fig. 4C) ....................................................... Ameletopsinae.....Chaquihua – Filamento terminal subigual en longitud que los cercos; fórceps genitales tetra- segmentados (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 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 53 – 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. 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. A B C 54 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América delSur KEY TO CHAQUIHUA Adults 1 Wing membranes and veins orangish ............................................... Chaquihua bullocki – Wing membranes hyaline, veins brownish ........................................... Chaquihua penai CLAVE 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. ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 55 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. 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. DB A C 56 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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. Plate 6. Chiloporter eatoni, nymph. Fig. 6A: habitus. Drawing by A. Dupuy. A ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 57 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 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 2(1) Body with small reddish brown spots ............................................................ Callibaetis – Body without small spots ................................................................................................. 3 3(2) Fore wings with spots ........................................................................................ Moribaetis – Fore wings without spots (Fig. 62B) .............................................................................. 4 4(3) Hind wings absent ............................................................................................................. 5 – Hind wings present .......................................................................................................... 11 5(4) Basal segment of forceps with distomedial projection (Fig. 20C) .......................... 6 – Basal segment of forceps without projection (Fig. 47B) ........................................... 7 58 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 6(5) Postero lateral margin of styliger plate with small projection (Fig. 64C) ................ ......................................................................................................................................Varipes – Posterior margin of styliger plate without projection (Fig. 20C) ................. Baetodes 7(5) Segment II of forceps with distomedial projection (Fig. 47B) ................................ 8 – Segment II of forceps without projection ................................................................... 9 8(7) 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 9(7) Apical margin of styliger plate with a subtriangular projection (Fig. 16A) ............. .................................................................................................................................. Apobaetis – Styliger plate without projection (Fig. 42D) ............................................................... 10 10(9) Segment III of forceps short and rounded (Fig. 42D) ................... Cloeodes (in part) – Segment III of forceps elongated (Fig. 8B) ............................................. Americabaetis 11(4) 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 12(11) Forceps apparently four-segmented .....................................................................Fallceon – Forceps three-segmented (Fig. 54C) ............................................................................ 13 13(12) Hind wings with subquadrangular costal projection (Fig. 54F) .............. Paracloeodes – Hind wings with pointed costal projection (Fig. 30C) ............................................. 14 14(13) Segment III of forceps short and rounded (Fig. 42D) ............................................ 15 – Segment III of forceps elongate (Fig. 30E) ............................................................... 16 15(14) 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 16(14) Fore wings with anal margin rounded (Fig. 30A) ................................Camelobaetidius – Fore wings with anal margin relatively straight (Fig. 62B) ............................. Tupiara Nymphs 1 Tarsal claws spatulate (Fig. 32I), with 5 to 40 denticles .....................Camelobaetidius – Tarsal claws apically pointed (Fig. 19H) ........................................................................ 2 2(1) Gills on abdominal segments I-V ........................................................................ Baetodes – Gills on abdominal segments I-VI, I-VII or II-VII .................................................... 3 3(2) Abdominal gills with two lamellae (Figs. 29M-O) ....................................... Callibaetis – Abdominal gills with a single lamella (Fig. 9J) ............................................................. 4 4(3) Tarsal claws without denticles (Fig. 44J) ....................................................................... 5 – Tarsal claws with denticles (Figs. 9D, 14D) although these can be very small (Fig. 19H) ...................................................................................................................................... 6 5(4) Tarsal claws long (0.75 to 0.8 times length of tarsi) (Fig. 16I); tibiae without an arc of long setae (Fig. 16I) ....................................................................................... Apobaetis ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 59 – 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 6(4) 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 7(6) 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 8(7) 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 9(8) 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 10(8) 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 11(10) Tarsal claws with 2 rows of denticles (Fig. 60I) ............................................. Spiritiops – Tarsal claws with 1 row of denticles (Fig. 21K) ........................................................ 12 12(11) 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 13(12) Hind wing pads rudimentary if present; terminal filament– Gills present on abdominal segments I or II-VII; labrum variable but not as above .............................................................................................................................................. 15 1 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. 60 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 61 CLAVES PARA LOS GÉNEROS SUDAMERICANOS Adultos 1 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 2(1) Cuerpo con pequeñas manchas castaño-rojizas ........................................... Callibaetis – Cuerpo sin tales manchas ................................................................................................. 3 3(2) Alas anteriores con manchas ............................................................................ Moribaetis – Alas anteriores sin manchas (Fig. 62B) .......................................................................... 4 4(3) Alas posteriores ausentes .................................................................................................. 5 – Alas posteriores presentes .............................................................................................. 11 5(4) Segmento I de los fórceps con una proyección lateral (Fig. 20C) ........................... 6 – Segmento I de los fórceps sin proyección (Fig. 47B) ................................................. 7 6(5) 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 7(5) Segmento II de los fórceps con una proyección distomedial (Fig. 47B) ............... 8 – Segmento II de los fórceps sin proyección .................................................................. 9 8(7) 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 9(7) 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 10(9) Fórceps con segmento III corto y redondeado (Fig. 42D) ........ Cloeodes (en parte) – Fórceps con segmento III alargado (Fig. 8B) .......................................... Americabaetis 11(4) 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 12(11) Fórceps aparentemente tetra-segmentados ........................................................Fallceon – Fórceps tri-segmentados (Fig. 54C) ............................................................................. 13 13(12) Proyección costal del ala posterior subcuadrangular (Fig. 54F) .............. Paracloeodes – Proyección costal del ala posterior aguda (Fig. 30C) ............................................... 14 14(13) Fórceps con el segmento III corto y redondeado (Fig. 42D) ................................ 15 – Fórceps con el segmento III alargado (Fig. 30E) ..................................................... 16 15(14) 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 62 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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 Uñas tarsales espatuladas, con 5 a 40 dentículos (Fig. 32I) ...............Camelobaetidius – Uñas tarsales aguzadas hacia el ápice (Fig. 19H) ......................................................... 2 2(1) Branquias en los segmentos I-V solamente ....................................................... Baetodes – Branquias en los segmentos abdominales I-VI, I-VII ó II-VII ............................... 3 3(2) Branquias abdominales con dos láminas (Figs. 29M-O) ............................ Callibaetis – Branquias abdominales con una lámina (Fig. 9J) ........................................................ 4 4(3) Uñas tarsales sin dentículos (Fig. 44J) ........................................................................... 5 – Uñas tarsales con dentículos, aunque pueden ser pequeños (Fig. 19H) ................ 6 5(4) 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. 63C- D); 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 Cloeodes y Bernerius están en revisión y existen dudas respecto a la validez de éste último. 4 Paracloeodes e Iguaira están en revisión y existen dudas respecto a la validez de éste último, ambos géneros parecen indistinguibles. ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 63 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(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 64 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur – Labro dorsalmente sin setas aplanadas (Figs. 53A-B); palpos maxilares bi- segmentados (Fig. 53E) ................................................................................................... 23 23(22) Uñas tarsales con 2 hileras de dentículos (Fig. 63I) ........................................ Tupiara – Uñas tarsales con 1 hilera de dentículos (Fig. 53J) ................................................... 24 24(23) 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 25(21) 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 26(25) 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 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 65 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. 7A- I modified from LUGO-ORTIZ & MCCAFFERTY, 1995a. A D B C E F H GI D I III II SL L P P I TS 66 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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) Plate 8. Americabaetis alphus, male imago. Figs. 8A-B. 8A, fore wing; 8B, genitalia. (IV = intercalary veins). A B IV IV ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 67 with setae between prostheca and mola, incisors fusedapically, 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 Frontal keel absent ............................................................................................................ 2 – Frontal keel present ........................................................................................................... 4 2(1) 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 3(2) 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 4(1) 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 5(4) 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 CLAVE PARA LOS AMERICABAETIS SUDAMERICANOS Ninfas 1 Quilla frontal ausente ........................................................................................................ 2 – Quilla frontal presente ...................................................................................................... 4 2(1) 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 68 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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). A BE C D F I J G H K P P S S SLL III II I D ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 69 3(2) 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 4(1) 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 5(4) 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. 70 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur Taxonomy: This species was originally described from male and female imagos by ESBEN- PETERSEN (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) FrontalABLA 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: matthias.wantzen@uni- konstanz.de 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: Michel.sartori@vd.ch ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 11 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 12 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 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 13 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.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. ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 71 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. A C B D E F H I L G KJ SLL III II I D III II I SLL III II I III II I III II I SP SP 72 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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- BERGH (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. 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 ESBEN- PETERSEN (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 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 73 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 inconsequen- tial 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 speciesbelongs 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. 74 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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 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. A D B C E F IV IV IV IV ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 75 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. Plate 12. Andesiops sp., nymphal habitus. Figs. 12A-B. 12A, dorsal view; 12B, lateral view. Drawings by C. Molineri. A B 76 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur KEY TO SOUTH AMERICAN ANDESIOPS Nymphs 1 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 2(1) 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 CLAVE PARA LOS ANDESIOPS SUDAMERICANOS Ninfa 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 2(1) 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 inter- costal area with six crossveins; 2) hind wings that are small and narrow, without cross veins. Distribution: Chile. ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 77 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. DA CF B E H G J K I L P P T SP L SP L S D III II I III II I 78 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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 fromnymphs 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) ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 79 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. A B C E FG D J I H D S P LSL S D 80 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 81 Plate 15. Apobaetis sp. (undescribed). Figs. 15A-B, nymphal habitus: 15A, dorsal view; 15B, ventral view. Drawings by C. Molineri. A B 82 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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. A C E B D FGH I J P I S AS SLL SP P I S III II I ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 83 CLAVE PARA LOS APOBAETIS 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 galea- lacinia; 3) segment III of labial palp as in Fig. 17C; 4) color pattern as in Fig. 17D. Distribution: Brazil. 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 = shortsetae; Roman numbers = segment numbers). Figs. 17A-D modified from SALLES & LUGO-ORTIZ, 2002; 17E-17G from LUGO-ORTIZ & MCCAFFERTY, 1997. A B C D F GE SS III II I III II I 84 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 85 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 PARA LOS 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 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). A E B F C D G H SP III II I IV CP CPIV IV SP III II I S 86 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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 & MCCAFFER- TY, 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, ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 87 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. 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 A B C D G E H J I K F P P TS III II I III II I D SLL 88 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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). 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. A B D C E IV IV P P ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 89 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 Abdominal tubercles present on segments II-V .................................. Baetodes spinifer – Abdominal tubercles absent ............................................................................................. 2 2(1) 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 3(2) 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 Ocelli absent .......................................................................................... Baetodes itatiayanus – Ocelli present ...................................................................................................................... 2 2(1) Coxal gills present (Figs. 25F, L) ..................................................................................... 3 – Coxal gills absent (Fig. 21I) ............................................................................................ 15 3(2) One finger-like gill on each coxa (Fig. 25F) ................................................................. 4 – Two finger-like gills on each coxa (Fig. 25L) ............................................................... 6 4(3) Abdominal tubercles absent ......................................................... Baetodes sancticatarinae – Abdominal tubercles present (Figs. 22L, 23C) ............................................................ 5 90 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 5(4) 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 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 91 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 non- pectinate 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 Tubérculos abdominales presentes en los segmentos II-V ............... Baetodes spinifer – Tubérculos abdominales ausentes .................................................................................. 2 2(1) 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 3(2) 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 Ocelos ausentes .................................................................................... Baetodes itatiayanus – Ocelos presentes ................................................................................................................ 2 2(1) Branquias coxales presentes (Figs. 25F, L) ................................................................... 3 – Branquias coxales ausentes (Fig. 21I) ......................................................................... 15 3(2) Cada coxa con una branquia digitiforme (Fig. 25F) ................................................... 4 – Cada coxa con dos branquias digitiformes (Fig. 25L) ................................................ 6 4(3) Tubérculos abdominales ausentes .............................................. Baetodes sancticatarinae – Tubérculos abdominales presentes (Figs. 22L, 23C) .................................................. 5 92 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 5(4) 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 cinco- seis 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 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 93 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 94 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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. A B C D EF G H J K I L D PS SS III II I SL LP P ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 95 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 distin- guished 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 palpiwith 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. 96 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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. BA DC E F G K H M J L I T T T T CG IP III II PS ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 97 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 distin- guished from other species of the genus by the following combination of characters: 1) 98 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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 finger- like 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: ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 99 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. A B C D G F HE I T T CG CG TT 100 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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. ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 101 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. A D B C E G F H K J I L M P P CG T T T T PS II I III T 102 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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. ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 103 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. A C B ED F G H I J L K N M PS SL L CG CGIII II I T 104 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur Baetodes uruguai NIETO Baetodes uruguai NIETO, 2004a: 75. Taxonomy: Baetodes uruguai, known from male imagos and nymphs, can be distin- guished 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 8- 9 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 6- 7 short clavate setae, less than half the length of fine setae (Fig. 25M); 4) tarsal claws with 5- 6 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 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 105 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). A B C D J E G H I F A III II I C P P L SL 106 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 107 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. A B D C E F G K L I J H IV IV IV IV IV IV IV IV CP CP VV 108 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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 spine- like 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 SOUTH AMERICAN CALLIBAETIS Imagos5 1 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 2(1)14 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 15 1 INTRODUCTION 17 2 MORPHOLOGY 18 Nymph 18 Adult 20 Eggs 22 3 BIOLOGY 23 4 WORKING WITH SPECIMENS 25 5 HIGHER CLASSIFICATION 27 6 DISTRIBUTION 28 7 LIST OF FAMILIES, SUBFAMILIES, GENERA AND SPECIES 29 8 SYSTEMATICS 43 Keys to South American families 43 Claves para las familias sudamericanas 48 Family Ameletopsidae 52 Family Baetidae 57 Family Caenidae 194 Family Coloburiscidae 224 Family Coryphoridae 227 Table of Contents 16 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur Family Ephemeridae 230 Family Euthyplociidae 234 Family Leptohyphidae 244 Family Leptophlebiidae 331 Family Melanemerellidae 533 Family Nesameletidae 536 Family Oligoneuriidae 538 Family Oniscigastridae 555 Family Polymitarcyidae 557 9 COLOR PLATES 589 10 GUIDE TO CHANGES IN SCIENTIFIC NAMES 607 11 REFERENCES 611 12 SUBJECT INDEX 635 13 ADDENDUM 645 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 17 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). 18 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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 (foreWings in both sexes completely hyaline (Fig. 28G) .......................... Callibaetis pollens – Wings at least of females with dark bands ................................................................... 3 3(2) 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 5 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. ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 109 4(3) 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 5(1) 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 6(5) 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 7(6) 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 8(6) 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 9(8) 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 10(9) 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 Maxillary palpi shorter than galea-lacinia ........................................ Callibaetis willineri – Maxillary palpi longer than galea-lacinia (Fig. 29F) .................................................... 2 2(1) 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 110 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur CLAVES PARA LOS CALLIBAETIS SUDAMERICANOS Imagos6 1 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 2(1) Alas en ambos sexos completamente hialinas (Fig. 28G) ............... Callibaetis pollens – Alas por lo menos en las hembras con bandas oscura .............................................. 3 3(2) 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 4(3) 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 5(1) 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 6(5) 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 7(6) 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) ...................................................................Callibaetissellacki 8(6) 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 6 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. ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 111 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. C B D A E G H M L J I F O Q K N P IV IV IV IV IV 112 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 9(8) 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 10(9) 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 Palpos maxilares más cortos que la galea-lacinia ........................... Callibaetis willineri – Palpos maxilares más largos que la galea-lacinia (Fig. 29F) ...................................... 2 2(1) 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 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 113 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 guttatus NAVÁS, 1915a: 120; GILLIES, 1990: 25; DA-SILVA, 1991: 346. Callibaetis apicatus NAVÁS, 1917: 189. Callibaetis bruchius NAVÁS, 1920e: 55. Callibaetis zonatus NAVÁS, 1929: 224. 114 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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. A C B D EF K H L G J I M ON P Q R S P P T D III II I L SL P S P S ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 115 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 jocosa NAVÁS, 1912b: 195. Callibaetis stictogaster NAVÁS, 1915a: 121. Callibaetis jaffueli NAVÁS, 1918: 244. Callibaetis spegazzinus NAVÁS, 1920d: 36. Callibaetis rimatus NAVÁS, 1932a: 113. Callibaetis 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. 116 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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. ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 117 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 fromthe 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. 118 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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 & MCCAF- FERTY, 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 1.5- 2.5 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 two- segmented, 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. ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 119 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. A B C E FD G J K LI H CP III II I IV IV A 120 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur KEYS TO SOUTH AMERICAN CAMELOBAETIDIUS Male Imagos7 1 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 2(1) Genitalia: segment II of forceps strongly curved (Fig. 30E) ...... Camelobaetidius anubis – Genitalia: segment II of forceps not as above (Fig. 30I) .......................................... 3 3(2) 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 4(3) 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 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 2(1) Coxal gills although small present; prosternum with single, medial protuberance ........................................................................................................ Camelobaetidius leentvaari – Coxal gills absent; prosternum without protuberance ............................................... 3 3(2) 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 4(1) Coxal gills present (Fig. 32H) .......................................................................................... 5 – Coxal gills absent (Fig. 34M) ......................................................................................... 14 5(4) Terminal filament short, subequal in length to abdominal segment X (Fig. 33F) ...... 6 – Terminal filament long, subequal in length to cerci ................................................... 7 6(5) 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 7(5) Pronotum with medial pair of tubercles ................................ Camelobaetidius tuberosus – Pronotum without tubercles ............................................................................................ 8 7 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 distinguishthese species from the others. ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 121 Plate 31. Camelobaetidius phaedrus. Fig. 31A, nymphal habitus. Drawing by C. Molineri. A 122 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 8(7) Maxillae with segment II of palpi bifid (Figs. 35L-M) ............. Camelobaetidius janae – Maxillae with segment II of palpi simple (Fig. 32E) .................................................. 9 9(8) Tarsal claws with less than 10 denticles ...................................................................... 10 – Tarsal claws with more than 15 denticles ................................................................... 11 10(9) 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 11(9) 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 12(11) 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 13(11) 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 14(4) 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 15(14) 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 16(14) 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 17(16) 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 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 123 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. CBA D I E F G H J K L M S CG SL L P P III II I 124 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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 CAMELOBAETIDIUS SUDAMERICANOS Imagos Machos8 1 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 2(1) Genitalia: segmento II de los fórceps fuertemente arqueado (Fig. 30E) ................. ............................................................................................................. Camelobaetidius anubis – Genitalia: segmento II de los fórceps levemente arqueado (Fig. 30I) .................... 3 3(2) 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 4(3) 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 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 2(1) Branquias coxales, aunque pequeñas presentes; proesterno con una protuberancia medial ........................................................................................... Camelobaetidius leentvaari – Branquias coxales ausentes; proesterno sin protuberancia ....................................... 3 3(2) 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 8 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. ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 125 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. A C B E D F G H J K L M I CG CG III II I TF 126 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 4(1) Branquias coxales presentes (Fig. 32H) ......................................................................... 5 – Branquias coxales ausentes (Fig. 34M) ........................................................................ 14 5(4) 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 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 127 (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 16(14) 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 17(16) 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 18(16) 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 palpolabial con una fuerte proyección distomedial aguda como en la Fig. 37I; uñas tarsales con 30 o menos dentículos .......................................... 19 19(18) 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 128 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 129 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) trans- ferred 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 130 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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 clawI; 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. A C D F G H B E I J K L M N O P Q III II I III II I III II I ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 131 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 132 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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. A C D F G H B E I J K L M N O P Q R III II I III II I III II I T E P ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 133 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. 134 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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; 36L- P from DOMINIQUE & THOMAS, 2001. A C D F G H B E I J K L M N O P III II I III II I III II I ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 135 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. 136 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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. A C D F G H B E I J K L M III II I CG III II I ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 137 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. 138 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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. A C DF G H B E I JK L M N O P Q R III II I III II I IC P P IC CG ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 139 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 & MCCAFFERTYCamelobaetidius 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. 140 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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. A C D F G H B E I J K L M N O P Q RCG CG III II I III II I L SL ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 141 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 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. A C B IV IV 142 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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. 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. A C D F G H B E I J K III II I MPP S D SD S P ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 143 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 & LUGO- ORTIZ, 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 nearlegs), 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 19 Plate 1. Figs. 1A-B, nymph. 1A, habitus, dorsal view; 1B, mouth parts. A B ClawTarsus Gills Femur Tibia Cerci Terminal Filament Hind Leg Middle Leg Fore Leg Ocelli Labrum Antenna Incisor Prostheca Molar SuperlinguaLingua Incisor Prostheca Molar Galea-lacinia Stipes Glossa Paraglossa Palpi Palpi Clypeus Labrum Hypopharynx Left Mandible Labium Right Mandible Right MaxillaLeft Maxilla HeadMesonotum Pro- notum Fore Wing Pads Abdomen 20 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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, Polymitarcy- idae) 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 21 Plate 2. Figs. 2A-D, adult. 2A, lateral view; 2B, fore wing; 2C, hind wing; 2D, genitalia, ventral view. B A C D Hind Leg Middle Leg CoxaFrons Trochanter Tarsi Fore Leg Tibia Antenna Femur Ocelli Eye Thorax Bullae Costal Projection Fore Wing Abdomen Genitalia Caudal FilamentsHind Wing 10 1 2 3 4 5 6 7 8 9 R4+5 R1 R2 R3 C Sc MA1 MA2 MA MP1 IMP MP2 CuAMP R1 C Sc R2 R3R4+5A Rs CuP CuA MA 1MA 2 MP 1MP 2 Forceps Penes Subgenital Plate Forceps Socket C Sc Rs R A MACuACuP MP 22 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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 longitudi- nal 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 areamidline, 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- 144 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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 I- VII, 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 Hind wings present ........................................................................................................... 2 – Hind wings absent ............................................................................................................. 4 2(1) 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 3(2) Segment II of forceps with a basal constriction (Fig. 42K) ........... Cloeodes hydation – Segment II of forceps without a basal constriction (Fig. 42G) ................................. ........................................................................................................................ Cloeodes aymara 4(1) 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 Hind wing pads absent ...................................................................................................... 2 – Hind wing pads present .................................................................................................... 3 2(1) 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 3(1) 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 4(3) 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 9 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. ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 145 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. A C D F G H B E I J K L M III II I IV IV A1 C IV C S 146 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 5(4) 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 CLAVES PARA LOS CLOEODES SUDAMERICANOS Imagos Machos10 1 Alas posteriores presentes ................................................................................................ 2 – Alas posteriores ausentes .................................................................................................. 4 2(1) 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 3(2) 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 4(1) 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 Pterotecas posteriores ausentes ....................................................................................... 2 – Pterotecas posteriores presentes ..................................................................................... 3 2(1) 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 3(1) 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 4(3) 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 5(4) 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 10 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. ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 147 Plate 43. Cloeodes sp. Fig. 43A, nymphal habitus. Drawing by C. Molineri. A 148 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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. ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 149 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 150 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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). 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. A C D F G H B E I J KL M N III II I P P C L SL S AS LS ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 151 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 examina- tion 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 152 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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. A C DF G H B E I J III II I AS P P ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 153 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 & FRAN- CISCHETTI (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. 154 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur KEY TO SOUTH AMERICAN CRYPTONYMPHA Nymphs 1 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 thumb- like 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 CLAVE 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. ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 155 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. A C D F G H B E I J K L M N O P III II I D III II I P SL L TS PP S 156 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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. 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. A B IV IP IV P ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 157 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 emargin- ation, 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 TO SOUTH AMERICAN GUAJIROLUS Nymphs 1 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 CLAVE 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. 158 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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. A C D F G H B E I J K L M SLLIII II I SLL SS P P SD D ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 159 Guajirolus queremba NIETO Guajirolus queremba NIETO, 2003b: 65. Taxonomy: This species was described based on the nymph and female subimago. Guajirolus queremba canbe 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. 49L- M) with glossae subequal to paraglossae, both slender elongate with a row of long spine- like 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. 160 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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. A C D F G H B E I J K L MN O P III II I I S PS SS D PG G P P SL L ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 161 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 & MCCAFFER- TY (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). 162 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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 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). A C D F G H B E I J K L SL L III II I D I P I P ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 163 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. 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. A C D B IV A1 IV P 164 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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). A C D F G H B E I J K III II I SP SS SP D S MS S MS C P P SL L ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 165 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, MCCAFFER- TY (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. 166 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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). A C D F G H B E I J K L III II I S D P P ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 167 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). 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. A C D F B E I III II IV CP IV 168 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur Nymphal Characteristics: 1) Antennae variable, 2-4 times of head capsule; 2) labrum (Fig. 55A) widerthan 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 2(1) 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 3(2) 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 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 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 169 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. A C D F G H B E I J K L M N O P Q R III II I SL L I P I P SL L I P D T T 170 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 2(1) 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 3(2) 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, 2.7- 3.4 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 10- 12 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 distin- guished 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 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 171 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. A C D F G H B E I J III II I D SL L I P P I 172 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera deAmérica del Sur 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, 3.9- 4.1 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 galea- lacinia; 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. ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 173 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). A C D F G H B E I J K III II I SS D T L SL I P P 174 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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. 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). A C D B III II I T ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 175 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 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). A C D F G H B E I J K III II I D P I P 176 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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. ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 177 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. 178 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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). 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 A C D F G H B E I J K III II I C D SS SL L S TSP P ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 179 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. 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. 61A- J modified from LUGO-ORTIZ & MCCAFFERTY, 1995a. A C D F G H B E I J III II I SL L S P P S 180 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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. 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. A C D B IV A ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 181 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 partiallycleft 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 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). A C D F G H B E I J KL III II I D SL L P P 182 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 183 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 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. IP P A C B IV 184 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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 3(2) 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 4(2) Labrum: dorsal subapical setae as in Fig. 66G; hypopharynx: superlingua rounded (as in Fig. 65F), maxillae with palpi elongate, reaching beyond apex of galea- lacinia (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 CLAVE PARA LOS VARIPES SUDAMERICANOS Ninfas 1 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 2(1) 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) ..............................................................................................................of the egg chorion (Figs. 221C, E). ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 23 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. 24 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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 genera- tion 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). ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 25 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 battery- operated 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 the4 3(2) 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 4(2) 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 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 185 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. A C D F G H B E I J K L M N III II I L SL S I P P D TS TS 186 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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 four- five 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 distin- guished 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 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 187 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. A C D F G H B E I J K L M N O III II I D TS TS P D SL L 188 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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 distomedialnymphs were collected. In general, if the water 26 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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 40- 50X 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. ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 27 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, grantp@swosu.edu. 28 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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). Representa- tives 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 biogeo- graphic 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. ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 29 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 7 LIST OF FAMILIES, SUBFAMILIES, GENERA AND SPECIES 30 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur • 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 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 31 • 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 32 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur • 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 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 33 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 34 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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) ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 35 • 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 36 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur 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 37 • 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) 38 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur • 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 39 • 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) 40 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur • 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 41 • 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 42 ABLA Vol. 2 – Domínguez et al.: Ephemeroptera de América del Sur • 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) ABLA Vol. 2 – Domínguez et al.: Ephemeroptera of South America 43 8 SYSTEMATICS KEYS TO SOUTH AMERICAN FAMILIES Adults 1 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 2 (1) 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 3 (2) Middle and hind legs of male and all legs of female atrophied, nonfunctional .................................................................................................................. Polymitarcyidae – All legs of both sexes well-developed, functional ...................................................... 4 4(3) 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) 5(2) 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
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