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F.A.A. Lencioni

DAMSELFLIES OF BRAZIL AN ILLUSTRATED IDENTIFICATION GUIDE

SOUTHEAST REGION

1st edition

Jacareí - São Paulo Frederico Augusto de Atayde Lencioni

2017

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Damselflies of Brazil - An illustrated identification guide Southeast Region Copyright © 2017 Frederico A.A. Lencioni All rights reserved. No part of this book may be reproduced or utilized in any form or by any means, electronic or mechanical, including photocopying, recording, or by any information storage and retrieval system, without written permission of the author.

Dados Internacionais de Catalogação na Publicação (CIP) (Câmara Brasileira do Livro, SP, Brasil)

Lencioni, F.A.A., 1970- Damselflies of Brazil : an illustrated identification guide [livro eletrônico] : southeast region / F.A.A. Lencioni. -- 1. ed. -- Jacareí, SP : Ed. do Autor, 2017.

395.776 Kb ; PDF

Bibliografia ISBN: 978-85-905339-2-4

1. Brasil - Região Sudeste - Guia 2. Libélulas - Brasil - Identificação 3. Zoologia - Brasil I. Título.

17-09168 CDD-595.7330981

Índices para catálogo sistemático:

1. Brasil : Libélulas : Identificação : Entomologia : Zoologia 595.7330981 2. Libélulas : Identificação : Brasil : Entomologia : Zoologia 595.7330981 Photographs, illustrations, layout and text by Frederico A.A. Lencioni (except when noticed).

Abstract: This is the first of a three-volume work covering all the described species of damselflies (Zygoptera) of Brazil. The present volume includes all 159 species found in the Southeast region (69 non-Coenagrionidae + 90 Coenagrionidae). To facilitate the identification of species, we have been incorporating keys for families and genera, and diagnostic illustration to all species. The diagnostic illustration now includes high-resolution scans of habitus of approximately 90% of the species (some acetoned and some from live specimens), and 92 photos of live specimens (covering 74 species, representing almost 50% of the species). The publication data also includes the etymology of all species-group and genus-group names. The volume concludes with a glossary and a bibliography of more than 180 papers, cross-referenced with the book. Obs.: This book was designed to be viewed in "2-page" mode with the diagnostic illustrations (and publication data) on the left side of the screen and scans of the specimens on the right side, so several blank pages were inserted to make this happen. All the chapters and new topics also start on odd pages, and for that purpose, blank pages have been inserted.

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I dedicate this book to my wife Angela Schmidt Lourenço Rodrigues, for her company and dedication;

To my wonderful kids Nathalia and Frederico which are my contribution to a better and prettier world;

To my parents Frederico and Luiza that gave me a different vision of the world and to my brothers Eduardo and Eric;

To Angelo B.M. Machado, colleague, friend, and pioneer in the study of Odonata in Brazil.

To Bastiaan & Marianne Kiauta, two very good friends who incentivized me all the time.

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Preface

When I started with the idea of writing a book gathering all species of Zygoptera with distribution in Brazil, all the scientists with which I talked about the idea said I was crazy as the information was so sparse that it would be difficult to gather all of it, and even if I were able to get all the information, the volume would be too big to analyze and arrange in a didactic way.

It was hard, but after two long years of sleepless nights, organizing the data, I published two volumes, the first in 2005 with all Non-Coenagrionidae species and the second in 2006 with all Coenagrionidae species, and four appendixes (1 - Non-Coenagrionidae described after volume 1; 2 - All described larvae; 3 - Description of Mecistogaster asticta Selys, 1860 larvae, oviposition and larval behavior, and 4 - Damselflies pictures).

Twelve years have passed since the publication of the first part of the guide of Zygoptera of Brazil, and many papers on the South American Zygoptera fauna have been published in this period, with material ranging from the larval description, new species, and several generic revisions.

The first edition of this book was the first guide to the species of Odonata published on species of South America (von Ellenrieder & Garrison, 2007, IDF report 11), covering the largest and most biodiverse country on the South American continent.

Of course, on that first attempt to bring together data on 320 species for the first edition, there were some errors and omissions.

To gather data for the two books, we analyzed over 300 papers and books (over 12,000 pages), and I scanned more than 2,500 illustrations; the vast majority had to be cleaned and prepared to assemble over 400 figures.

The simple work of structuring a book to contain 320 species of Zygoptera, separating the data I understood to be the most important for a publication of this kind (original citation, original genus, type locality, where the types are housed, geographic distribution in Brazil and South America, larval description, and revision) was not the easiest task, and in the final moments of the work, I had to choose to split it into two separate books, because the volume of information was enormous, and the simple job of reviewing the data of the book was insane.

During the structuring work of the two guides (mainly on Coenagrionidae), I had several discussions on the generic division of species with several scientists and, in many cases, we simply could not reach a consensus, primarily by the description failures, poor figures, and absence of characters that support the proper separation in any described genera.

Fortunately, after the publication of my books, N. von Ellenrieder, R.W. Garrison, P. Pessacq, A.B.M. Machado, Janira M. Costa, and Federico Lozano solved several problems related to various genera, revalidating some, describing others, and readjusting several species in genera already described.

The guides were cited in more than 50% of the publications on South America's Zygoptera, which shows that they were widely used and that, in my opinion, was necessary. Criticisms have obviously been made, both positive and negative, and all have led me to correct errors and include everything that has been added to Zygoptera's knowledge in Brazil so far.

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In this 2nd edition, I included all papers published on the Zygoptera fauna of Brazil until August 2016 and decided not to divide them by families, but by geographic region of the occurrence of the species in Brazil, separating into three volumes. This first one deals with 159 species found in the Southeast Region of Brazil, the second will include the 144 species found in the South, Northeast and Central-west regions of Brazil, and the third volume will deal with the 186 species found in the North of Brazil.

The discrepancy in the number of species found in different regions of Brazil is caused more by the absence of researchers in various states than the real fauna distributed in those states. The environmental policies instituted in Brazil in the last decade, instead of saving endangered species from extinction, will wipe out researchers and research institutes. Soon, we will not have any more endangered species lists because there will be no one to write them, and we will probably find lists enumerating some animal groups in which all the researchers are extinct. These are very sad days.

In this book, several techniques were used to produce the best-quality images possible in each case, ones that show specific and relevant characteristics to identify the species, varying from pen and ink drawings to auto-montage (syncroscopy) images. I also include, when possible, scanned images of live or acetoned specimens.

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Index Index ................................................................................................................ 01 Chapter 1 - Introduction ............................................................................... 07 How to collect, handle, conserve and store adults ................................. 09 Adult morphology .................................................................................. 11 How to collect, rearing and prepare larvae for study ............................ 17 Larval morphology ................................................................................ 21 Museums acronyms ............................................................................... 25 Map of South America .......................................................................... 27 Chapter 2 - Key to families of Zygoptera found in the Southeast of Brazil .. 29 Chapter 3 - Dicteriadidae ............................................................................... 33 Heliocharis ............................................................................................ 37 Heliocharis amazona ............................................................................. 39 Live photos ............................................................................................ 41 Chapter 4 - Calopterygidae ............................................................................ 43 Key to genera of Coenagrionidae found in the Southeast of Brazil ..... 45 Bryoplathanon ....................................................................................... 47 Bryoplathanon globifer ........................................................................... 49 Hetaerina ............................................................................................... 51 Hetaerina auripennis ............................................................................. 53 Hetaerina brightwelli ............................................................................ 55 Hetaerina hebe ...................................................................................... 57 Hetaerina longipes ................................................................................ 59 Hetaerina proxima ................................................................................. 61 Hetaerina rosea ..................................................................................... 63 Hetaerina simplex .................................................................................. 65 Mnesarete .............................................................................................. 67 Mnesarete borchgravii .......................................................................... 69 Mnesarete guttifera ................................................................................ 71 Mnesarete hyalina ................................................................................. 73 Mnesarete lencionii ............................................................................... 75 Mnesarete pruinosa ............................................................................... 77 Mnesarete pudica phryne ...................................................................... 79 Mnesarete pudica pudica ....................................................................... 81 Mnesarete rhopalon ............................................................................... 83 Live photos ............................................................................................ 85 Chapter 5 - Lestidae ...................................................................................... 91 Key to genera of Lestidae found in the Southeast of Brazil .................. 93 Archilestes ............................................................................................. 95

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Archilestes exoletus ............................................................................... 97 Lestes ..................................................................................................... 99 Lestes auritus ......................................................................................... 101 Lestes bipupillatus ................................................................................. 103 Lestes dichrostigma ............................................................................... 105 Lestes forficula ...................................................................................... 107 Lestes minutus ........................................................................................ 109 Lestes paulistus ...................................................................................... 111 Lestes pictus ........................................................................................... 113 Lestes tricolor ........................................................................................ 115 Live photos ............................................................................................ 117 Chapter 6 - Perilestidae .................................................................................. 119 Perilestes ............................................................................................... 121 Perilestes fragilis ................................................................................... 123 Chapter 7 - Megapodagrionidae .................................................................... 125 Key to genera of Megapodagrionidae found in the Southeast of Brazil ..................................................................................................... 127 Allopodagrion ........................................................................................ 129 Allopodagrion contortum ...................................................................... 131 Heteragrion ........................................................................................... 133 Heteragrion aurantiacum ...................................................................... 135 Heteragrion beschkii ............................................................................. 137 Heteragrion brianmayi .......................................................................... 139 Heteragrion cinnamomeum .................................................................. 141 Heteragrion consors .............................................................................. 143 Heteragrion cyane .................................................................................. 145 Heteragrion dorsale .............................................................................. 147 Heteragrion flavovittatum ..................................................................... 149 Heteragrion freddiemercuryi ................................................................. 151 Heteragrion gracile ............................................................................... 153 Heteragrion mantiqueirai ...................................................................... 155 Heteragrion muryense ........................................................................... 157 Heteragrion petiense ............................................................................. 159 Heteragrion rogertaylori ....................................................................... 161 Heteragrion thais ................................................................................... 163 Heteragrion tiradentense ....................................................................... 165 Heteragrion triangulare ........................................................................ 167 Live photos ....................................................................................................... 169 Chapter 8 - Pseudostigmatidae ...................................................................... 175

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Mecistogaster ......................................................................................... 177 Mecistogaster amalia ............................................................................ 179 Mecistogaster asticta ............................................................................. 181 Mecistogaster linearis ........................................................................... 183 Mecistogaster pronoti ............................................................................ 185 Live photos ............................................................................................ 187 Chapter 9 - Protoneuridae ............................................................................. 189 Key to genera of Protoneuridae found in the Southeast of Brazil ........ 191 Epipleoneura ......................................................................................... 193 Epipleoneura machadoi ......................................................................... 195 Epipleoneura venezuelensis ................................................................... 197 Epipleoneura williamsoni ...................................................................... 199 Forcepsioneura ...................................................................................... 201 Forcepsioneura garrisoni ...................................................................... 203 Forcepsioneura grossiorum .................................................................. 205 Forcepsioneura itatiaiae ....................................................................... 207 Forcepsioneura lucia ............................................................................. 209 Forcepsioneura sancta .......................................................................... 211 Idioneura ............................................................................................... 213 Idioneura ancilla ................................................................................... 215 Idioneura celioi ...................................................................................... 217 Neoneura ............................................................................................... 219 Neoneura confundens ............................................................................ 221 Neoneura fulvicollis ............................................................................... 223 Neoneura kiautai ................................................................................... 225 Neoneura sylvatica ................................................................................ 227 Neoneura waltheri ................................................................................. 229 Peristicta ................................................................................................ 231 Peristicta aeneoviridis ........................................................................... 233 Peristicta forceps ................................................................................... 235 Peristicta jalmosi ................................................................................... 237 Peristicta janiceae ................................................................................. 239 Roppaneura ........................................................................................... 241 Roppaneura beckeri ............................................................................... 243 Live photos ............................................................................................ 245 Chapter 10 - Coenagrionidae ......................................................................... 251 Key to genera of Coenagrionidae found in the Southeast of Brazil ..... 253 Acanthagrion ......................................................................................... 257 Acanthagrion aepiolum ......................................................................... 259

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Acanthagrion ascendens ........................................................................ 261 Acanthagrion cuyabae ........................................................................... 263 Acanthagrion franciscoi ........................................................................ 265 Acanthagrion gracile ............................................................................. 267 Acanthagrion lancea .............................................................................. 269 Acanthagrion minutum .......................................................................... 271 Acanthagrion temporale ........................................................................ 273 Acanthagrion truncatum ......................................................................... 275 Acanthagrion viridescens ...................................................................... 277 Aceratobasis .......................................................................................... 279 Aceratobasis cornicauda ....................................................................... 281 Aceratobasis macilentum ....................................................................... 283 Aceratobasis mourei ............................................................................... 285 Aceratobasis nathaliae .......................................................................... 287 Angelagrion ........................................................................................... 289 Angelagrion fredericoi .......................................................................... 291 Angelagrion nathaliae ........................................................................... 293 Argentagrion .......................................................................................... 295 Argentagrion ambiguum ........................................................................ 297 Argia ...................................................................................................... 299 Argia claussenii ..................................................................................... 301 Argia croceipennis ................................................................................. 303 Argia lilacina ......................................................................................... 305 Argia modesta ........................................................................................ 307 Argia mollis ........................................................................................... 309 Argia reclusa ......................................................................................... 311 Argia smithiana ..................................................................................... 313 Argia sordida ......................................................................................... 315 Argia tamoyo ......................................................................................... 317 Cyanallagma .......................................................................................... 319 Cyanallagma angelae ............................................................................ 321 Cyanallagma nigrinuchale .................................................................... 323 Cyanallagma trimaculatum ................................................................... 325 Enallagma .............................................................................................. 327 Enallagma novaehispaniae .................................................................... 329 Fluminagrion ......................................................................................... 331 Fluminagrion taxaense .......................................................................... 333 Franciscagrion ...................................................................................... 335 Franciscagrion franciscoi ..................................................................... 337

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Franciscagrion longispinum .................................................................. 339 Franciscobasis ....................................................................................... 341 Franciscobasis franciscoi ...................................................................... 343 Franciscobasis sonia .............................................................................. 345 Homeoura .............................................................................................. 347 Homeoura chelifera ............................................................................... 349 Homeoura lindneri ................................................................................ 351 Homeoura nepos .................................................................................... 353 Ischnura ................................................................................................. 355 Ischnura capreolus ................................................................................ 357 Ischnura fluviatilis ................................................................................. 359 Ischnura ramburii .................................................................................. 361 Leptagrion ............................................................................................. 363 Leptagrion acutum ................................................................................. 365 Leptagrion afonsoi ................................................................................. 367 Leptagrion andromache ........................................................................ 369 Leptagrion bocainense .......................................................................... 371 Leptagrion capixabae ............................................................................ 373 Leptagrion elongatum ............................................................................ 375 Leptagrion macrurum ............................................................................ 377 Leptagrion perlongum ........................................................................... 379 Leptagrion porrectum ............................................................................ 381 Leptagrion vriesianum ........................................................................... 383 Metaleptobasis ....................................................................................... 385 Metaleptobasis selysi ............................................................................. 387 Minagrion .............................................................................................. 389 Minagrion caldense ............................................................................... 391 Minagrion canaanense .......................................................................... 393 Minagrion franciscoi ............................................................................. 395 Minagrion mecistogastrum .................................................................... 397 Minagrion ribeiroi ................................................................................. 399 Minagrion waltheri ................................................................................ 401 Nehalennia ............................................................................................. 403 Nehalennia minuta selysi ....................................................................... 405 Oxyagrion .............................................................................................. 407 Oxyagrion basale ................................................................................... 409 Oxyagrion brevistigma .......................................................................... 411 Oxyagrion chapadense .......................................................................... 413 Oxyagrion evanescens ........................................................................... 415

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Oxyagrion franciscoi .............................................................................. 417 Oxyagrion haematinum ......................................................................... 419 Oxyagrion hempeli ................................................................................ 421 Oxyagrion impunctatum ........................................................................ 423 Oxyagrion machadoi ............................................................................. 425 Oxyagrion microstigma ......................................................................... 427 Oxyagrion mirnae ................................................................................... 429 Oxyagrion pavidum ............................................................................... 431 Oxyagrion pseudocardinale .................................................................. 433 Oxyagrion santosi .................................................................................. 435 Oxyagrion simile .................................................................................... 437 Oxyagrion sulinum ................................................................................ 439 Oxyagrion terminale .............................................................................. 441 Telagrion ............................................................................................... 443 Telagrion longum .................................................................................. 445 Telebasis ................................................................................................ 447 Telebasis carmesina .............................................................................. 449 Telebasis coccinea ................................................................................. 451 Telebasis corallina ................................................................................ 453 Telebasis erythrina ................................................................................ 455 Telebasis filiola ...................................................................................... 457 Telebasis griffinii ................................................................................... 459 Telebasis obsoletum ............................................................................... 461 Telebasis paraensei ............................................................................... 463 Telebasis vulcanoae ............................................................................... 465 Telebasis willinki ................................................................................... 467 Tigriagrion ............................................................................................. 469 Tigriagrion aurantinigrum .................................................................... 471 Tuberculobasis ....................................................................................... 473 Tuberculobasis costalimai ..................................................................... 475 Tuberculobasis guarani ......................................................................... 477 Live photos ............................................................................................ 479 Acknowledgements ........................................................................................ 513 Figure sources .................................................................................................. 515 Glossary ........................................................................................................... 527 Bibliography .................................................................................................... 537

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DICTERIADIDAE

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Dicteriadidae28

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DICTERIADIDAE Endemic to South America, this very small family has only two monotypic genera, both are characterized by extremely long and almost bare legs and a large moveable hook on each labial palp, forcep-shaped appendages whose medial face is finely toothed and paraproct rudimentary. Medium sized damselflies (34 to 58 mm), specimens of both genera perch with wings opened horizontally and had the larvae knew, Heliocharis (Santos & Costa, 1988 and Geijskes, 1986) and Dicterias (Fleck et al, 2012). Only Heliocharis is found in the Southeast Region of Brazil.

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Dicteriadidae30

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Heliocharis Selys, 1853 Type species: Heliocharis amazona Selys, 1853 Etymology: Helio, Greek (Ήλιος) for sun + charis, Greek (χάρις) for grace. A monotypic genus whose species (Heliocharis amazona) is widely distributed (Colombia, Ecuador, Peru, Bolivia, Guyana, Suriname, French Guiana, Brazil, Paraguay and Argentina). Male Heliocharis are more common than females, and I have never found females in open sunny places, contrasting with information published by Dunkle (1991) who reported finding them in both sunny and shady areas. Observing fights between the Heliocharis amazona males is a fascinating experience. They touch each other with their long legs and fly upwards in circles, always in very sunny places. Larva found in slow-flowing creeks and rivers with litter substrate bordered by abundant riparian vegetation (Ferreira-Peruquetti & Fonseca-Gessner, 2006). The genus can be characterized by: - Medium to large damselflies (45 to 58 mm); - Perch with wings open horizontally; - Greenish thorax, blue abdomen; - Long legs with vestigial spurs; - Ventro-anteriorly projection of the penis with spatulate apex.

Fig. B1 - Heliocharis amazona male, showing the vestigial spurs (A) in contrast with regular spurs (B) of the prothoracic femur of Allopodagrion contortum.

Fig. B2: Comparison of the Labial palp of Dicteriadidae (A) with Coenagrionidae (B) in the same scale: 1 - right hook of Heliocharis amazona, 2 - right hook of Leptagrion elongatum (transposed).

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Heliocharis amazona Selys, 1853 Original description in: Bull. Acad. r. Belg. 20(Annexe): 55 Male described: Yes Female described: Yes Type locality: border of the Amazon in Bolivia. Types housed in: IRSN - st Etymology: amazona, in reference to the environment of the type locality. Synonyms: Heliocharis brasiliensis Hagen in Selys, 1859; Heliocharis libera Selys, 1869; Cyanocharis valga Needham, 1903; Neocharis cothurnata Förster, 1906; Dicterias umbra Ris, 1918; Dicterias peruviana Navás, 1920; Heliocharis paraensis Costa & Santos, 1991. Distribution in South America: Brazil, Argentina, Colombia, Ecuador, Peru, Bolivia, Venezuela, Guyana, Surinam, Guyane Française Distribution in Brazil: Goiás, Mato Grosso, Minas Gerais, São Paulo Larva described: Yes (Geijskes, 1986, Santos & Costa, 1988) Revision: Yes (Dunkle, 1991)

Fig. 01: A - Male head; B - Penis; C - Appendages lateral view; D - Appendages mediodorsal view.

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Dicteriadidae34

Fig. 02: Heliocharis amazona - Male habitus (A - alive and C - acetoned); B- female habitus (acetoned).

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Fig. BL-01: Heliocharis amazona male (Brotas - SP).

Fig. BL-02: Heliocharis amazona male, note the odd position of the hind legs (Brotas - SP).

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