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Keys to Neotropical Hexapoda
Thorp and Covich’s Freshwater Invertebrates
A Global Series of Books on the Identification,

Ecology, and General Biology of Inland Water Invertebrates
by Experts from Around the World
Fourth Edition
Series Editor: James H. Thorp
Volume I: Ecology and General Biology

Edited by James H. Thorp and D. Christopher Rogers
Published 2015
Volume II: Keys to Nearctic Fauna

Edited by James H. Thorp and D. Christopher Rogers
Published 2016
Volume III: Keys to Neotropical Hexapoda

Edited by Neusa Hamada, James H. Thorp, and D. Christopher Rogers
Published 2018
Volume IV: Keys to Palaearctic Fauna

Edited by D. Christopher Rogers and James H. Thorp
In Press: 2019
Volumes in Preparation and Under Contract

Keys to Neotropical and Antarctic Fauna
Keys to Australasian Fauna
Keys to Afrotropical Fauna
Possible Future Volumes of the Fourth Edition

Keys to Oriental and Oceanic Hexapoda
Keys to Oriental and Oceanic Fauna
Keys to Palaearctic Hexapoda
Keys to Nearctic Hexapoda
Related Publications
Field Guide to Freshwater Invertebrates of North America
by J.H. Thorp and D.C. Rogers
Keys to Neotropical Hexapoda
Thorp and Covich’s Freshwater
Invertebrates - Volume III
Fourth Edition
Edited by
Neusa Hamada
Coordenação de Biodiversidade, Instituto Nacional de Pesquisas da Amazônia
Manaus, Amazonas, Brazil
James H. Thorp
Kansas Biological Survey and Department of Ecology and Evolutionary Biology
University of Kansas, Lawrence, KS, USA
D. Christopher Rogers
Kansas Biological Survey and The Biodiversity Institute
University of Kansas, Lawrence, KS, USA
Academic Press is an imprint of Elsevier
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Cover Photos’ credits
Odonata Picture
Fernando da Costa Pinheiro
Conselho Nacional de Desenvolvimento Cientı́fico e Tecnológico (CNPq), Brazil
Trichoptera Picture
Ana Maria Oliveira Pes
Instituto Nacional de Pesquisas da Amazônia (INPA), Brazil
Collembola Picture
Diego D. da Silva
Instituto Nacional de Pesquisas da Amazônia, Manaus (INPA), Brazil
Publisher: Candice Janco

Acquisitions Editor: Louisa Hutchins
Editorial Project Manager: Emily Thomson
Production Project Manager: Paul Prasad Chandramohan
Cover Designers: Greg Harris, with contributions from the academic editors
Typeset by TNQ Technologies

Dedications from the Editors
To all the authors for agreeing to face this journey,

making the construction of this book possible
Neusa Hamada
To where the wild things are and the children,

who grow up to study and protect them
James H. Thorp
To our authors who made this book possible

This page intentionally left blank
Contents
Contributors to Volume III xiii Division II

About the Editors xvii
Preface to the Fourth Edition xix
Subclass Insecta
Preface to Volume III xxi
Acknowledgments for Volume III xxiii Section II.1
Orders Ephemeroptera,
Division I Plecoptera, Orthoptera, Blattodea,
Introduction and Subclass Hemiptera, Neuroptera,
Entognatha Megaloptera, Hymenoptera,
Trichoptera, Lepidoptera, and
1. Introduction Mecoptera
James H. Thorp, D. Christopher Rogers,
Neusa Hamada
3. Order Ephemeroptera
Frederico Falcão Salles, Eduardo
Introduction to This Series,
Domıńguez, Carlos Molineri,
Volume, and Chapter 3
Rafael Boldrini, Carolina Nieto,
Nature and Fauna of the Neotropics 4
Lucimar Gomes Dias
Components of Taxonomic Chapters 4
How to Use This Volume 4 Introduction 61
Keys to Major Taxa of Hexapoda 5 Limitations 62
References 9 Terminology and Morphology 62
Material Preparation and Preservation 63
2. Class Collembola Keys to the Order Ephemeroptera 63
References 115
Nikolas G. Cipola, Diego D. da Silva,
Bruno C. Bellini
4. Order Plecoptera
Introduction 11
Fernanda Avelino-Capistrano, Pablo Pessacq,
Limitations 16
Leandro Silva Barbosa
Terminology and Morphology 18
Material Preparation and Preservation 30 Introduction 119
Key to Neotropical Freshwater Collembola 39 Limitations 120
Acknowledgments 53 Terminology and Morphology 120
References 53 Material Preparation and Preservation 131
vii
viii Contents
Keys to Plecoptera 132 Acknowledgments 226

Acknowledgments 139 References 226
References 139
9. Order Neuroptera
5. Order Orthoptera
Caleb C. Martins, Adrian Ardila-Camacho
Ana Lúcia Nunes Gutjahr,
Introduction 229
Carlos Elias de Souza Braga
Limitations 231
Introduction 143 Terminology and Morphology 232
Limitations 145 Material Preparation and
Terminology and Morphology 145 Preservation 234
Material Preparation and Key to Aquatic Larvae of Neuroptera 234
Preservation 147 Acknowledgments 235
Keys to Semiaquatic Grasshoppers 149 References 235
Acknowledgments 167
References 167 10. Order Trichoptera
Ana Maria Pes, Ralph W. Holzenthal, Julieta V.
6. Order Blattodea
Sganga, Allan P.M. Santos, Patrik Barcelos-Silva,
Sonia Maria Lopes Lucas M. Camargos
Introduction 169 Introduction 237
Limitations 170 Limitations 238
Terminology and Morphology 170 Terminology and Morphology 239
Material Preparation and Preservation 170 Material Preparation and
Keys to Blattaria 172 Preservation 239
Acknowledgments 172 Keys to Trichoptera 245
References 172 Acknowledgments 319
References 320
7. Order Hemiptera
11. Order Lepidoptera
Felipe Ferraz Figueiredo Moreira,
Higor Daniel Duarte Rodrigues, Márlon Breno Graça, M. Alma Solis
Robert W. Sites, Isabelle da Rocha Silva
Introduction 325
Cordeiro, Oséias Martins Magalhães
Limitations 328
Limitations 176 Material Preparation and Preservation 330
Terminology and Morphology 176 Keys to Larval Lepidoptera 331
Material Preservation and Preparation 177 Acknowledgments 336
Keys to Freshwater Hemiptera 177 References 336
Acknowledgments 214
References 215 12. Order Hymenoptera
Daniell R.R. Fernandes, Ranyse B. Querino,
8. Order Megaloptera
Neusa Hamada
Adrian Ardila-Camacho,
Introduction 339
Atilano Contreras-Ramos
Limitations 342
Limitations 220 Material Preparation and
Terminology and Morphology 221 Preservation 343
Material Preparation and Preservation 223 Key to Aquatic Hymenoptera 343
Keys to Larval Megaloptera 225 References 346
Contents ix
13. Order Mecoptera 14.3. Odonata: Superfamily

Renato Jose Pires Machado
Libelluloidea
Ulisses Gaspar Neiss, Gunther Fleck,
Introduction 349
Pablo Pessacq, Kenneth J. Tennessen
Limitations 350
Terminology and Morphology 350 Introduction 399
Material Preparation and Limitations 400
Preservation 350 Keys to Libelluloidea 401
Acknowledgments 351 Acknowledgments 446
References 351 References 446
Section II.2 14.4. Superfamily Calopterygoidea

Order Odonata Danielle Anjos-Santos, Ulisses Gaspar Neiss,
Pablo Pessacq
14. Order Odonata
Introduction 449
Pablo Pessacq, Javier Muzón, Limitations 450
Ulisses Gaspar Neiss Terminology and Morphology 450
Material Preparation and
Introduction 355
Preservation 450
Limitations 356
Keys to Calopterygoidea 451
Acknowledgments 467
References 467
Keys to Odonata 363
Acknowledgments 364
References 365 14.5. Superfamily Lestoidea
Javier Muzón
14.1. Superfamily Aeshnoidea
Introduction 469
Alcimar do Lago Carvalho, Pablo Pessacq Limitations 470
Introduction 367
Limitations 368
Preservation 470
Keys to the Superfamily Lestoidea 470
Acknowledgments 472
Keys to Aeshnoidea 368
References 472
Acknowledgments 375
References 375
14.6. Superfamily Coenagrionoidea
14.2. Superfamily Gomphoidea Federico Lozano, Javier Muzón,
Danielle Anjos-Santos, Pablo Pessacq
Rodolfo Novelo-Gutiérrez, Alonso Ramı´rez,
Enrique González-Soriano Introduction 475
Limitations 476
Introduction 377
Limitations 379
Preservation 476
Key to Coenagrionidae Genera 476
Keys to Gomphidae 380
Acknowledgments 493
Acknowledgments 396
References 493
References 396
x Contents
Section II.3 15.4. Family Hydrophilidae

Order Coleoptera Bruno Clarkson, Miguel
Archangelsky, Patrıćia L.M. Torres,
15. Order Coleoptera: Introduction Andrew E.Z. Short
Cesar J. Benetti, Mariano C. Michat, Introduction 561
Miguel Archangelsky Limitations 562
Introduction 497
Limitations 499
Preservation 571
Key to Hydrophilidae 571
Acknowledgments 575
Key to Aquatic Coleoptera Families 504
References 575
Acknowledgments 516
References 516
15.5. Family Hydraenidae
15.1. Families Hydroscaphidae and Juan A. Delgado, Josefina Garrido,
Torridincolidae Albert Deler-Hernández,
Luis F. Valladares
Brunno H.L. Sampaio, Andrew E.Z. Short
Introduction 577
Introduction 519
Limitations 578
Limitations 520
Preservation 581
Keys to Suborder Myxophaga 520
Key to Hydraenidae 581
References 524
Acknowledgments 581
References 581
15.2. Families Gyrinidae, Haliplidae,
and Noteridae 15.6. Families Dryopidae, Elmidae,
Cesar J. Benetti, Mario Toledo, and Psephenidae
Daniara Colpani, Bruno A.C. Guimarães
Maria Inês S. Passos, Veronica Manzo,
Introduction 527 Crystal A. Maier
Limitations 528
Introduction 583
Limitations 584
Keys to Gyrinidae, Haliplidae, and
Noteridae 529
Preservation 584
Acknowledgments 536
Keys to Dryopidae, Elmidae, and
References 536
Psephenidae 584
Acknowledgments 598
15.3. Family Dytiscidae References 598
Cesar J. Benetti, Mariano C. Michat,
Nelson Ferreira, Jr. Rafael Benzi Braga, 15.7. Family Scirtidae
Yoandri S. Megna, Mario Toledo
Marıá L. Libonatti, Rafał Ruta
Introduction 539
Introduction 599
Limitations 540
Limitations 600
Keys to Neotropical Dytiscidae 541
Preservation 602
Acknowledgments 558
References 602
References 559
Contents xi
Section II.4 16.4. Family Thaumaleidae

Order Diptera Luiz Carlos Pinho, Heron Huerta
Introduction 719
16. Order Diptera
Limitations 719
Lı´via Maria Fusari, Galileu P.S. Dantas, Terminology and Morphology 719
Luiz Carlos Pinho Material Preparation and Preservation 720
References 721
Introduction 607
Limitations 610
Terminology and Morphology 610 16.5. Family Culicidae
Material Preparation and Jeane Marcelle Cavalcante
Preservation 612 do Nascimento, Ruth Leila
Keys to Larvae of Aquatic and Ferreira Keppler, Neusa Hamada
Semiaquatic Diptera 619
References 621 Introduction 723
Limitations 724
16.1. Family Ceratopogonidae
Maria Marcela Ronderos, Florentina Dıáz, Preservation 726
Pablo Ignacio Marino, Keys to Culicidae 729
Ruth Leila Ferreira-Keppler Acknowledgments 744
References 744
Introduction 625
Limitations 632
Terminology and Morphology 647 16.6. Family Dixidae
Material Preparation and Preservation 656 Caio C.D. Corrêa, Leonardo H.
Keys to Ceratopogonidae 658 Gil-Azevedo
Acknowledgments 659
References 659 Introduction 747
Limitations 747
16.2. Family Chironomidae
Fabio Laurindo da Silva, Luiz Carlos Pinho, Keys to Genera of Dixidae 749
Sofia Wiedenbrug, Galileu P.S. Dantas, References 749
Augusto Siri, Trond Andersen, Susana
Trivinho-Strixino 16.7. Family Corethrellidae
Introduction 661 André Pereira Amaral, Heron Huerta,
Limitations 662 Luiz Carlos Pinho
Material Preparation and Preservation 664 Introduction 751
Keys to Chironomidae 665 Limitations 752
References 698 Terminology and Morphology 752
References 753
16.3. Family Simuliidae
Neusa Hamada, Jeane Marcelle 16.8. Family Chaoboridae
Cavalcante do Nascimento
Luiz Carlos Pinho, André Pereira Amaral
Introduction 701
Limitations 701 Introduction 755
Terminology and Morphology 702 Limitations 755
Material Preparation and Preservation 703 Terminology and Morphology 755
Keys to Simuliidae 704 Material Preparation and
Acknowledgments 718 Preservation 757
xii Contents
16.9. Family Blephariceridae 16.12. Family Syrphidae

Leonardo H. Gil-Azevedo Gil Felipe Gonçalves Miranda,
Graham Rotheray
Introduction 759
Material Preparation and Terminology and Morphology 782
Preservation 760 Preparation and Preservation 783
Keys to Blephariceridae 761 Acknowledgments 783
16.10. Family Psychodidae 16.13. Family Periscelididae

Danilo P. Cordeiro, Rüdiger Wagner Rosaly Ale-Rocha, Ruth Leila
Ferreira-Keppler
Introduction 765
Material Preparation and Terminology and Morphology 786
Preservation 766 Material Preparation and
Key to Larvae of Neotropical Preservation 787
Psychodidae Subfamilies 767 References 787
References 770
16.14. Family Ephydridae
16.11. Family Stratiomyidae
Rosaly Ale-Rocha, Ruth Leila
Fabio Siqueira Pitaluga de Godoi, Ferreira-Keppler
José Roberto Pujol-Luz
Introduction 789
Introduction 771 Limitations 791
Limitations 773 Terminology and Morphology 792
Terminology and Morphology 773 Material Preparation and Preservation 793
Material Preparation and Acknowledgments 793
Preservation 773 References 793
Key to Stratiomyidae 774
Acknowledgments 776
References 776 Taxonomic Index 795
Contributors to Volume III
Rosaly Ale-Rocha [Chapters 16.13, 16.14] Coordenação Cesar J. Benetti [Chapters 15, 15.2, 15.3] Coordenação de
de Biodiversidade, Instituto Nacional de Pesquisas da Biodiversidade, Instituto Nacional de Pesquisas da
Amazônia, Manaus, Amazonas, Brazil; email: Amazônia, Manaus, Amazonas, Brazil; email:
alerocha@inpa.gov.br cjbenetti@gmail.com
André Pereira Amaral [Chapters 16.7, 16.8] Departa- Rafael Boldrini [Chapter 3] Centro de Estudos da Bio-
mento de Ecologia e Zoologia, Universidade Federal diversidade, Universidade Federal de Roraima, Boa
de Santa Catarina, Florianópolis, Santa Catarina, Vista, Roraima, Brazil; email: rafael.boldrini@ufrr.br
Brazil; email: depereira@gmail.com Carlos Elias de Souza Braga [Chapter 5] Department of
Trond Andersen [Chapter 16.2] Department of Natural Natural Sciences, State University of Pará, Belém, Pará,
History, University Museum of Bergen, Bergen, Brazil; email: bragaelias@yahoo.com.br
Norway; email: trond.andersen@uib.no Rafael Benzi Braga [Chapter 15.3] Laboratório de
Danielle Anjos-Santos [Chapters 14.4, 14.6] Centro de Entomologia, Instituto de Biologia, Universidade
Investigaciones Esquel de Montaña y Estepa Patagóni- Federal do Rio de Janeiro, Rio de Janeiro, Rio de
cas (CIEMEP), Esquel, Chubut, Argentina; email: Janeiro, Brazil; email: rafaelbenzi@gmail.com
danielleanjos2@yahoo.com.br Lucas M. Camargos [Chapter 10] Department of
Miguel Archangelsky [Chapters 15, 15.4] Laboratorio de Entomology, University of Minnesota, St. Paul, MN,
Investigaciones en Ecologı́a y Sistemática Animal United States; email: lmcamargosbio@gmail.com
(LIESA), Centro de Investigación Esquel de Montaña Alcimar do Lago Carvalho [Chapter 14.1] Departamento
y Estepa Patagónica (CIEMEP, CONICET-UNPSJB), de Entomologia, Museu Nacional, UFRJ, Rio de Janeiro,
Esquel, Chubut, Argentina; email: hydrophilinae@ Rio de Janeiro, Brazil; email: alagoc@acd.ufrj.br
gmail.com
Jeane Marcelle Cavalcante do Nascimento [Chapters
Adrian Ardila-Camacho [Chapters 8, 9] Laboratorio de 16.3, 16.5] Coordenação de Biodiversidade, Curso de
Entomologı́a, Centro de Investigaciones en Microbio- Pós-Graduação em Entomologia, Instituto Nacional
logı́a y Parasitologı́a Tropical e CIMPAT, Universi- de Pesquisas da Amazônia, Manaus, Amazonas, Brazil;
dad de los Andes, Bogotá, Colombia; email: email: jeanemarcelle@gmail.com
aardilac88@gmail.com
Nikolas G. Cipola [Chapter 2] Coordenação de Bio-
Fernanda Avelino-Capistrano [Chapter 4] Faculdades diversidade, Curso de Pós-Graduação em Entomologia,
São José, and Laboratório de Entomologia, UFRJ, Instituto Nacional de Pesquisas da Amazônia, Manaus,
Rio de Janeiro, Rio de Janeiro, Brazil; email Amazonas, Brazil; email: nikolasgc@gmail.com
fernandaacsilva@yahoo.com.br
Bruno Clarkson [Chapter 15.4] Laboratório de Sistemá-
Leandro Silva Barbosa [Chapter 4] Departamento de tica e Bioecologia de Coleoptera, Departamento de
Entomologia, Museu Nacional, UFRJ, Rio de Janeiro, Zoologia, Universidade Federal do Paraná; Curitiba,
Brazil; email: leanbarbosa@gmail.com Paraná, Brazil; email: brclarkson@gmail.com
Patrik Barcelos-Silva [Chapter 10] Divisão do Curso de Daniara Colpani [Chapter 15.2] Coordenação de
Pós-graduação em Entomologia, Instituto Nacional de Biodiversidade, Curso de Pós-Graduação em
Pesquisas da Amazônia, Manaus, Amazonas, Brazil; Entomologia, Instituto Nacional de Pesquisas da
email: patrikbarcelos@gmail.com Amazônia, Manaus, Amazonas, Brazil; email:
Bruno C. Bellini [Chapter 2] Universidade Federal do Rio daniara.entomo@gmail.com
Grande do Norte, Natal, Rio Grande do Norte, Brazil;
email: entobellini@gmail.com
xiii
xiv Contributors to Volume III
Atilano Contreras-Ramos [Chapter 8] Departamento de Ruth Leila Ferreira-Keppler [Chapters 16.1, 16.5, 16.13,
Zoologı́a, Instituto de Biologı́a, Universidad Nacional 16.14] Coordenação de Biodiversidade, Instituto
Autónoma de México, Ciudad Universitaria, Ciudad Nacional de Pesquisas da Amazônia, Manaus, Amazo-
de México, México; email: acontreras@ib.unam.mx nas, Brazil; email: ruth@inpa.gov.br
Danilo P. Cordeiro [Chapter 16.10] Coordenação de Nelson Ferreira Jr. [Chapter 15.3] Laboratório de Ento-
Biodiversidade, Instituto Nacional de Pesquisas da mologia, Instituto de Biologia, Universidade Federal
Amazônia, Manaus, Amazonas, Brazil; email: do Rio de Janeiro, Rio de Janeiro, Rio de Janeiro,
d.pacheco.c@gmail.com Brazil; email: nferrejr@gmail.com
Caio C.D. Corrêa [Chapter 16.6] Departamento de Ento- Gunther Fleck [Chapter 14.3] Ver à soie Musée-
mologia, Museu Nacional, Universidade Federal do Magnanerie, Lagorce, Ardèche, France; email:
Rio de Janeiro, Rio de Janeiro, Rio de Janeiro, Brazil; fleckgunther@gmail.com
email: caio.entomologia@gmail.com Lı́via Maria Fusari [Chapter 16] Departamento de
Galileu P.S. Dantas [Chapters 16, 16.2] Coordenação de Hidrobiologia, Universidade Federal de São Carlos,
Biodiversidade, Instituto Nacional de Pesquisas da São Carlos, São Paulo, Brazil; email:
Amazônia, Manaus, Amazonas, Brazil; email: liviafusari@gmail.com
galileu.psd@gmail.com Josefina Garrido [Chapter 15.5] Departamento de Ecolo-
Diego D. da Silva [Chapter 2] Coordenação de Bio- gı́a y Biologı́a Animal, Facultad de Biologı́a, Universi-
diversidade, Curso de Pós-Graduação em Entomologia, dad de Vigo, Vigo, Spain; email: jgarrido@uvigo.es
Instituto Nacional de Pesquisas da Amazônia, Manaus, Leonardo H. Gil-Azevedo [Chapters 16.6, 16.9] Departa-
Amazonas, Brazil; email: diegocollembola@gmail.com mento de Entomologia, Museu Nacional, Universidade
Fabio Siqueira Pitaluga de Godoi [Chapter 16.11] Federal do Rio de Janeiro, Rio de Janeiro, Rio de
Departamento de Biologia, Universidade Federal do Janeiro, Brazil; email: lhgazevedo@mn.ufrj.br
Amazonas, Manaus, Amazonas, Brazil; Email: Enrique González-Soriano [Chapter 14.2] Instituto de Bio-
fgodoi@gmail.com logı́a, Departamento de Zoologı́a, Universidad Nacional
Albert Deler-Hernández [Chapter 15.5] Department of Autónoma de México, Ciudad de México, México;
Zoology, Faculty of Science, Charles University, Prague, email: esoriano@ib.unam.mx
Czech Republic; email: adeler1982@gmail.com Márlon Breno Graça [Chapter 11] Coordenação de Bio-
Juan A. Delgado [Chapter 15.5] Departamento de diversidade, Curso de Pós-Graduação em Entomologia,
Zoologı́a y Antropologı́a Fı́sica, Facultad de Biologı́a, Instituto Nacional de Pesquisas da Amazônia, Manaus,
Universidad de Murcia, Murcia, Spain; email: Amazonas, Brazil; email: marlon_lgp@hotmail.com
jdelgado@um.es Bruno A.C. Guimarães [Chapter 15.2] Laboratório de
Lucimar Gomes Dias [Chapter 3] Departamento de Cien- Entomologia, Departamento de Zoologia, Instituto de
cias Biológicas, Facultad de Ciencias Exactas y Natu- Biologia, Universidade Federal do Rio de Janeiro,
rales, Grupo de Investigación Bionat, Universidad de Rio de Janeiro, Rio de Janeiro, Brazil; email:
Caldas, Manizales, Caldas, Colômbia; email: brunoaguilar0044@gmail.com
lucimar.dias@ucaldas.edu.co Neusa Hamada [Chapters 1, 12, 16.3, 16.5] Coordenação
Florentina Dı́az [Chapter 16.1] Centro de Estudios Para- de Biodiversidade, Instituto Nacional de Pesquisas da
sitológicos y de Vectores (CEPAVE), Facultad de Cien- Amazônia, Manaus, Amazonas, Brazil; email:
cias Naturales y Museo (UNLP), Consejo Nacional de nhamada@inpa.gov.br
Investigaciones Cientı́ficas y Técnicas (CONICET), Ralph W. Holzenthal [Chapter 10] Department of Ento-
Buenos Aires, Argentina, UNLP, FCNYM, División mology, University of Minnesota, St. Paul, MN, United
Entomologı́a, Museo de La Plata, Buenos Aires, States; email: holze001@umn.edu
Argentina; email: mfdiaz@fcnym.unlp.edu.ar
Heron Huerta [Chapters 16.4, 16.7] Instituto de
Eduardo Domı́nguez [Chapter 3] Instituto de Bio- Diagnóstico y Referencia Epidemiológicos, Laboratorio
diversidad Neotropical (IBN). CONICET - U.N.T., & de Entomologı́a, Ciudad de México, México; email:
Facultad de Ciencias Naturales, Universidad Nacional cerato_2000@yahoo.com
de Tucumán, San Miguel de Tucumán, Argentina;
email: eduardo.mayfly@gmail.com Maria Inês S Passos [Chapter 15.6] Laboratório de Inse-
tos Aquáticos, Departamento de Zoologia/ Instituto de
Daniell R.R. Fernandes [Chapter 12] Coordenação de Biociências, Universidade Federal do Estado do Rio de
Biodiversidade, Instituto Nacional de Pesquisas da Janeiro (Unirio), Rio de Janeiro, Rio de Janeiro, Brazil;
Amazônia, Manaus, Amazonas, Brazil; email: email: minespassos@gmail.com
daniellrodrigo@hotmail.com
Contributors to Volume III xv
Fabio Laurindo da Silva [Chapter 16.2] Department of Gil Felipe Gonçalves Miranda [Chapter 16.12] Gerência
Zoology, Institute of Biosciences, University of São da Região Administrativa Calha Norte III, Diretoria
Paulo, São Paulo, São Paulo, Brazil; email: de Gestão e Monitoramento de Unidades de Conser-
fabiologia@gmail.com vação, Instituto de Desenvolvimento Florestal e da
Marı́a L. Libonatti [Chapter 15.7] Laboratory of Ento- Biodiversidade, Monte Alegre, Pará, Brazil; email:
mology, IBBEA, CONICET-UBA, DBBE-FCEN, gilfgm@gmail.com
University of Buenos Aires, Buenos Aires, Argentina; Carlos Molineri [Chapter 3] Instituto de Biodiversidad
email: libonatti.marialaura@gmail.com Neotropical (IBN). CONICET - U.N.T., & Facultad
Sonia Maria Lopes [Chapter 6] Laboratório de Bio- de Ciencias Naturales, Universidad Nacional de Tucu-
diversidade e Sistemática de Blattaria, Departamento mán, San Miguel de Tucumán, Tucumán, Argentina;
de Entomologia, Museu Nacional, Universidade email: carlosmolineri@gmail.com
Federal do Rio de Janeiro, Rio de Janeiro, Rio de Felipe Ferraz Figueiredo Moreira [Chapter 7] Labora-
Janeiro, Brazil; email: sonialfraga@gmail.com tório de Biodiversidade Entomológica, Fundação
Federico Lozano [Chapter 14.6] Laboratorio de Bio- Oswaldo Cruz, Rio de Janeiro, Rio de Janeiro, Brazil;
diversidad y Genética Ambiental (BioGeA), Universi- email: felipe.moreira@ioc.fiocruz.br
dad Nacional de Avellaneda (UNDAV), Buenos Javier Muzón [Chapter 14, 14.5, 14.6] Laboratorio de
Aires, Buenos Aires, Argentina; email: Biodiversidad y Genética Ambiental (BioGeA), Univer-
lozano@undav.edu.ar sidad Nacional de Avellaneda (UNDAV), Buenos Aires,
Renato Jose Pires Machado [Chapter 13] Instituto de Buenos Aires, Argentina; email: jmuzon@gmail.com
Biociências, Universidade Federal do Mato Grosso, Ulisses Gaspar Neiss [Chapters 14, 14.3, 14.4] Instituto
Cuiabá, Mato Grosso, Brazil; email: rjpmachado@ de Criminalı́stica, Departamento de Polı́cia Técnica-
gmail.com Cientı́fica, Manaus, Amazonas, Brazil; email:
Oséias Martins Magalhães [Chapter 7] Laboratório de ulisses.neiss@gmail.com
Biodiversidade Entomológica, Fundação Oswaldo Carolina Nieto [Chapter 3] Instituto de Biodiversidad
Cruz, Rio de Janeiro, Rio de Janeiro, Brazil; email: Neotropical (IBN). CONICET - U.N.T., & Facultad
oseiasmm@bol.com.br de Ciencias Naturales, Universidad Nacional de Tucu-
Crystal A. Maier [Chapter 15.6] Gantz Family Collection mán, San Miguel de Tucumán, Tucumán, Argentina;
Center, Field Museum of Natural History, Chicago, IL, email: macarolina_nieto@yahoo.com.ar
United States; email: cmaier@fieldmuseum.org Rodolfo Novelo-Gutiérrez [Chapter 14.2] Instituto de
Veronica Manzo [Chapter 15.6] Instituto de Biodiversidad Ecologı́a, A. C., Xalapa, Veracruz, México; email:
Neotropical (IBN), CONICET - UNT. Horco Molle, rodolfo.novelo@inecol.mx
Yerba Buena, Tucumán, Argentina; email: Ana Lúcia Nunes Gutjahr [Chapter 5] Department of
vmanzo@csnat.unt.edu.ar Natural Sciences, State University of Pará, Belém,
Pablo Ignacio Marino [Chapter 16.1] UNLP, FCNYM, Pará, Brazil; email: melcam@uol.com.br
División Entomologı́a, Museo de La Plata, La Plata, Ana Maria Pes [Chapter 10] Coordenação de
Buenos Aires, Argentina; email: pmarino@fcnym. Biodiversidade, Instituto Nacional de Pesquisas da
unlp.edu.ar Amazônia, Manaus, Amazonas, Brazil; email:
Caleb C. Martins [Chapter 9] Laboratório de Morfologia anampes@gmail.com
e Evolução de Diptera, Departamento de Biologia, Pablo Pessacq [Chapters 4, 14, 14.1, 14.3, 14.4, 14.6]
FFCLRP, Universidade de São Paulo, Ribeirão Preto, Centro de Investigaciones Esquel de Montaña y Estepa
São Paulo, Brazil; email: calebcalifre@gmail.com Patagónicas (CIEMEP), Esquel, Chubut, Argentina;
Yoandri S. Megna [Chapter 15.3] Departamento de email: pablopessacq@yahoo.com.ar
Biologı́a, Facultad de Ciencias Naturales y Exactas, Luiz Carlos Pinho [Chapters 16, 16.2, 16.4, 16.7, 16.8]
Universidad de Oriente, Santiago de Cuba, Cuba; Departamento de Ecologia e Zoologia, Universidade
email: ysuarez1976@yahoo.es Federal de Santa Catarina, Florianópolis, Santa Cata-
Mariano C. Michat [Chapters 15, 15.3] Laboratory of Ento- rina, Brazil; email: luiz.pinho@ufsc.br
mology, IBBEA, CONICET-UBA., DBBE-FCEN, José Roberto Pujol-Luz [Chapter 16.11] Departamento
University of Buenos Aires, Buenos Aires, Argentina; de Zoologia, Universidade de Brası́lia, Brası́lia, Dis-
email: marianoide@gmail.com trito Federal, Brazil; email: jrpujolluz@gmail.com
xvi Contributors to Volume III
Ranyse B. Querino [Chapter 12] Empresa Brasileira de Andrew E.Z. Short [Chapters 15.1, 15.4] Department of
Pesquisa Agropecuária, Embrapa Meio-Norte, Tere- Ecology & Evolutionary Biology, Division of Ento-
sina, Piauı́, Brazil; email: ranyse.silva@embrapa.br mology, Biology Institute, University of Kansas, Law-
Alonso Ramı́rez [Chapter 14.2] Department of Environ- rence, KS, United States; email: aezshort@ku.edu
mental Sciences, University of Puerto Rico, San Juan, Isabelle da Rocha Silva Cordeiro [Chapter 7] Labora-
Puerto Rico; email: aramirez@ramirezlab.net tório de Biodiversidade Entomológica, Fundação
Higor Daniel Duarte Rodrigues [Chapter 7] Museu de Oswaldo Cruz, Rio de Janeiro, Rio de Janeiro, Brazil;
Zoologia, Universidade de São Paulo, São Paulo, São email: rocha.belle@gmail.com
Paulo, Brazil; email: higorddr@gmail.com Augusto Siri [Chapter 16.2] Instituto de Limnologı́a ‘Dr.
D. Christopher Rogers [Chapter 1] Kansas Biological Raúl A. Ringuelet’, (ILPLA, CONICET, UNLP), La
Survey and The Biodiversity Institute, University of Plata, Buenos Aires, Argentina; email: augustosiri@
Kansas, Lawrence, KS, United States; email: yahoo.com.ar
branchiopod@gmail.com Robert W. Sites [Chapter 7] Enns Entomology Museum,
Maria Marcela Ronderos [Chapter 16.1] Centro de Division of Plant Sciences, University of Missouri,
Estudios Parasitológicos y de Vectores (CEPAVE), Columbia, MO, United States; email:
Facultad de Ciencias Naturales y Museo (UNLP), SitesR@missouri.edu
Consejo Nacional de Investigaciones Cientı́ficas y M. Alma Solis [Chapter 11] MAS, Systematic Ento-
Técnicas (CONICET), Buenos Aires, Argentina, mology Laboratory, USDA, National Museum of
UNLP, FCNYM, División Entomologı́a, Museo de La Natural History, Washington, DC, United States; email:
Plata, La Plata, Buenos Aires, Argentina; email: alma.solis@ars.usda.gov
ronderos@fcnym.unlp.edu.ar Kenneth J. Tennessen [Chapter 14.3] Florida State Collec-
Graham Rotheray [Chapter 16.12] Department of Natural tion of Arthropods, Gainesville, Florida, United States;
Sciences, National Museums, Scotland, Edinburgh, email: ktennessen@centurytel.net
Great Britain; email: grahamrotheray@gmail.com James H. Thorp [Chapter 1] Kansas Biological Survey
Rafa1 Ruta [Chapter 15.7] Department of Biodiversity and and Department of Ecology and Evolutionary Biology,
Evolutionary Taxonomy, Faculty of Biological University of Kansas, Lawrence, KS, United States;
Sciences, Wroclaw University, Poland; email: email: thorp@ku.edu
rafal.ruta@uwr.edu.pl Mario Toledo [Chapters 15.2, 15.3] Museo di Storia
Frederico Falcão Salles [Chapter 3] Departamento de Naturale, Parma, Italy; email: toledo.pinguicula.
Ciências Agrárias e Biológicas, Universidade Federal mario3@gmail.com
do Espı́rito Santo, São Mateus, Espı́rito Santo, Brazil; Patrı́cia L.M. Torres [Chapter 15.4] IBBEA, CONICET-
email: ffsalles@gmail.com UBA. Laboratory of Entomology, DBBE-FCEN,
Brunno H.L. Sampaio [Chapter 15.1] Laboratório de University of Buenos Aires, Buenos Aires, Buenos
Entomologia, Departamento de Zoologia, Instituto de Aires, Argentina; email: patricialmtorres@yahoo.com.ar
Biologia, Universidade Federal do Rio de Janeiro, Susana Trivinho-Strixino [Chapter 16.2] Department of
Rio de Janeiro, Rio de Janeiro, Brazil; email: Hydrobiology, Federal University of São Carlos, São
brunnosampaio@ufrj.br Carlos, São Paulo, Brazil; email: strixino@ufscar.br
Allan P.M. Santos [Chapter 10] Departamento de Luis F. Valladares [Chapter 15.5] Departamento de Bio-
Zoologia, Universidade Federal do Estado do Rio de diversidad y Gestión Ambiental, Facultad de Ciencias
Janeiro e UNIRIO, Rio de Janeiro, Rio de Janeiro, Biológicas y Ambientales, Universidad de León,
Brazil; email: allanpms@gmail.com León, Spain; email: lfvald@unileon.es
Julieta V. Sganga [Chapter 10] Departamento de Rüdiger Wagner [Chapter 16.10] Schlitz, Germany;
Biodiversidad y Biologı́a Experimental, Facultad de email: ruediger2.wagner@t-online.de
Ciencias Exactas y Naturales, Universidad de Buenos
Aires, Buenos Aires, Buenos Aires, Argentina; email: Sofia Wiedenbrug [Chapter 16.2] Zoologische Staats-
jsganga@gmail.com sammlung München, Munich, Germany; email:
sofia.wig@gmail.com
About the Editors
Dr. Neusa Hamada has been working for more than Dr. James H. Thorp has been Professor in the Department
30 years at the National Institute for Research in Amazonia of Ecology and Evolutionary Biology at the University of
(INPA) in Manaus, Amazonas, Brazil, where she is Kansas (Lawrence, KS, United States) and a senior scientist
a Research Professor. She received her Master’s degree in in the Kansas Biological Survey since 2001. Prior to
Biological Sciences from INPA’s Entomology graduate returning to his alma mater, Prof. Thorp was a Distinguished
program and her Ph.D. in Entomology from Clemson Professor and Dean at Clarkson University, Department
University, Clemson, SC, United States. Neusa’s research Chair and Professor at the University of Louisville, Asso-
began with black flies (Diptera: Simuliidae) and subse- ciate Professor and Director of the Calder Ecology Center of
quently widened to other aquatic taxonomic groups. Since Fordham University, Visiting Associate Professor at Cornell,
1999 she has advised students at both the Master’s and Ph.D. and Research Ecologist at the University of Georgia’s
levels in INPA’s Entomology graduate program and collab- Savannah River Ecology Laboratory. He received his
orates with other graduate schools in the Amazon region, Baccalaureate from the University of Kansas (KU) and both
advising students on the taxonomy, biology, and ecology of Master’s and Ph.D. degrees from North Carolina State
aquatic insects. She has published more than 200 refereed University. Those degrees focused on zoology, ecology, and
journal articles, books, and book chapters on aquatic insects. marine biology, with an emphasis on the ecology of fresh-
She has also been active in outreach initiatives, including water and marine invertebrates. Dr. Thorp has been on the
publishing children books and creating exhibits that use editorial board of three freshwater journals and is a former
aquatic insects as model to popularize aquatic science President of the International Society for River Science. He
among school-age children and the general public. teaches freshwater, marine, and invertebrate courses at KU,
and his Master’s and Doctoral graduate students work on
various aspects of the ecology of communities through
macrosystems in rivers, reservoirs, and wetlands. Prof.
Thorp’s research interests and background are highly diverse
and span the gamut from organismal biology to community,
ecosystem, and macrosystem ecology. He works on both
fundamental and applied research topics using descriptive,
experimental, and modeling approaches in the field and lab.
While his research emphasizes aquatic invertebrates, he also
xvii
xviii About the Editors
studies fish ecology, especially as related to food webs. He

has published more than 100 refereed journal articles, books,
and chapters, including three single-volume editions of
Ecology and Classification of North American Freshwater
Invertebrates (edited by J.H. Thorp and A.P. Covich) and
two volumes in the current fourth edition of Thorp and
Covich’s Freshwater Invertebrates.
Dr. D. Christopher Rogers is a research zoologist at the

University of Kansas with the Kansas Biological Survey
and is affiliated with the Biodiversity Institute. He received
his Ph.D. degree from the University of New England in
Armidale, NSW, Australia. Christopher specializes in
freshwater crustaceans (particularly Branchiopoda and
Malacostraca) and the invertebrate fauna of seasonally
astatic wetlands on a global scale. He has numerous peer
reviewed publications in crustacean taxonomy and inver-
tebrate ecology, as well as published popular and scientific
field guides and identification manuals to freshwater
invertebrates. Christopher is an associate editor for the
Journal of Crustacean Biology and a founding member of
the Southwest Association of Freshwater Invertebrate
Taxonomists. He has been involved in aquatic invertebrate
conservation efforts all over the world.
Preface to the Fourth Edition
Those readers familiar with the first three editions of our zoogeographic regions covered by the various future
invertebrate book (Ecology and Classification of North volumes.
American Freshwater Invertebrates, edited by J.H. Thorp Our concept for T&C IV included producing one book
and A.P. Covich in 1991, 2001, and 2010) will note that the (Volume I, published in 2015) with six chapters on general
fourth edition has expanded from a North American focus environmental issues applicable to many invertebrates,
to worldwide coverage of inland water invertebrates. We followed by 35 chapters devoted to individual taxa at
gave this revised book series the name Thorp and Covich’s various levels (order to phylum, or even multiple phyla in
Freshwater Invertebrates to: (1) associate present with past the case of the protozoa). Volume I was designed both as an
related editions, unite current volumes, and link all to future independent book on ecology and general biology of
editions; (2) establish a connection between the ecological various invertebrate taxa and as a companion volume for
and general biology coverage in Volume I with the taxo- users of the keys in the regional taxonomic volumes,
nomic keys in the remaining volumes; and (3) give credit to thereby reducing the amount of information duplicated in
Professor Alan Covich for his work on the first three the taxonomic volumes. The perhaps 10e11 taxonomic
editions. For the sake of brevity, we refer to the current volumes to be published in the next decade or so will
edition as T&C IV. Whether the fifth edition of T&C will contain both keys for identifying invertebrates in specific
ever appear is certainly problematic, but who knows! At zoogeographic regions and descriptions of detailed
present we are considering producing 10e12 total volumes anatomical features needed to employ those keys. While
in the fourth edition. the vast majority of authors in T&C editions IeIII were
While I am the sole editor of the overall book series, from the United States or Canada, we attempted in T&C IV
Christopher Rogers has been a major and highly valued to attract as many editors and authors as possible from
partner in developing ideas for the fourth edition and has around the world but especially from the zoogeographic
been a senior or coeditor on all our volumes. He will also regions described in the keys. Our goals for T&C IV are to
play a major role in many of the remaining volumes improve the global state of taxonomic and ecological
because of his diverse and global knowledge of freshwater knowledge of inland water invertebrates, to help protect our
invertebrates, especially in the area of taxonomy. After aquatic biodiversity, and to encourage more students to
publishing the first two volumes, we began contacting some devote their careers to working with these fascinating
potential coeditors and authors to develop volumes for organisms. These goals are especially important because
other zoogeographic regions. Out of the seven volumes in the verified and probable percent losses of species in
this edition, three are now published and four are under wetlands, ponds, lakes, creeks, and rivers around the globe
contract. In addition, we have agreements from scientists exceed those in most terrestrial habitats.
willing to lead development of two volumes in another
bioregion. However, we are still seeking experts in fields of James H. Thorp
invertebrate taxonomy for various zoogeographic regions
to serve as highly dependable authors or even coeditors,
especially those who both study fauna and live in the
xix
Preface to Volume III
This is the third volume of the fourth edition of Thorp scientific journals which follow the International Code
and Covich’s Freshwater Invertebrates (T&C IV), and of Zoological Nomenclature. Thus, no taxa that have
the first to focus on a bioregion outside the Nearctic. merely been proposed should have been included even if
Information on the ecology and general biology of the they have been identified by the world’s expert on that
groups can be found in Volume I (Ecology and General group. “Common” species are not designated in the keys
Biology, edited by Thorp & Rogers, 2015), the companion themselves because a common species in one area may not
text for the current and all remaining books in this be common in another, and this designation can easily lead
series. All taxonomic volumes are expected to consist of to false identifications. Authors have been encouraged to
an introductory chapter followed by chapters on different end the keys at the point where further identification
taxonomic groups (phyla, class, orders, or smaller levels), without genetic analysis is not practical or when it is clear
depending on the nature of the volume. Some chapters that too many of the extant fauna have yet to be described
in the current volume are very small and others are in scientific publications. Where the majority of taxa at
quite large, such as coverage of all Trichoptera in a given taxonomic level (e.g., genus) have not been
Chapter 10. identified, we have requested that the author’s key end at
A typical chapter includes a short introduction, a brief the taxonomic level above that (e.g., subfamily).
discussion of limits to identification of taxa in that chapter, Users of these keys need to realize that taxonomy is
important information on terminology and morphology a growing and vibrant field in which new taxa are being
that is needed to use the keys, techniques for preparing described and previously accepted relationships reeval-
and preserving material for identification, the taxonomic uated. For some users, this volume may be sufficient for
keys, and a few references. Three chapters serve as their needs, but for others a companion text listing known
introductions to an order, which is then covered in greater species in a smaller geographic region may also be
detail in subsequent chapters; these consist of Chapters 14 helpful.
(Odonata), 15 (Coleoptera), and 16 (Diptera). This edition is focused on species found in fresh surface
The multilevel keys are formatted to enable users to waters, thereby reflecting the bias of existing scientific
work easily at the level of their taxonomic expertise and the literature and the habitats of springtails and insects. The
needs of their project. For that reason, we separated keys by majority of taxa described in this volume spend their larval
major taxonomic divisions. For example, a student in stage entirely in water, and some also occur there as adults.
a college course might work through one or more of the Most of the taxa, however, have an aerial adult life stage
initial Coleoptera keys to determine the family in which that rarely enters the water.
a freshwater beetle belongs. In contrast, someone working It is our hope that scientists and students from around
on an environmental monitoring project might need to the world will benefit from this volume, especially those
identify a dragonfly or midge to genus, and thus would use studying organisms in the Neotropics. Suggestions for
the relevant, detailed keys that require more background improving future volumes are welcome.
experience. We also designed most keys, where possible, to
proceed from a general to a specific character within Editors
a couplet. Neusa Hamada
We have asked authors to include only taxa that are James H. Thorp
recognized internationally by publication in reputable D. Christopher Rogers
xxi
Acknowledgments for Volume III
Many people contributed to this volume in addition to the Elsevier editors Emily Thomson and Louisa Hutchins, and
chapter authors and those acknowledged in individual the production teams in Great Britain, the United States,
chapters. We greatly appreciate all our colleagues who have and other parts of the globe. We also thank the hard workers
contributed information, figures, or reviews for Volume III in India led by Paul Prasad Chandramohan who efficiently
and also thank those who provided similar services for the produced the proofs for this volume.
earlier editions, upon which the present book partially
Neusa Hamada
relies. We are again grateful to the highly competent people
James H. Thorp
at Academic Press/Elsevier who helped in many aspects of
the book’s production from the original concept to the final
marketing. In particular, we appreciate our association with
xxiii
Division I
Introduction and Subclass

Entognatha
INTRODUCTION
Chapter 1
Introduction1
James H. Thorp
Kansas Biological Survey and Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence, KS, United States
Kansas Biological Survey and The Biodiversity Institute, University of Kansas, Lawrence, KS, United States
Neusa Hamada
Coordenação de Biodiversidade, Instituto Nacional de Pesquisas da Amazônia, Manaus, Amazonas, Brazil
Chapter Outline
Introduction to This Series, Volume, and Chapter 3 Keys to Major Taxa of Hexapoda 5
Nature and Fauna of the Neotropics 4 Hexapoda: Classes 6
Components of Taxonomic Chapters 4 Hexapoda: Insecta: Orders 6
How to Use This Volume 4 References 9
INTRODUCTION TO THIS SERIES, Palaearctic Fauna (Rogers and Thorp, 2018); (5) Keys
VOLUME, AND CHAPTER to Neotropical and Antarctic Fauna (Hexapoda only to
order) (Damboreana et al., in prep.); and (6) Keys to
The first volume of Thorp and Covich’s Freshwater Australasian Fauna (Ahyong et al., in prep). Future vol-
Invertebrates (Thorp and Rogers, 2015) provided global umes will cover Oriental/Oceanic fauna and Afrotropical
coverage of the ecology, general biology, phylogeny, and fauna (each with 1e2 volumes) as well as more detailed
collection techniques for inland water invertebrates. All coverage of Hexapoda in the Nearctic and Palaearctic
subsequent volumes have now or will have focused on Bioregions. We consider this series of 10e12 volumes to
taxonomic keys for individual bioregions of the world. be a globally expanded, fourth edition of the earlier
The taxonomic keys provide the users with means to work Ecology and Classification of North American Freshwater
at the level of their need and expertise. Our authors have Invertebrates (edited by Thorp & Covich in 1991, 2001,
tried to limit the specificity of the keys to those taxa where and 2010). While some labs may have multiple copies of
comprehensive knowledge is available for most of the taxa the “Keys to Fauna” in their region, we also recommend
in a particular group and where users cannot identify all that they have at least one copy of Volume I to obtain
life stages. useful background information on each invertebrate
We hope to have six volumes in print by the end group.
of 2019: (1) an introductory volume on ecology and The current chapter is organized into an introduction, a
general biology of inland water invertebrates of the world description of the taxonomic focus of this particular
(Thorp and Rogers, 2015); (2) Keys to Nearctic Fauna volume, a section explaining the organization of most
(Thorp and Rogers, 2016); (3) the present volume, Keys to taxonomic chapters, and a key to larger taxonomic groups.
Neotropical Hexapoda (Hamada et al., 2018); (4) Keys to This chapter’s key is designed to help the reader locate the
most pertinent chapter (a task probably useful only for
students and relatively novice taxonomists) and begin
1 This chapter was written to be a useful starting point for the previous, identifying organisms in their samples. Readers will note
current, and future taxonomic volumes in all zoogeographic regions. that chapters within and among volumes vary in the spec-
Consequently, there will be only minor to moderate differences in this ificity of their taxonomic keys. This reflects both the likely
chapter among various volumes.
Thorp and Covich’s Freshwater Invertebrates, Fourth Edition. https://doi.org/10.1016/B978-0-12-804223-6.00001-9 3

4 Thorp and Covich’s Freshwater Invertebrates
INTRODUCTION
percent of the fauna that has been named and how easily details on collecting, preparation, and preserving major
taxa can be separated by alpha taxonomic methods and taxa. Figures in each chapter are limited to those needed
associated keys. While there are occasional references to for effective use of the keys. For additional anatomical
genetic separation of taxa, the keys are primarily based on information, including figures, see the relevant chapter in
microscope identifications. Volume I.
NATURE AND FAUNA OF THE HOW TO USE THIS VOLUME

NEOTROPICS
There is an old maxim that says “keys are written by
As is clear from the title, this volume covers inland water people who do not need them for people who cannot use
Hexapoda of the Neotropics, which to some readers, them.” Nonetheless, we have made every effort to make
especially those from temperate zones, may suggest a these keys as user friendly as publication limitations
warm climate. It is important to recognize, however, that would permit.
the Neotropical Bioregion stretches from northern Each chapter includes a basic introduction to the
Mexico (warm) through the Caribbean and equatorial morphology and terminology used in diagnosing the taxa of
countries (often hot) to the frigid areas of Tierra del that section. Limitations to the current state of taxonomic
Fuego (Argentina) and the islands of Cape Horn (Chile). knowledge are also presented so that the reader may gage
It also includes both low- and high-altitude sites which the reliability of the information presented. Only the
vary greatly in their temperature range. Moreover, some established, peer-reviewed scientific literature is used to
regions are hot or cold deserts and others range up to define the taxonomic categories and epithets included. All
very humid environments. The freshwater invertebrate names, as far as we are aware, conform to the International
fauna of these diverse regions will vary accordingly. Many Code of Zoological Nomenclature. All nomina and taxo-
of these diverse habitats are described in Chapter 2 of nomic arrangements used, as well as the rejection of old
Volume I. names are based on peer-reviewed scientific literature.
Names from unpublished manuscripts, dissertations, “in
COMPONENTS OF TAXONOMIC house” designations, or records that have not been validated
are not included. Provisional names and species designated
CHAPTERS “taxon 1” or “species 1” are not used unless they were
This volume is an identification manual to the inland previously recognized and accepted in the peer-reviewed
water hexapods (springtails and insects) of the Neotropics scientific literature (Richards and Rogers, 2011). No new
where we present information needed to diagnose and species descriptions or previously unpublished taxonomic
assign these organisms to various taxonomic levels. Other arrangements are presented.
information concerning ecology, morphology, physiology, The keys are dichotomous (no triplets or quadruplets
phylogeny, and both collecting and culturing techniques are used) and are hierarchical. Thus, for a given group,
can be found in Volume I of this series. Each of the the first keys are to the highest taxonomic category. The
remaining 42 chapters in the current volume is limited to a second set of keys is to the next level, the third set to
single order or lower taxonomic level, except for Chapter the level below that one, and so on, down to the lowest
2’s coverage of all orders of Collembola. We have justifiable taxonomic level based on current knowledge of
attempted to include the following six sections in those that group. This final level varies for different groups
chapters: (1) a brief overview of the higher-level taxon; (2) depending upon the state of resolution in the scientific
a description of identification limitations for each taxon; literature. Organisms not identifiable beyond a particular
(3) details of pertinent terminology and morphology; (4) taxonomic level are left at that level.
information on preparing and preserving specimens for Properly prepared keys typically employ specific,
identification; (5) taxonomic keys (separated by level of primary, diagnostic characters. Older keys often use
identification); and (6) limited list of references. Readers different characters than the more recent keys. This shift
can find a much more extensive list of references to their in primary characters results from systematists and tax-
group in Volume I (Chapters 3 and 7e41) along with more onomists testing the importance of characters. The
Chapter | 1 Introduction 5
INTRODUCTION
ultimate goal of the systematist is to have the taxonomic that the specimen to be identified is new, a hybrid,
characters used in our keys converge as closely as possible anomalous, or a recent invasive colonist is always a
with biological reality. To a nontaxonomist, the resulting possible answer. This is fundamental to using any
process may erroneously seem merely to be “lumping and identification key.
splitting” taxonomic groups rather than the result of
Once one arrives at a name or group of possible names for
employing the scientific method to reveal natural
a specimen in hand, the specimen should then be
relationships.
compared against descriptions, distribution maps, and
Surprisingly, many users do not know how to interpret
figures of that and other taxa in that group. The
a dichotomous key, making the fundamental assumption
descriptions, figures, and maps are other tools to be used in
that a correct identification answer is always present in the
identification. Direct comparison of the specimen at hand
key. This assumption generally takes one of the following
with identified museum material or using molecular
three forms:
comparisons is also sometimes necessary for a correct
1. All species are identifiable using a given key. Many new identification.
species have yet to be discovered, let alone described. Species are not immutable, fixed in location and
Generalized geographic ranges are provided for most form. They change constantly and will continue to do so,
taxa presented herein, yet species ranges shrink, swell, confounding keys and any other identification method, such
and change elevation constantly, particularly as as trait tables, character matrices, or even genetic analyses.
weather and climate patterns shift. Species disperse, This is one reason why biology is far behind physics in the
colonize, and suffer stochastic local extinctions. In development of unified theories: biology is far more com-
addition to these natural processes, some species are plex than physics, as it involves more mutable interacting
introduced intentionally or accidentally by humans, and parts and processes.
sometimes their establishment allows other species to
invade as well. Furthermore, some regions are better
studied than others: many parts of the world have not
been explored, nor the fauna described. There are still
plenty of new taxa to encounter.
2. All variation is accounted for in the key. As stated,
KEYS TO MAJOR TAXA OF HEXAPODA
identification keys use specific, primary, diagnostic A major change in the identification keys for our fourth
characters. Problems in identification are compounded edition has been to include multiple keys per chapter that
by taxa that: (a) have different character states at generally start with a class-level key (order in this volume)
different times; (b) only have diagnostic characters at and proceed to finer and finer divisions. These allow users
certain life stages or in certain genders; and/or (c) have to work at their levels of interest, need, and skill without
severely truncated morphology (often due to lack of having to wade through extraneous taxa not in the direct
sexual selection) and lack morphological characters to line to the taxon of interest.
separate the species. Furthermore, new variation within The general characteristics of Hexapoda (insects
taxa is continually developing, and thus one cannot and springtails) are: genitalia positioned at abdominal ter-
assume that species are immutable or develop tools minus; limbs present or absent, if present, not more than
predicting those changes. three pairs; pseudolegs may be present in larvae of some
3. The key is a sufficient identification tool in and of itself. orders; one pair of antennae; wings generally present in
A key is just a tool. The fact that one has a bolt adults.
that needs removing and a wrench of the correct The following keys will allow you to move to the
size does not mean that the bolt can be loosened. next level of keys, which are in the individual chapters.
Similarly, identification keys are tools to aid in taxon Chapters 14 (Odonata), 15 (Coleoptera), and 16 (Diptera)
identification. They are primarily tools to eliminate are introductory chapters to their groups and lead to 6, 7, or
incorrect taxa from the range of possible choices, nar- 14 additional chapters, respectively, for those orders. The
rowing the field to the names that may be applicable. key to larval insects was derived in part from Chapter 1 in
Keys are the process of elimination. The possibility Volume I of the fourth edition.
INTRODUCTION
Hexapoda: Classes
1 Mouthparts internal; abdomen forked distally, and bent ventrally to meet the collophore (tubular projection) depending from the first
abdominal segment; wings absent (Fig. 1.1) ............................................................................................................ Collembola [Chapter 2]
1’ Mouthparts external; abdomen various, collophore absent; wings present or not................................................................... Insecta [p. 6]
Hexapoda: Insecta: Orders

Based in part on DeWalt et al. (2010).
1 Thorax without jointed legs ........................................................................................................................................................................... 2
1’ Thorax with three pairs of jointed legs.......................................................................................................................................................... 3
2(1) Mummy-like pupa, with developing adult structures enveloped in a case which may contain vegetable or mineral matter bound by silk
secretions; occur in holometabolous insects within inland waters (Coleoptera, Diptera, Hymenoptera, Lepidoptera, Mecoptera, Mega-
loptera, Neuroptera, and Trichoptera); generally not keyed below, but pupal keys present in some dipteran chapters ........... insect pupae
2’ Not completely enclosed in a case; mobile larvae, mostly with legs, prolegs, pseudopods, or creeping welts on one or more segments; larvae
......................................................................................................................................................................................... Diptera [Chapter 16]
3(1) With large functional wings; forewings may be rigid and chitinized, at least at the base........................................................................... 4
3’ Wingless, or with developing wings (wingpads) or brachypterous............................................................................................................... 7
4(3) Not the combination of characteristics described below............................................................................................................................... 5
4’ Abdomen narrowly constricted at junction with thorax; basal half of forewings parallel to the leading edge; last leg pair not greatly enlarged
for jumping; usually <3 mm; enter water to deposit eggs on or in aquatic insect hosts; adult Fig. 1.2 .........Hymenoptera [Chapter 12]
5(4) Mesothoracic wings rigid and chitinized (Figs. 15.7e9) or thickened in basal half (Figs. 7.29e30)......................................................... 6
5’ Meso- and metathoracic wings completely membranous with numerous distinct veins, forewings leathery; hind legs with enlarged femora
suitable for jumping; order primarily terrestrial but with some semiaquatic adult species in the tropics; Fig. 1.3.......................................
.................................................................................................................................................................................... Orthoptera [Chapter 5]
6(5) Chewing mouthparts; or if appearing as a tube, then antennae are situated half way down the tube; mesothoracic wings as elytra (e.g., rigid
and chitinized) covering metathoracic wings, elytra may be abbreviated; adults; Fig. 1.4 .................................. Coleoptera [Chapter 15]
6’ Sucking mouthparts formed into a broad or narrow tube; mesothoracic wings hardened in basal half; adults; Fig. 1.5 ..............................
......................................................................................................................................................................................Hemiptera [Chapter 7]
7(3) Without extensile labium; abdomen not terminating in lamellae or points .................................................................................................. 8
7’ Labium formed into elbowed geniculate, extensile grasping organ; abdomen terminating in three lamellae or five usually short, triangular
points; larvae; Fig. 1.6................................................................................................................................................. Odonata [Chapter 14]
8(7) Mouthparts for sucking, formed into a broad or narrow tube or a pair of long stylets................................................................................ 9
FIGURE 1.1 Lateral view of the springtail Symphypleona (Arlesminthurus, Collembola) showing its general morphology. From Chapter 2.
Bar ¼ 0.5 mm.
INTRODUCTION
FIGURE 1.2 Lateral view of the head and part of the thorax of a semiaquatic aquatic wasp (Hymenoptera) Chalcididae. From Chapter 12.
FIGURE 1.3 External structures of a semiaquatic grasshopper (Orthoptera, Acrididae). From Chapter 5.
8’ Mouthparts not for sucking, not formed into a tube or pair of stylets........................................................................................................ 10
9(8). Mouthparts a pair of long stylets, feeds on sponges; all tarsi with one claw; larvae; Fig. 1.7.............................. Neuroptera [Chapter 9]
9’ Mouthparts a broad or narrow tube; mesotarsi with two claws; adults and nymphs; Fig. 1.5 ................................ Hemiptera [Chapter 7]
10(9) Abdomen without ventral prolegs on abdominal segments 3e6 ................................................................................................................ 11
10’ Ventral prolegs on abdominal segments 3e6, each with a ring of fine hooks; larvae; Fig. 1.8 ......................... Lepidoptera [Chapter 11]
11(10) Antennae elongate, with three or more antennomeres ................................................................................................................................ 12
11’ Antennae extremely small, inconspicuous, with one antennomere; larvae; Fig. 1.9............................................ Trichoptera [Chapter 10]
12(11) Terminal appendages absent, or composed of one or two short appendages uni or bi-articulated............................................................ 13
12’ Terminal appendages composed of two or three long, multiarticulated filaments ..................................................................................... 15
13(12) Antennae with three or greater than five antennomeres .............................................................................................................................. 14
13’ Antennae with four antennomeres; larvae; Fig. 1.10............................................................................................. Megaloptera [Chapter 8]
14 (13) Abdomen posterior end with a pair of hooks inserted dorsally, curved ventrally with the apices pointing anteriorly; larvae......................
.................................................................................................................................................. Mecoptera (Nannochoristidae) [Chapter 13]
14’ Abdomen posterior end not as described above larvae .......................................................................................... Coleoptera [Chapter 15]
15(12) Abdomen with plumose, forked, or lamellar gills; with two cerci and a median caudal filament, occasionally latter is reduced to stub; tarsus
with one claw; larvae; Fig. 1.11........................................................................................................................ Ephemeroptera [Chapter 3]
15’ Abdomen with or without gills; if present gills digitiform in shape on abdominal segments 1e2 or 3, and/or at the base of the cerci; two
cerci, no median filament; tarsi with two claws; larvae; Fig. 1.12 ........................................................................... Plecoptera [Chapter 4]
INTRODUCTION
FIGURE 1.4 Example of an adult aquatic beetle (Coleoptera, Cneo-

glossidae). From Chapter 15.
FIGURE 1.6 Dorsal view of the dragonfly larva Epigomphus subsimilis
Calvert (Odonata, Gomphidea) showing wing sheaths strongly divergent.
From Chapter 14.2.
FIGURE 1.7 Lateral view of the larval stage of the Spongilla fly
Climacia (Neuroptera, Sisyridae). Bar ¼ 0.5 mm. From Chapter 9.
FIGURE 1.5 Dorsal view of the adult, aquatic bug Ambrysus sp. FIGURE 1.8 Lateral view of the moth larva Petrophila sp. (Lepidoptera,
(Hemiptera, Naucoridae). From Chapter 7. Crambidae). From Chapter 11.
INTRODUCTION
FIGURE 1.11 Nymph of the mayfly Cloeon smaeleni Lestage
(Ephemeroptera, Baetidae). From Chapter 3.
FIGURE 1.9 Lateral view of the larva of the Cheumatopsyche

(Trichoptera, Hydropsychidae). From Chapter 10.
FIGURE 1.10 Dorsal view of a larval Ilyobius sp. (Megaloptera,

Sialidae). From Chapter 8. FIGURE 1.12 Stonefly nymph (Plecoptera, Notoperla). From Chapter 4.
REFERENCES Rogers, D.C. & J.H. Thorp (eds.). 2019. Keys to Palaearctic Fauna.
Volume IV in: Thorp and Covich’s Freshwater Invertebrates, Fourth
Fraulob, M., B. Wipfler, F. Hünefeld, H. Pohl & R.G. Beutel. 2012. The Edition. Academic Press, Elsevier, Boston, MA. In Press.
larval abdomen of the enigmatic Nannochoristidae (Mecoptera, Thorp, J.H. & D.C. Rogers (eds.). 2015. Ecology and General Biology.
Insecta). Arthropod Structure & Development 41: 187e198. Volume I in: Thorp and Covich’s Freshwater Invertebrates, Fourth
Richards, A.B. & D.C. Rogers. 2011. Southwest Association of Fresh- Edition. Academic Press, Elsevier, Boston, MA. 1118 p.
water Invertebrate Taxonomists (SAFIT) list of freshwater macro- Thorp, J.H. & D.C. Rogers (eds.). 2016. Keys to Nearctic Fauna. Volume
invertebrate taxa from California and adjacent states including II in: Thorp and Covich’s Freshwater Invertebrates, Fourth Edition.
standard taxonomic effort levels. 266 p. Academic Press, Elsevier, Boston, MA. 740 p.
Chapter 2
Class Collembola
Nikolas G. Cipola
Coordenação de Biodiversidade, Curso de Pós-Graduação em Entomologia, Instituto Nacional de Pesquisas da Amazônia, Manaus, Amazonas, Brazil
Diego D. da Silva
Coordenação de Biodiversidade, Curso de Pós-Graduação em Entomologia, Instituto Nacional de Pesquisas da Amazônia, Manaus, Amazonas, Brazil
CLASS COLLEMBOLA
Bruno C. Bellini
Universidade Federal do Rio Grande do Norte, Natal, Rio Grande do Norte, Brazil
Chapter Outline Collembola: Poduromorpha: Onychiuridae: Genera 48

Introduction 11 Collembola: Poduromorpha: Tullbergiidae: Genera 49
Systematics of Collembola Orders 12 Collembola: Entomobryomorpha: Families 49
Order Poduromorpha (sensu Janssens & Christiansen, 2011) 14 Collembola: Entomobryomorpha: Isotomidae: Genera 49
Order Entomobryomorpha (sensu Soto-Adames et al., 2008) 15 Collembola: Entomobryomorpha: Entomobryidae: Genera 50
Order Symphypleona (sensu Bretfeld, 1994, 1999) 15 Collembola: Entomobryomorpha: Paronellidae: Genera 51
Order Neelipleona, Family Neelidae 16 Collembola: Symphypleona: Families 51
Limitations 16 Collembola: Symphypleona: Sminthurididae: Genera 51
Terminology and Morphology 18 Collembola: Symphypleona: Dicyrtomidae: Genera 52
Collembola Terminology 27 Collembola: Symphypleona: Sminthuridae: Genera 52
Material Preparation and Preservation 30 Collembola: Symphypleona: Bourletiellidae: Genera 52
Key to Neotropical Freshwater Collembola 39 Collembola: Symphypleona: Spinothecidae: Genera 53
Collembola: Orders 39 Collembola: Symphypleona: Katiannidae: Genera 53
Collembola: Poduromorpha: Families 39 Collembola: Neelipleona: Neelidae: Genera 53
Collembola: Poduromorpha: Odontellidae: Genera 40 Acknowledgments 53
Collembola: Poduromorpha: Brachystomellidae: Genera 43 References 53
Collembola: Poduromorpha: Neanuridae: Genera 45
Collembola: Poduromorpha: Hypogastruridae: Genera 48
INTRODUCTION to a surface and used Latin/Greek terminology to system-

atically redefine Podura: colla (Latin) or kolla (Greek)
Springtails (Hexapoda: Collembola) are small terrestrial which means glue, and embolon (Greek) which means “that
arthropods similar to insects, possessing three tagmata, one which has been thrown into something,” in this case, like a
pair of antennae, and three pairs of thoracic legs. They differ plug or tube (Bellinger et al., 2017).
from all other hexapods by the presence of three abdominal Collembola is an independent class of arthropods with
appendages: a ventral tube (or collophore), the tenaculum four orders (Poduromorpha, Entomobryomorpha, Sym-
(or retinaculum), and the furcula. These appendages are phypleona, and Neelipleona, Fig. 2.1) which currently has
considered as synapomorphies of the taxon (Hopkin, 1997). about 8700 described species in 708 genera and 29 families
The tenaculum and furcula are secondarily lost in some taxa. (Soto-Adames et al., 2008; Bellinger et al., 2017). How-
The taxonomic name “Collembola” was proposed by Lub- ever, this substantial species richness probably represents a
bock in 1870 based on the presence of the collophore in small slice of the real diversity of the class, which may
species of the genus Podura Linnæus. Lubbock observed surpass 50,000 species (Hopkin, 1997; Deharveng et al.,
that the appendage was responsible for attaching specimens 2008; Cicconardi et al., 2013).
Thorp and Covich’s Freshwater Invertebrates, Fourth Edition. https://doi.org/10.1016/B978-0-12-804223-6.00002-0 11

(A) (B)
(C) (D)
CLASS COLLEMBOLA
FIGURE 2.1 (A) Poduromorpha (Neotropiella carli) habitus, lateral view, general morphology. Bar ¼ 1 mm. (B) Entomobryomorpha (Seira dowlingi)
habitus, lateral view, general morphology. Bar ¼ 0.5 mm. (C) Symphypleona (Arlesminthurus) habitus, lateral view, general morphology. Bar ¼ 0.5 mm.
(D) Neelipleona (Neelus murinus) habitus, lateral view, general morphology. (D) Photo courtesy of Andy Murray. (https://www.chaosofdelight.org).
The potentially high species richness of springtails is condition that has evolved independently several times in
possibly related to small body size, restricted dispersal springtails (D’Haese, 2002).
capacity, widespread distribution in terrestrial and aquatic Although several species of Collembola can be
surface ecosystems, and specialization to different micro- hygrophilous and may be found both in land or directly
habitats. Concerning aquatic habitats, springtails also are associated with water habitats, these aquatic associations are
specialized for different microhabitats and can be ecolog- infrequently recorded in taxonomic or ecological papers. At
ically categorized as primary (water-dependent) or sec- present, 525 water-dependent species of springtails are
ondary (epigean hydrophilous) aquatic-associated species known worldwide, of which 465 are freshwater-dependent.
(Waltz & McCafferty, 1979; Deharveng et al., 2008). The In the Neotropical Region approximately 45 water-
primary aquatic-associated springtails can be cryophilous dependent species have been reported, of which 28 are
species, which live permanently on the top of snow in freshwater-dependent (Deharveng et al., 2008).
temperate mountains or glaciers; epineustic species, found
on the surface of freshwater, usually with adaptive
morphological structures, such as in some species of Systematics of Collembola Orders
Sminthurides, Arlesminthurus and Pseudobourletiella; and Phylogenetic relationships among Collembola orders are
intertidal (or littoral) species, which are associated with much discussed and numerous hypotheses have been pro-
beach soil, rocks, and water puddles and are capable of posed so far. Poduromorpha and Neelipleona are possibly
being submerged, such as Spinactaletes and Isotogastrura. monophyletic (D’Haese, 2002, 2003; Gao et al., 2008;
The secondary aquatic associated springtails are usually Xiong et al., 2008; Schneider et al. 2011; Schneider &
hygrophilous epiedaphic species that accidentally fall on D’Haese, 2013). On the other hand, Entomobryomorpha
the surface of freshwater (Arlé, 1971; Waltz & McCafferty, and Symphypleona appear as either monophyletic or
1979; Christiansen & Snider, 2008; Palacios-Vargas, 2013). polyphyletic taxa in different approaches (D’Haese, 2002;
Adaptation to aquatic or semiaquatic habitats is a derived Xiong et al., 2008; Schneider & D’Haese, 2013).
Chapter | 2 Class Collembola 13
CLASS COLLEMBOLA
FIGURE 2.2 (A) Ceratophysella (Hypogastruridae), mandible apex. (B) Brachystomella (Brachystomellidae), maxilla. (C) Superodontella nana
(Cassagnau, 1954) (Odontellidae), maxilla. (D) Friesea jeanneli Denis, 1947 (Neanuridae), maxilla capitulum. (E) Anurida maritima (Guérin-Méneville,
1836) (Neanuridae), maxilla capitulum. (F) Pseudachorutes auricularius Cassagnau & Rapoport, 1962 (Neanuridae), maxilla capitulum. (G) Neotropiella
murphyi Massoud, 1965 (Neanuridae), maxilla capitulum. (C) Modified from Jordana et al. (1997). (DeG) Modified from Massoud (1967).
Due to body similarities (including elongated trunk with Schneider & D’Haese, 2013; Bellinger et al., 2017).
clear tergal segmentation), Entomobryomorpha and D’Haese (2003) considered Poduromorpha as the sister
Poduromorpha were once grouped as the order Arthro- group of the other orders, and Entomobryomorpha as the
pleona. Since this group is not supported by phylogenetic sister group of Symphypleona plus Neelipleona. In the
analyses and an elongated segmented trunk is considered a same sense, the relationships between springtails with
plesiomorphic character among springtails, Arthropleona is globular body (Symphypleona and Neelipleona) are also
no longer used (Xiong et al., 2008; Schneider et al., 2011; uncertain. Due to the globular body form Symphypleona
CLASS COLLEMBOLA
FIGURE 2.3 (A) Ceratophysella (Hypogastruridae), dens and mucro. (B) Podura aquatica Linnaeus, 1758 (Poduridae), dens and mucro. (C) Ony-
chiurus argus Denis, 1925 (Onychiuridae), head and trunk dorsal pseudocelli (setae omitted). (D) Onychiurus argaensis Arbea & Jordana, 1994
(Onychiuridae), third antennal segment sense organ. (E) Mesaphorura critica Ellis, 1976 (Tullbergiidae), third antennal segment sense organ. (BeE)
Modified from Jordana et al. (1997).
has been considered a sister group of Neelipleona denticles and large fringed lamellae; and mucro, if present,
(D’Haese, 2003; Xiong et al., 2008). Other analyses indi- with fewer than three lamellae (Figs. 2.2 DeG). Brachys-
cate that Symphypleona is the sister group of Podur- tomellidae has about 130 species in 18 genera; it is the
omorpha plus Entomobryomorpha (Gao et al., 2008), or all sister group of Neanuridae, with absent mandibles and
other orders (Schneider & D’Haese, 2013). typical maxilla structure (Fig. 2.2 B), which serves as the
main characteristics for differentiating the two families
(Massoud, 1967; D’Haese, 2002; Najt et al., 2005;
Order Poduromorpha (sensu Janssens & Bellinger et al., 2017). Odontellidae presents 135 species
Christiansen, 2011) and differs from these other Neanuroids by maxilla lacking
Poduromorpha (Fig. 2.1 A) specimens are characterized by the cardo (Fig. 2.2 C) (Jordana et al., 1997; Bellinger et al.,
a strongly granulated cuticle, distinct tergal segmentation 2017). Genera such as Aethiopella, Anurida, Friesea,
and presence usually of setae on the protergite or rarely Furculanurida, Halachorutes, Neotropiella, Pseudanurida,
papillae (e.g., Isotogastruridae) (Palacios-Vargas et al., and Pseudachorutes of Neanuridae; Brachystomella, Bra-
2013). The order has about 3342 species in 333 genera; the chystomellides, and Rapoportella of Brachystomellidae;
Neotropics has at least 70 genera (Mari-Mutt & Bellinger, and Odontella and Superodontella of Odontellidae may
1990; Bellinger et al., 2017). Poduromorph species are have aquatic or semiaquatic species in the Neotropical
classified in 6 superfamilies and 11 families: Neanuroidea Region, but most are known from the Nearctic Region
(Brachystomellidae, Neanuridae, Odontellidae), Podur- (Waltz & McCafferty, 1979; Christiansen & Snider, 2008).
oidea (Poduridae), Hypogastruroidea (Hypogastruridae, Poduridae has only one recognized species, Podura
Pachytullbergiidae), Gulgastruroidea (Gulgastruridae), aquatica, with Holarctic distribution, and it is typically
Isotogastruroidea (Isotogastruridae), and Onychiuroidea found on water. Podura is characterized by dentes more
(Onychiuridae, Paleotullbergiidae, Tullbergiidae) (Bellinger than three times as long as the manubrium and surpassing
et al., 2017). the ventral tube, with distal rings of granules and modified
Neanuridae is the most diverse family, with 167 genera mucro (Fig. 2.3 B).
and about 1400 species described worldwide. They are Hypogastruridae is the second largest family of
characterized by reduced (rarely totally absent) mandibles Poduromorpha, with about 700 species in 40 genera, of
without a molar plate; modified maxillae with or without which Ceratophysella, Hypogastrura, Schoettella, and
Xenylla are found in aquatic environments in the Neotrop- Microfalculidae is a monotypic family restricted to
ical Region. Hypogastruridae is defined on symplesiomor- Magadascar.
phies such as presence of the molar plate on mandibles Isotomoidea (Actaletidae and Isotomidae) and Coena-
(Fig. 2.2. A) and absence of pseudocelli (D’Haese, 2002; letoidea (Coenaletidae) are characterized by having the
Deharveng, 2004). Pachytullbergiidae groups 13 species in body without scales and setae usually smooth or unilater-
three genera characterized by lateral projections on the ally ciliate. The dorsal chaetotaxy is more homogenous
abdomen. Sensiphorura (Pachytullbergiidae) apparently is a when compared to Entomobryoidea and macrosetae, if
semiaquatic genus. present, are always acuminate. The phylogenetic relation-
Isotogastruridae is represented by the monotypic genus ships among these groups are still unclear, but Coenaletidae
Isotogastrura with 10 species found in beaches intertwined differs from Isotomoidea by having the third abdominal
of tropical and subtropical regions (Bellinger et al., 2017). segment fused with the fourth. Actaletidae and Coenaleti-
They are recognized by the prothorax devoid of setae dae are families of marine Collembola with a few species
(Palacios-Vargas et al., 2013). recorded from Mexico, Central America, and Caribbean
Onychiuridae is the most diverse family of Onychiur- Islands, which are present in marine water the first, and
CLASS COLLEMBOLA
oidea with approximately 650 species in 56 genera, while associated with hermit crabs of the genus Coenobita
Tullbergiidae has 226 species and 32 genera. Onychiuridae (Crustacea: Decapoda) the second. Within Isotomoidea,
is apparently polyphyletic, due to divergence in ancestry Isotomidae differs from Actaletidae by tenent hairs never
between the subfamilies Onychiurinae and Tetrodonto- leaf-like and fourth to sixth abdominal segments distinctly
phorinae (D’Haese, 2002, 2003). Onychiurids differ by separated; when fused, then segments are shorter than the
third antennal article sense organ with five conical papillae length of mesothorax to third abdominal segment. Iso-
and five setae protecting the microsensilla or rod-like tomidae is the second most diverse family of the order, with
sensilla, while tullbergids lack such structures and present 112 genera and about 1,400 species, which are usually
two microsensilla and two or three dorsal sensilla (Jordana related to the soil environment (Soto-Adames et al., 2008;
et al., 1997). Onychiuroids genera that have semiaquatic Bellinger et al., 2017). However, at least 30 genera of
members are: Lophognathella, Onychiurus, Thalassa- Isotomidae have been recorded in freshwater habitats in
phorura and Tullbergia (Waltz & McCafferty, 1979; North America (Waltz & McCafferty, 1979; Christiansen &
Christiansen & Snider, 2008). Paleotullbergiidae is a Snider, 2008).
monotypic family from Africa without any known aquatic Tomoceroidea (Oncopoduridae and Tomoceridae) is
association (Bellinger et al., 2017). possibly a monophyletic group, but with unclear relation-
ships among other groups of Entomobryomorpha. They are
similar to Entomobryidae in having ciliate body setae, but
Order Entomobryomorpha (sensu differ by lacking bothriotricha on the head, second, and
third abdominal segments. Tomoceridae species are
Soto-Adames et al., 2008)
distinguished by dens basally with spines, while in Onco-
Entomobryomorpha (Fig. 2.1 B) is the most diverse order poduridae the dens lacks spines, or if present, on the distal
of Collembola, with more than 4000 described species in half (Soto-Adames et al., 2008; Bellinger et al., 2017).
200 genera, distributed among 8 families and 4 superfam- Genera found in freshwater habitats are Tomocerus from
ilies: Coenaletoidea, Entomobryoidea, Isotomoidea, and North America and Oncopodura from the Brazilian
Tomoceroidea (Soto-Adames et al., 2008). Amazon and Mexico (Arlé, 1960; Waltz & McCafferty,
Entomobryoidea (Entomobryidae, Microfalculidae, and 1979).
Paronellidae) is a monophyletic group, characterized by
mostly multilaterally ciliate body setae, cylindrical or Order Symphypleona (sensu Bretfeld,
broadening macrosetae, presence of a postocular bothrio-
1994, 1999)
trichum, and second and third abdominal segments with 2
and 3 bothriotricha, respectively (Zhang et al., 2015). Symphypleona (Fig. 2.1 C) springtails have antennae
Entomobryidae is the largest family of Collembola, with longer than head and a globular body, formed by fusion of
1839 species in 64 genera, widely distributed and found in most thoracic and abdominal segments. They are widely
almost all terrestrial and freshwater environments. Paro- distributed and quite diverse with about 1,300 species in
nellidae is also distributed worldwide, with about 550 115 genera, 10 families, and 5 superfamilies: Dicyrto-
species in 38 genera. The two families are distinguished moidea (Dicyrtomidae), Katiannoidea (Arrhopalitidae,
from each other by having crenulate dentes with small Collophoridae, Katiannidae, Spinothecidae), Sminthur-
falcate or bidentate mucrones in Entomobryidae, while the idoidea (Mackenziellidae, Sminthurididae), Sturmioidea
Paronellidae have smooth dentes and square-to-elongate (Sturmiidae), and Sminthuroidea (Bourletiellidae, Smin-
mucrones with up to 10 teeth (Soto-Adames et al., 2008). thuridae) (Bellinger et al., 2017).
Sminthuridoidea differs from other superfamilies in the Currently Neelidae comprises 44 species in 5 genera, of
presence of a clasping modification of the second and third which Megalothorax, Neelides, Neelus, and Spinaethorax
antennal articles of males. Sminthurididae differs from are recorded in the Neotropical Region. The species are
Mackenziellidae by the presence of well-developed man- usually associated with euedaphic environments and caves,
dibles and a hypognathous head. Sminthurididae has 154 but species such as Megalothorax aquaticus Stach, 1951
species in 12 genera, but in Neotropical Region species are found on freshwater in Europe, as well as Neelus spp. in
often found on water surfaces belong to Denisiella, North America (Christiansen & Snider, 2008; Bellinger
Sminthurides, and Sphaeridia spp. (Waltz & McCafferty, et al., 2017).
1979; Bretfeld & Gauer, 1994; Fjellberg, 2007; Christian-
sen & Snider, 2008).
The remaining superfamilies of Symphypleona form a
LIMITATIONS
monophyletic clade, the Appendiciphora, characterized by
the presence of a pair of modified setae (subanal append- Identification of springtails is challenging because of
ages) on the inferior anal lobes of the small abdomen of chaetotaxy, use of immature specimens, polymorphisms,
CLASS COLLEMBOLA
females. The subanal appendix may be directed toward the inattention to type specimens, and imprecise location of
anus (up) in Dicyrtomoidea and Sminthuroidea, posteriorly collected individuals. Improved descriptions should
in Sturmioidea, or toward the genital opening (down) in include chaetotaxy (distribution, morphology, size and
Katiannoidea (Bretfeld, 1999). presence or absence of some head, trunk or appendicular
Sminthuridae is one the most diverse family of Sym- setae) as well as other morphological characters. The lack
phypleona, with 260 species in 33 genera, followed by of details in descriptions can hide a range of similar species
Bourletiellidae with 241 species in 35 genera and Dicyr- under a single name or make separation of two or more
tomidae 207 species in 8 genera. Sminthuridae differs from previously described species difficult. In Brazil, a clear
Bourletiellidae in having a pair of bothriotricha on fifth example is seen with Entomobrya species, which in most
abdominal segment and pretarsus usually with a pair of cases were described mostly based in color patterns, a
setae. Dicyrtomidae resembles Sminthuridae but the fourth character which may be useful to depict species, but seen as
antennal article is much shorter than the third. highly plastic at least in some taxa (Heckman, 2001; Katz
In Katiannoidea, Katiannidae is the most diverse family et al., 2015). Descriptions of immature forms lead to
with about 210 species in 19 genera, with at least six genera uncertain species identities since they differ from adults in
recorded from the Neotropical Region. Katiannidae differs size, color, morphometry, absence of genital plate, and pore
from other families of the group by the presence of 6e8 chaetotaxy. For example, within populations of some
eyes and spatulate setae on tibiotarsus. Collophoridae has Entomobryoidea species variations occur in color pattern,
only 9 species all of genus Collophora, while Arrhopaliti- dorsal body setae, labial chaetotaxy, and number of teeth on
dae 150 species in 3 genera, both families are characterized unguis (Cipola et al., 2016, 2017; Soto-Adames, 2016).
by reduction of eyes (0e4). Collophoridae has a capitate Juveniles of many Poduromorpha have reduced chaetotaxy
bothriotrix on each side of the small abdomen; these setae and less sensilla on fourth antennal article, as well as less
are pointed in Katiannidae and Arrhopalitidae (Bretfeld, setae on dens and sometimes different type of anal spines.
1999; Bellinger et al., 2017). Few specimens can even present punctual asymmetries or
Genera of Appendiciphora considered aquatic or semi- polymorphisms, especially regarding chaetotaxic elements
aquatic in North America are: Arrhopalites (Arrhopalitidae), on the left and right sides.
Collophora (Collophoridae), Sminthurinus (Katiannidae), The misuse of corrosive chemicals during clarification,
Bourletiella, Deuterosminthurus, Heterosminthurus, Pseu- improper choice of conservation medium or storage of
dobourletiella (Bourletiellidae), Neosminthurus, Sminthu- slides under inadequate conditions can lead to material
rus, and Sphyrotheca (Sminthuridae) (Waltz & McCafferty, degradation. Another problem is the absence of important
1979; Christiansen & Snider, 2008). collecting data attached to the material, especially
regarding type locality, which hinders recollection of spe-
cies from their original collection sites. An example of this
Order Neelipleona, Family Neelidae problem is Dicranocentrus silvestrii Absolon, with the type
Neelidae (Fig. 2.1 D) is the only family in Neelipleona, and locality given as “South America” leading to a very unclear
similarly to Symphypleona, species have globular bodies. type locality (Mari Mutt, 1979). Mastigoceras camponoti
They differ from Symphypleona by minute body size (some Handschin, a monotypic endemic taxon from Brazil, was
Symphypleona species may be also minute), antennae stated by the author to have been collected from “South of
shorter than head, absence of eyes, sensory fields on the Minas Gerais state,” which is one of the largest federal units
large abdomen, and reduced small abdomen hidden of Brazil (Heckman, 2001). Another example is provided
ventrally (Schneider & D’Haese, 2013). by Börner (1906) who described three species from
“São Franscisco, Brazil,” which could be several locations phytogeographic domains in Central and South America
in different regions of the country. are much understudied. Most of the species recorded from
Neotropical springtails are not well known compared the New World have been described from the United
to other regions. Numerous ecosystems and even large States, Mexico, Puerto Rico, and Brazil, with few or no
species from other countries (Mari-Mutt & Bellinger,
1990). At the same time, large countries with compara-
tively well-known faunas like Brazil are still under-
sampled in many regions and/or federal units (Zeppelini
et al., 2017). Second, the Neotropical Region potentially
has a high biodiversity of land arthropods, including
Collembola, so it is expected that many new species will
be found in unexplored habitats. Lack of knowledge
combined with a large undescribed fauna limits the overall
comprehension of the distribution and composition of
CLASS COLLEMBOLA
Collembola fauna in Neotropical Region. For instance,
regarding the aquatic fauna, only 45 species in 15 genera
have been recorded so far, which we believe does
not reflect the real species richness of aquatic springtails
in the region since these numbers were gathered from
isolated data, not from directed collections (Deharveng
FIGURE 2.4 (A) Rapoportella mucronata Najt & Massoud, 1974
(Brachystomellidae), maxilla capitulum. (B) Brachystomellides navar-
et al., 2008).
inensis Weiner & Najt, 1997 (Brachystomellidae), fourth antennal segment All of these factors lead to the lack of identification keys
ventral side. (A, B) Modified from the original description. and specific diagnoses for freshwater springtails from the
FIGURE 2.5 (A) Ceratophysella (Hypogastruridae), sixth abdominal segment spines. (B) Triacanthella nivalis Cassagnau & Deharveng, 1974
(Hypogastruridae), sixth abdominal segment spines. (C) Paraxenylla affiniformis Stach, 1929 (Hypogastruridae), fifth and sixth abdominal segments and
anus position (dorsal view). (D) Xenylla capixaba Fernandes & Mendonça, 2010 (Hypogastruridae), sixth abdominal segment and anus position (ventral
view). (E) Ceratophysella (Hypogastruridae), third tibiotarsus distal region, and empodical complex. (F) Microgastrura sofiae Vázquez & Palacios-
Vargas, 1996 (Hypogastruridae), mandible apex. (G) Ceratophysella (Hypogastruridae), fourth and third antennal segments ventral faces. (B, C)
Modified from Thibaud et al. (2004). (D, F) Modified from the original description.
Neotropics. The only specific key to freshwater springtails segments may be subdivided, creating the impression of
of the New World is for the Nearctic taxa (Waltz & Vor VI segments (Figs. 2.8 D, E). The antennal article III is
McCafferty, 1979), since the key to Neotropical species usually undivided and possesses an apical sense organ with
provided by Heckman (2001) is not specific for aquatic distinctive setae or and associated sensilla (Figs. 2.3 D, E).
taxa, therefore, is not directly relevant to Neotropical In males of Sminthurididae (e.g., Sminthurides) and in
Collembola fauna. Coenaletidae (Coenaletes) the antennal articles IIeIII are
modified into a clasping organ for holding hold females
during courtship (Fig. 2.46). The antennal article IV can be
TERMINOLOGY AND MORPHOLOGY
undivided (more common), annulated (e.g., Dicranocen-
The general morphology of Collembola is based mostly on trus and Mastigoceras) or pseudosegmented (e.g.,
the following references: Hopkin (1997), Christiansen & Sminthurides, Temeritas) (Figs. 2.8 DeE, 55). This
Bellinger (1998), Fjellberg (1999, 2007), Heckman (2001), segment often has an apical bulb and numerous sensory
Deharveng (2004), and Bellinger et al. (2017). Specific modifications (sensilla) of taxonomic significance, espe-
references for each order follows: Massoud (1967), Jordana cially in Poduromorpha (Figs. 2.4 B, 2.5 G, 6 F, 7 A, 21).
CLASS COLLEMBOLA
et al. (1997), Thibaud et al. (2004), and Najt et al. (2005) In most Collembola taxa the fourth segment is the longest,
for Poduromorpha; Potapov (2001) and Soto-Adames et al. but in Dicyrtomidae and most Tomoceridae it is much
(2008) for Entomobryomorpha; Richards (1968), Betsch shorter than the third segment (Fig. 2.38).
(1980), and Bretfeld (1999) for Neelipleona and The eyepatches are dorsal and bear a small set of eyes
Symphypleona. (ommatidia) with up to 8 lenses on each side (Figs. 2.1
Springtails, as well as other hexapods, have the body AeC, 18, 25, 33 B, 57 A, 58 A). Eyepatches are usually
divided into three tagmata: head, thorax and abdomen pigmented (black, brown, or dark blue). Reduction in
(Figs. 2.1 A, B). The body is entirely covered by a thin number of ommatidia is common and nearly all families
granulous cuticle, which is variably hydrophobic. The have taxa with fewer than 8 þ 8 eyes. In a few families the
granules of epicuticle are typically hexagonal patches number is always reduced, as in Collophoridae (4 þ 4
formed by the connection of primary triangular micro- eyes), Arrhopalitidae (2 þ 2 or less), and Neelidae and
granules. This pattern of cuticular organization is quite Cyphoderinae (Paronellidae) (without eyes) (Figs. 2.1 D,
evident in scanning electron microscope photographs of 32 A, 33 A, 66 A).
Poduromorpha species. Other taxa may have reduction or Between the eyes and antenna in some families a
fusion of such granules giving the epicuticle another postantennal organ (PAO) may be present (e.g., Brachysto
microprofile. In some species of Symphypleona, the mellidae, Neanuridae, Isotomidae, and rarely in Entomo-
cuticular granules can also be square (formed by four bryidae), simple and oval (e.g., part of Isotomidae) to quite
linked granules), not hexagonal. The cuticle has various complex or formed by many circular vesicles (moruliform,
sizes of setae (usually smooth or ciliated), spines, sensilla e.g., part of Anurida, Morulina, and Neotropiella) (Figs. 2.6
and/or scales, which combined are mostly diagnostic for GeH, 7 E, 18e19). The structure of the PAO is variable and
different taxonomical groups (Figs. 2.3, 4 B, 5 AeD, 6, 7, of taxonomic importance for being a hydrophilic structure
9, 13, 16, 20e24, 34, 37, 42, 45e48, 50, 52, 53, 58e63, 65, and it is possibly a chemo, thermo, or hygroreceptor.
66). An example of this is the presence of scales in some The dorsal region of the mouth is covered by a variably
Entomobryomorpha (Figs. 2.8 AeD, 9 B, 23). developed clypeus usually carrying some setae and termi-
The head is dorsally rounded, elliptical, or subtriangular nating in the prelabrum and an anterior labrum, both with a
and bears a pair of antennae, eyepatches and postantennal variable number of setae (Fig. 2.67). The labium is found
organs (which can be totally absent), and mouthparts on the ventral side of the head. In most Collembola it
(Figs. 2.1, 8, 18, 25, 32, 33, 38, 48, 55, 57, 66A). Springtails consists of a pair of palps often with five main papillae and
are prognathous (most Poduromorpha and Entomo- a variable number of accessory (guard) setae. Each palp is
bryomorpha) or hypognathous (Neelipleona and most associated with three labial plates, each with a few setae.
Symphypleona) (Fig. 2.1). The maxillary palps (or maxillary outer lobes) are basically
Antennae are inserted anterodorsally on head and have conical projections seen laterally of labial papillae. Each
intrinsic muscles that allow independent movement of all palp usually has one apical and one basal setae, with rare
four antennal articles (IeIV). Antenna size is variable exceptions of one or two additional setae in Isotomidae
within different taxa and they can be shorter than the head (e.g., Desoria). The sublobal plate may have up to four
(e.g., Neelidae, Figs. 2.1 D, 66 A) to much longer than the setae-like projections which can be totally absent. Finally,
whole body (e.g., Campylothorax, Salina, and Temeritas) labial and postlabial regions present several typical setae of
(Figs. 2.32 B, C, 55 A). The antennal articles IeII are taxonomic importance.
usually simple and undivided, but in some cases the first Inside the cephalic capsule are the mouthparts, mostly
(e.g., Mastigoceras) and the second (e.g., Dicranocentrus) formed by a pair of strong chewing mandibles and a pair of
(A) (B)
(C) (D)
CLASS COLLEMBOLA
(E) (F) (G) (H)
(I)
FIGURE 2.6 (A) Neanura muscorum (Templeton, 1836) (Neanuridae), fifth and sixth abdominal segments (dorsal view). (B) Paranura magdalenae
Benito & Palacios-Vargas, 2008 (Neanuridae), sixth abdominal segment (dorsal view). (C) Paranura colombiana Palacios-Vargas & Peñaranda-Parada,
2005 (Neanuridae), sixth abdominal segment (dorsal view). (D) Pseudachorutes parvulus Börner, 1901 (Neanuridae), fifth and sixth abdominal segments
(dorsal view). (E) Neanura muscorum (Templeton, 1835) (Neanuridae), head tubercles, setae omitted (dorsal view). Arrow points to unpaired antenno-
frontal tubercule (clypeal). (F) Sensillanura santizaragozai Palacios-Vargas & Catalán, 2010 (Neanuridae), fourth antennal segment sensilla (setae
omitted). (G) Aethiopella condei Massoud, 1963 (Neanuridae), frontal eyes and postantennal organ. (H) Pseudachorutes subabdominalis Steiner, 1958
(Neanuridae), frontal eyes and postantennal organ. (I) Hylaeanura mendoncae Zeppelini & Palacios-Vargas, 2013 (Neanuridae), fourth abdominal
segment ventral side (setae omitted) and reduced furcula. (A, G) Modified from Massoud (1967). (B, C, F, I) Modified from the original description.
(D, E, H) Modified from Jordana et al. (1997).
CLASS COLLEMBOLA
FIGURE 2.7 (A) Odontellina nivalis (Cassagnau, 1959) (Odontellidae), fourth antennal segment apex. (B) Thalassaphorura brevisetosa Sun, Gao &
Potapov, 2013 (Onychiuridae), fourth abdominal segment ventral view (furcula area). (C) Onychiurus gulinensis Sun & Zhang, 2012 (Onychiuridae),
fourth abdominal segment ventral view (furcula area). (D) Agraphorura calvoi Arbea, 2005 (Onychiuridae), postantennal organ. (E) Thalassaphorura
brevisetosa Sun, Gao & Potapov, 2013 (Onychiuridae), postantennal organ. (F) Mesaphorura matilei Thibaud & Palacios-Vargas, 2000 (Tullbergiidae),
sixth abdominal segment dorsal view. (A) Modified from Massoud (1967). (BeF) Modified from the original description.
lateral maxillae (Figs. 2.2, 4 A, 5 F, 64). The mandibles enlarged and strongly projected along the midline
consist apically by an inner row of incisors teeth (4e6) (e.g., Lepidocyrtoides, Lepidocyrtinus, and Campylo-
followed by a basal molar plate with numerous conical thorax) (Figs. 2.8 A, B, 32 C).
denticles (Fig. 2.2 A). Mandibles are absent in Each leg is formed by epicoxa, subcoxa (epicoxa or
Brachystomellidae (Fig. 2.2 B) and lacking molar plates in both may be fused with body), coxa, trochanter, femur,
Neanuridae (Figs. 2.2 DeE). The maxillae are usually tibiotarsus, pretarsus, and one claw (with or without an
formed by a variable number of apical teeth plus some empodial appendagedunguiculus). Coxae usually have
fringed associated lamellae (Figs. 2.2 BeE). Maxillae teeth only a few setae. In some Symphypleona, such as Smin-
and lamellae could also be reduced or modified, as seen in thuridae, the hind subcoxa possesses a robust spine-like
styliform maxillae of Pseudachorutinae (Neanuridae) seta (Fig. 2.40). The next segment, the trochanter, usually
(Figs. 2.2 F, G). The mandibles and maxillae can be elon- presents few setae, but in Entomobryoidea and Tomocer-
gated in Rhynchocyrtus, Stenognathriopes, and Sten- oidea the hind trochanter may have a field of small, smooth
ognathellus (Figs. 2.63 B and 64). spine-like setae (trochanteral organ). The Symphypleonan
The trunk is formed by a thorax with ambulacral legs families Katiannidae and Sminthuridae may also have
and a legless abdomen. It can be cylindrical or elongated trochanteral organ composed by a single robust spine
and have distinct segments (Entomobryomorpha and (Fig. 2.40). In these families such organ can occur on both
Poduromorpha), or the segments can be mostly fused to middle and hind trochanters, and may not be homologous
each other into a rounded or globose body (Symphypleona to the trochanteral organ in Entomobryoidea.
and Neelipleona) (Fig. 2.1). The femur (plural, femora) is an undivided segment
The thorax has three segments, prothorax (I), meso- with setae and/or scales (Fig. 2.1). This segment usually has
thorax (II), and metathorax (III), each with a pair of legs. minor taxonomic value but a remarkably exception is the
The prothorax is the shortest segment. Poduromorpha has a entomobryid Tyranosseira, in which males have broadened
distinct plate-like pronotum with setae and papillae femora with distinct spines on the first pair of legs (Bellini
(rarely), while Entomobryomorpha lacks a distinct prono- & Zeppelini, 2011).
tum and dorsal prothoracic setae (Figs. 2.1 A, B). The The tibiotarsus (plural, tibiotarsi) is presumed to be
mesothorax and metathorax are simple and similar to each formed by the fusion of tibia and tarsomeres seen in other
other, although in few cases one or the other may be hexapods. This segment usually is long and covered by
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“May I ask if my learned friend has any evidence to back up that
suggestion?” Sir Richard Tester asked quietly.
But Mr. Ruggles made no answer.
“The insinuation is in keeping with the rest of the case—
groundless,” Sir Richard continued.
“The bail will be as before,” the magistrate announced abruptly.
And John Gaunt, who stood in the dock, drew a deep breath of
relief.
CHAPTER XXXV
W HEN Lady Mildred was told that her sister had gone away in
the motor-car without leaving any message, she at once
jumped to the conclusion that her journey was to London. She had
been aware that Lady Ethel had been very anxious, and it was
natural to think that she might go to see John Gaunt with the idea of
affecting a reconciliation between herself and her husband.
The week’s solitude had somewhat softened her feelings, but
there still remained some bitterness. And perhaps, too, jealousy that
Gaunt would not make a sacrifice to retain her love.
But when the afternoon came to an end, she began to wait with
impatience her sister’s return, and her restlessness increased with
each passing hour. When at last the sound of the car could be heard
she ran eagerly to meet her.
“Where have you been? Why did you not tell me you were going?”
she demanded impetuously.
Lady Ethel smiled a little wanly as she threw off her furs.
“I have been to London. But let us go to your room, dear.”
The sisters did not speak again until they were alone and there
was great anxiety in Lady Mildred’s eyes.
“I have seen your husband.”
“Tell me at once. Why do you hesitate?” Lady Mildred cried
vehemently.
“It is difficult to explain what happened. It was as I thought. Your
letter has wounded him deeply, and I am afraid that you must go to
him, if you wish to be forgiven.”
“Forgiveness!” Lady Mildred repeated scornfully.
“Yes. You, I think, are in the wrong. If you take my advice you will
order the car and go to London at once. You would get there by ten
o’clock.”
“Why should I go to him? It is I who have been wronged, and he
must come to me.”
“If that is your state of mind, then I fear that you will remain apart.
It is a pity, for I am convinced that you are both passionately in love
with one another. Pride is responsible for the ruin of many lives, and
I pray you not to be stupid, dear,” Lady Ethel said gently.
Lady Mildred was silent, and it was evident that she was struggling
with herself. It was strange to find that her sister’s sympathy was so
decidedly in favor of her husband, and she began to wonder if, after
all, she herself might be to blame. But the scene in court came back
to her mind, and she could hear the counsel’s words as he described
the charge against Gaunt.
And while she fought with her pride, she was conscious that her
love for John Gaunt was burning fiercely, and that if he were to
appear she would receive him with widely opened arms.
“I went to Park Lane, and no one was there,” Lady Ethel continued
quietly. “I chanced to pass your husband’s room and I saw many
trunks there, so I imagine that he is going away.”
Lady Mildred drew a quick breath, and a look of fear crept into her
eyes when she remembered the talk in the library.
Could John be going to the Congo? Was it possible that they were
about to carry out the mad scheme that they discussed?
“Ethel, you must come back with me,” she cried eagerly.
“I am very tired, but I will gladly come,” Lady Ethel answered, and
all the listlessness vanished in an instant.
“Give orders for the car, and I will see about the packing. Baby will
stay here with nurse.”
“You are going to John?”
“Yes. Yes. I have been a fool, and I see it for the first time. You are
right, Ethel. There is only one thing that matters.”
“Love,” Lady Ethel repeated softly, and the color came to her
cheeks.
Half an hour later the sisters were driving back to London, and the
chauffeur had received instructions to travel as quickly as possible.
Now and then the car swayed ominously, but neither of them noticed
the tremendous rate at which they were going. Lady Mildred began
to tell of Gaunt’s plans to hold up the Congo Free State, and her
story was listened to with intense eagerness.
“Is Mr. Drake going?” she asked in a low voice.
“Yes, and his brother. It is a curious business for a parson to join
in.”
“Mildred, I warn you to prepare yourself for a disappointment if you
think that you will be able to persuade your husband to back out of it.
He loves you passionately; but there are some things that have more
weight with him than love.”
“I don’t think he will go, if I can only see him,” Lady Mildred
answered confidently.
The conversation languished and they were both relieved when at
last the car stopped at their house in Park Lane.
“Is your master in?” Lady Mildred demanded of the footman.
“No, my lady. He left a couple of hours ago,” was the answer.
She entered quickly and ran up to her room in the hope that there
might be a letter for her, but there was nothing there.
“I will ask the servants if there is any message. You had better rest
quietly; you look ill,” Lady Ethel said and hastened away.
She returned in a few minutes, and her face was grave as she sat
down by her sister.
“He has gone, and has left no message at all. I have been to his
rooms, and the trunks are not there,” she announced.
“What does it mean?” Lady Mildred asked hoarsely. “Do you think
that he can already have started?”
“No, but we must be very careful what we say. We must be loyal to
John, and we must not arouse any suspicion among the servants.”
“It is nearly eleven o’clock—too late to do anything to-night,” Lady
Mildred said miserably.
There was a knock at the door and Lord Lynton came in hurriedly.
“Have you any idea where Gaunt is?” he asked anxiously.
“No. Haven’t you seen him?”
“I have been dining with Sir Keith Hamilton, and afterwards he
talked to me very plainly. It appears that he has formed a great liking
for Gaunt, and he expressed his opinion very forcibly about this
police court business. He is convinced that Gaunt played no
unworthy part in that Marillier affair, and that the whole thing is a put
up job on the part of the Baron. I chanced to say that I had taken up
rather a strong attitude, and he promptly told me that I was a fool,
advising me to go and make my peace at once.”
“And I have come on a like errand, but I am afraid that we are too
late,” Lady Mildred said brokenly, and tears forced themselves to her
eyes.
“Surely there ought to be no difficulty in finding him,” the Earl said
decidedly.
“Perhaps he has already left the country?” Lady Ethel broke in.
“No, that’s impossible, for he is on bail,” Lord Lynton replied.
“Bail would not prevent him from going. I think I ought to tell you
——”
But Lady Mildred pulled her sister up sharply.
“We have no right to tell John’s secrets to any one. Perhaps we
are worrying ourselves about nothing. There may be news in the
morning,” she said hopefully, but there was no hope in her heart.
The morning came and the sisters breakfasted at an early hour.
But there was no letter.
“Shall we call at the Carlton? Captain Drake was staying there,”
Lady Ethel suggested, and they visited the hotel, only to find that he,
too, had left on the previous evening.
“I believe that he has gone to the Congo,” Lady Mildred cried
vehemently when they were once more in the car.
Next they visited Gaunt’s office in the city, where they were
received by Mr. Foster, the secretary, who could give them no
information.
“Mr. Gaunt has ceased to come here, my lady. In a couple of
months’ time the office will be finally closed.”
“When did you last see Mr. Gaunt?” Lady Mildred asked eagerly.
“Yesterday, but I have no idea where he is at present.”
“Cannot you tell me if he is going abroad?” she persisted.
“No, my lady. I have no information at all. It is not Mr. Gaunt’s
custom to tell me of his movements, unless he thinks it necessary for
me to know,” Mr. Foster answered with a smile.
They returned straight to Park Lane, for that was the most likely
place at which they would receive news, but when evening came
Lady Mildred was convinced that her suspicions were correct, and
she grew utterly despondent. It was revealed to her how great was
her love for her husband now that she believed that he had gone
deliberately into danger. All the dreadful things that she had heard of
West Africa came rushing to her mind, and she grew desperate
when she realized that he had left her with anger in his heart. And
she broke down completely, sobbing piteously; in vain Lady Ethel
strove to comfort her.
“I love him—oh, so much—and it is my fault that we quarreled. I
should have known that he would not do so dreadful a thing as that
wicked barrister charged him with. What shall I do? I must see him,”
Lady Mildred cried passionately.
“Perhaps he has only gone into the country for a rest,” her sister
suggested. “I don’t think that he could have left for the Congo without
saying a word.”
“You don’t understand John. It was my lack of trust, and that cruel,
cruel letter that hurt him. What a fool I have been.”
The morning came, bringing no news, and they wandered listlessly
about the house. Suddenly Lady Ethel gave a cry, and almost
fiercely clutched her sister’s arm.
“Did you not say that they intended to stop at the Canary Islands?
To-day is Saturday, and the Union-Castle Liner sails. Let us go to
Waterloo and see if John or one of the Drakes is there,” she cried
impetuously.
The car was ordered at once, and upon arriving at the station they
found the boat train drawn up, and by the bustle around it was
evidently soon to start. They walked eagerly along closely examining
each carriage, but saw no signs of either of them.
A warning whistle, and the train moved away.
“Of course they could join the Saxon at Southampton, for John
would not wish to be seen. Could we get there in time?” Lady
Mildred cried excitedly.
“At any rate we can try,” Lady Ethel answered eagerly. And then
began a mad race to the port. She urged the chauffeur to full speed,
and the powerful car tore along the road. Once or twice a policeman
held up a warning hand, but they went on heedlessly until they
reached the dock gates.
“Has the Saxon sailed?” they asked anxiously.
“Just about starting,” was the answer.
Now their progress was tantalizingly slow, and when at last the
wharf came in sight, they saw that the ropes had been thrown off
and that the big liner was moving towards Southampton water.
But Lady Mildred’s eyes were fixed on the upper deck, and she
could just make out the figure of a man leaning against the rails.
There were field-glasses in the car, and with trembling fingers she
focused them.
“It is John, and we are too late,” she said with a gasp.
But she would not give way, and turned to her sister her face white
and her lips trembling.
“Our yacht the Heron is lying here. She can be commissioned in a
few days,” she said very quietly.
“What are you going to do?” Lady Ethel asked in wonder.
“Follow them,” she answered curtly.
“And I will go with you,” Lady Ethel said in a low voice, but her
eyes were shining brightly.
CHAPTER XXXVI
T HERE was a feeling of constraint when Gaunt led Edward Drake

from the room after the latter’s interview with Lady Ethel.
“I am afraid that I came in at a rather inopportune moment, but this
is not the time for love-making,” Gaunt remarked kindly.
Half an hour later they were with Captain Armstrong, who had just
arrived from Newcastle to make final arrangements.
“We can sail in a couple of days,” the captain remarked cheerfully.
“And if we leave by the Saxon on Saturday, you can pick us up at
Las Palmas. With the superior speed of the Esmeralda you ought to
arrive about the same time we do,” Gaunt remarked.
“She did twenty-five knots in her last trial,” the captain answered
complacently.
“You are quite satisfied with the men you have got?”
“Yes, but I have had to be very cautious, for there are one or two
naval men at Elswick who have been mighty curious. I must say that
Señor Alvarez has played the game skillfully, for they all think that I
have entered the San Salvador navy. But I shall be glad when we
are outside British waters.”
“Then you had better get away as soon as possible. Of course you
must not come into Las Palmas harbor?” Gaunt asked.
“No, for one doesn’t generally go to South America via the
Canaries. I intend to lie a few miles off, and I shall come ashore in
the pinnace. I suppose I shall find you at the Santa Catalina Hotel?”
“Yes. Captain Drake will be with us, and he will look after the men
who are already there.”
They dined at the Carlton Hotel, and Captain Armstrong was in the
highest spirits, in contrast to the others. The meal had just come to
an end, and they had lighted their cigars when Edward Drake gave a
significant look at Gaunt. The latter turned round and saw that the
Baron de Croiseuil was seating himself at a table on the other side of
the room.
“It is as well that he should see me here. Hang it, I’ll go over and
speak to him,” Gaunt said suddenly, and immediately hastened
across the room.
“Good-evening, Baron,” he said suavely, and took a chair on the
other side of the table.
For once in his life the Baron lost his composure, and flushed
hotly.
“You have done your worst and have failed,” Gaunt continued
quietly.
“Not quite, for I have effectually closed your mouth, my friend. It
was very painful to me—the necessity of taking these proceedings.
But still, it is possible to end them. One word, and Mr. Ruggles will
explain that a dreadful mistake has been made,” the Baron said
coolly, after a supreme effort at self-control.
“I do not intend to say that word. You have done your worst. You
have put forward a false charge in an infamous way. You have
attempted to ruin me utterly. And do you think that I am the man to
take it lying down? No, my friend. It soon will be my turn,” Gaunt said
grimly.
“You can do nothing, and I am still willing to be friends.”
“Friends!” Gaunt cried contemptuously. “Such canaille as you, my
friend! I would prefer an out and out Apache. But I won’t detain you
from your dinner. Au revoir, Baron.”
And he was immediately ashamed of the outbreak, for he had
done no good save to relieve his feelings. He did not realize that he
had made the Baron thoroughly uncomfortable.
It was decided that they should leave Gaunt House that night and
go straight to Winchester, and there await the wire from Captain
Armstrong to the effect that the Esmeralda was about to sail.
The next day such a message came.
“You have still time to motor to Lynton House,” Edward Drake
suggested as they were getting ready to start for Southampton,
where they intended to stay the night.
“I have told you that I won’t discuss my wife with you,” Gaunt
answered coldly.
“You are very hard, and I think you will regret it. Try to put yourself
in her place. She was sorely tried,” Drake protested quietly.
In reply Gaunt took his wife’s letter from his pocket.
“Read that, and if you are a man you will understand.”
And Drake knew that it was hopeless to make another effort at
reconciliation. But that night was a very miserable one for all three of
them. Each was thinking of the woman that he loved, and
remembering that in the morning they sailed upon a desperate
adventure from which they might never return.
Passages had been booked under fictitious names, and they went
aboard the Saxon just before sailing time. However, there was
considerable delay, and they knew no peace until the last rope had
been thrown ashore.
And then Gaunt turned to Drake.
“We have started for good or ill,” he said with unwonted solemnity.
“It is for good, and God is with us,” Edward Drake answered
reverently.
And then he gave a cry.
“There is a motor-car with two ladies.”
Gaunt strained his eyes towards the shore and his face became
very white when he recognized his wife. He did not move his eyes
from the place where Lady Mildred stood, but his face did not soften.
The liner cleared the point, and began to travel more quickly, so that
in a very few minutes the wharf disappeared from view.
And then he turned away with a sigh and walked to the cabin
which had been reserved for him on the deck. There was a cold wind
blowing, and he closed the door; not till then did the mask fall from
his face. Love tugged at his heart, and he would have given all he
possessed to be face to face with his wife. His anger vanished,
never to return, and his longing for her came back with
overwhelming force. Now he blamed himself bitterly that he had not
gone to her before his departure, for now, many weary weeks must
elapse before he could see her again.
He sat there thinking deeply—of the vow he had made—his
crusade against the rulers of the Congo—and his wife. There was a
knock at the door, and the steward entered to unpack his trunk. Still
he did not move until the bugle sounded the hour for dressing.
Almost mechanically he put on his evening clothes, and went
down to the saloon, where the passengers were busily looking for
their places at the table. Edward Drake and his brother were already
there, but the meal was a very silent one, for each was busy with his
thoughts.
After dinner Gaunt and Edward Drake went to the comfortable
smoking-room on deck, and lighted their cigars.
“Now there can be no turning back,” Edward Drake remarked.
“Do you wish to?” Gaunt demanded abruptly.
“No. Except that I was sorry to leave England without obtaining the
answer to a question which I had asked.”
“I suppose you refer to the time when I interrupted you. I am very
sorry. Drake, you are a good fellow, and I could not wish a woman a
better husband.”
“Thank you,” Drake answered simply. “At one time I never dreamt
of telling Lady Ethel of my love. My brother Lindsay is my rival, and I
thought that I must leave the way clear to him. But love can be
selfish, and I intend to fight for my happiness. But it is hardly credible
that she can care for me.”
“You never can tell where a woman’s feelings are concerned.”
“Lady Ethel was with your wife on the wharf. I am sorry that they
did not come a quarter of an hour earlier. Lady Mildred will be very
miserable.”
“And so will her husband. But we won’t talk of it, Drake. I have
made up my mind to think of one thing, and one thing alone—the
Congo. This may be a rash adventure of ours, but it shall not fail
from any lack of energy on my part. And here’s luck to it,” Gaunt
remarked as he raised his glass to his lips.
As they steamed down the channel the weather grew so rough
that many of the passengers were confined to their cabins. But they
had determined to make no acquaintances, and kept themselves
strictly to themselves, so they were rather glad than otherwise. Upon
reaching the bay of Biscay, as is often the case, the sea was
smoother and they were able to spend more time in the open air. It
was not until the fourth day that anything of moment happened. It
was just after twelve o’clock that they saw smoke on the horizon,
and soon they could make out a steamer that was traveling very
rapidly.
“Looks like one of our small cruisers,” the first officer remarked to
them.
And it was not until the war-ship was but some two or three miles
astern that they knew that she was the Esmeralda.
“She is flying the San Salvador flag,” the officer remarked.
Gaunt ran for his glasses, and fixed them on the vessel of which
he was the temporary owner and his heart beat more rapidly as he
noticed her workmanlike appearance.
“She is doing well over twenty knots, and I should say she carries
turbines. A smart little ship,” the friendly officer continued.
The Esmeralda was quickly overhauling them, and soon signal
flags fluttered on the Saxon which were answered by the cruiser.
“What does she say?” Gaunt asked quickly.
“Only her name. She is not very communicative, and is evidently in
a hurry.”
The smoke was belching from her stacks and she seemed to spurt
forward so that the Saxon was rapidly left behind.
It was late at night when they reached Las Palmas, and they went
ashore at once, to be driven along pebbly streets to the Santa
Catalina Hotel. Immediately upon their arrival Captain Drake left to
visit the men who had already arrived and were awaiting them.
He ascertained from the officer who had been placed in charge
that there had been no mishap, and that they were ready to embark
at any moment.
Breakfast had been ordered for an early hour in the morning, and
they had not been at the table very long before Captain Armstrong
joined them.
“Everything all right?” asked Gaunt.
“Couldn’t be better,” the captain answered briskly. “I have a good
lot of men and they are shaping well. Haven’t had trouble of any
kind.”
“Good. Let us get breakfast over for we mustn’t stay here a
moment longer than we can help.”
Tugs had been chartered, and by eleven o’clock the whole party
had left the harbor. There had been many inquiries from the officials
of the port, which Captain Armstrong had deftly turned aside.
An hour later they reached the Esmeralda; the men were quickly
got on board and the tugs returned to Las Palmas.
Gaunt and the Drakes stood on the quarter-deck and looked
around curiously. Yes. Captain Armstrong had done wonders in the
short time at his disposal.
Six inch guns stared at them from squat turrets, and Edward
Drake looked at Gaunt with something akin to awe.
“I am beginning to realize what it means. Maybe death, but life for
the natives of the Congo,” he said in a low voice.
An order from the bridge and the flag of San Salvador was
lowered. Then an ensign took its place, which was white with a red
cross.
“She is doing close on twenty-five knots,” Captain Armstrong
remarked as he joined them, and there was a look of pride on his
face.
But no one answered, and the men looked at one another.
And Edward Drake sighed.
CHAPTER XXXVII
L ADY MILDRED did not enter the car until the Saxon had
disappeared, and to her sister’s surprise her face was almost
cheerful.
“Captain Samson lives in Southampton and I am going to see him
at once.”
“You really intend to go to the Congo?” Lady Ethel asked quickly.
“Yes, and shall start as soon as possible. The Heron is at
Summers and Payne’s yard, and we will go there and find out
Captain Samson’s address.”
An hour later they were in the house of the captain of Gaunt’s
yacht, and Lady Mildred at once came to the point.
“Captain, how soon can the Heron start on a long voyage?” she
asked.
“A matter of a fortnight, my lady. The engineer is tinkering with his
turbines and has got ’em to pieces,” Captain Samson replied.
“What is the shortest time, working night and day?” she insisted.
“It might be managed in a week, my lady,” the captain answered
reflectively.
“It must be quicker than that, captain. We will sail on Friday next.
Tell the steward to lay in stores for a long voyage, and the bunkers
must be filled up.”
“May I ask where we are going, my lady?”
“First of all to the Canary Islands. See the engineer and have the
work started at once. I shall be at the Station Hotel on Thursday
night, and I shall expect you there. Good-bye, captain.”
And with a cheerful nod Lady Mildred left him and gave the
chauffeur orders to drive back to London.
“I don’t understand you,” Lady Ethel said timidly. “A couple of
hours ago you were utterly miserable, while now you seem almost
happy.”
“I am not sure that I understand myself. I am only thinking that in a
week or two I shall see John. I have no fear how he will receive me,
for I’ve no pride left, and I am prepared to go on my knees and ask
his forgiveness for that cruel letter,” Lady Mildred said earnestly.
“I shall be glad to come with you, dear.”
“I am not sure that I ought to take you, for we cannot catch them
up until they are in the Congo, and there may be danger.”
“I am not afraid,” Lady Ethel answered quickly.
During the next few days they were busy making preparations for
the voyage, and it was not until Thursday that anything of importance
happened. It chanced that they lunched at the Carlton Hotel and
upon reaching the streets they heard the name of Gaunt being
shouted by the newspaper boys.
“Disappearance of John Gaunt. Bail estreated.”
And they read an account of the proceedings at the police court.
Mr. Ruggles, K. C., had been in great form and took full advantage of
the position.
“Listen to this,” Lady Mildred cried, and proceeded to read from
the paper. “The court was crowded this morning at the adjourned
hearing of the charge of murder brought by the Congo government
against Mr. John Gaunt. The magistrate took his seat and the name
of the millionaire was called, but there was no answer. At first it was
believed that he had been delayed by some accident, but when half
an hour had passed, it was murmured that he had left the country.
The magistrate adjourned the court for an hour, and when he again
sat the police gave evidence to the effect that Mr. Gaunt had not
been seen for some few days. Inquiries at his office and at Park
Lane gave no information, and it is assumed that, for some purpose
of his own, he has left the country. Mr. Ruggles proceeded to
address the court: ‘I must remind your Worship that I anticipated this
event and strongly urged that bail should not be allowed. In spite of
my urging you gave him his freedom, of which he appears to have
taken full advantage. I apply to your Worship to grant a warrant for
his arrest.’
“‘Have you anything to say?’ the magistrate asked of Sir Richard
Tester.
“‘No, your Worship. I can only think that some accident has
happened to my client.’
“‘We understand the warrant has been signed and is in the hands
of the police.’”
“They will have to go a long way to execute it,” Lady Mildred
remarked with a smile.
“You do not seem very distressed,” Lady Ethel answered drily.
Her answer was a laugh, and they alighted from the car to be met
by Lord Lynton, who was just coming out of the house.
“I see you have heard the news,” he remarked with a glance at the
paper which she still held in her hand. “I suppose the next we shall
hear will be of his arrest.”
“I don’t think so,” Lady Mildred answered. “Come into the house,
Geoffrey. I want to talk to you.”
Lord Lynton followed to her room, and it was evident that he was
very curious.
“You have not the air of a wife who has just heard that the police
are after her husband,” he remarked drily.
“Geoffrey, give me your word of honor that you will not breathe a
word of what I tell you,” she said earnestly.
“I can keep a secret. It is my business,” he answered.
“Well then, John has gone back to the Congo. Ethel and I are
going to follow him in the Heron. We sail to-morrow.”
“The devil you are!” he blurted out.
“Yes, and I think it would be as well if you came with us.”
“You may as well tell me the whole story.”
“Very well, but remember that it is in confidence,” she answered,
and proceeded to relate all that had happened.
He listened in open-eyed amazement, and when she had finished
he had lost some of his wonted composure.
“It is pure madness, and the man must be stopped,” he said hotly.
“Remember your promise,” she said warningly.
“Of course I will, but the thing is stupendous. It may cause a
European war.”
“Will you come with us? We need not consider Europe. I am only
thinking of my husband.”
“Yes. I will come if I can get leave of absence. When do you
leave?”
“We shall motor to Southampton in a couple of hours’ time.”
“I will be back before that,” he answered, and hastened away.
Sir Keith Hamilton did not keep him waiting very long, and
received him pleasantly.
“I suppose you have come about your brother-in-law. I shall not be
altogether sorry if he has left the country and doesn’t return,” Sir
Keith remarked drily.
“I have come to ask leave of absence for an indefinite time,” Lord
Lynton answered, and his voice was very grave.
“For what purpose?”
“To accompany my sister on her yacht.”
“A pleasure trip?”
“Not exactly, sir. In fact I am bound to silence.”
“I see. Does this voyage concern Gaunt?”
“I can say that our destination is the Congo,” the Earl announced
slowly.
“May I ask if Gaunt will be on board?”
“He will not. But please don’t cross-examine me, sir. I am in a
difficult position, and I must keep my promise.”
“Yes, that is necessary,” Sir Keith said reflectively. “You can have
your leave.”
“Thank you, sir. May I ask if you have confidence in my
judgment?”
“Yes—to a degree,” was the cautious answer.
“I can tell you nothing, but I would suggest that the first cruiser
squadron of the Atlantic fleet be sent to the Canary Islands for a few
weeks.”
“That is a large order,” Sir Keith said with a smile, but his eyes
were filled with curiosity.
“The squadron is now at Gibraltar. You know I would not suggest
this without a weighty reason. Admiral Stretton is a fine sailor, and
something of a diplomatist, I believe.”
The foreign secretary’s eyes did not leave the Earl’s face, and it
was a long time before he spoke.
“I think I may venture to say that the squadron will cruise in the
direction of the Canaries. May I understand that you will
communicate with me when you are at liberty to do so?”
“Yes, sir. At the first opportunity.”
“You can tell me nothing further?”
“I am not sure that I have not already said too much. I am very
anxious, and my responsibility is great.”
“May I ask if Gaunt has already gone to the Congo?” Sir Keith said
quietly.
“I cannot tell you, sir.”
“Do you anticipate international trouble? I must admit that you
have made me rather anxious. Cannot you give any further

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To all the authors for agreeing to face this journey,

To where the wild things are and the children,

To our authors who made this book possible

Contributors to Volume III xiii Division II

Keys to Plecoptera 132 Acknowledgments 226

13. Order Mecoptera 14.3. Odonata: Superfamily

Section II.2 14.4. Superfamily Calopterygoidea

Section II.3 15.4. Family Hydrophilidae

Section II.4 16.4. Family Thaumaleidae

16.9. Family Blephariceridae 16.12. Family Syrphidae

16.10. Family Psychodidae 16.13. Family Periscelididae

studies fish ecology, especially as related to food webs. He

Dr. D. Christopher Rogers is a research zoologist at the

especially those who both study fauna and live in the

Introduction and Subclass

Thorp and Covich’s Freshwater Invertebrates, Fourth Edition. https://doi.org/10.1016/B978-0-12-804223-6.00001-9 3

NATURE AND FAUNA OF THE HOW TO USE THIS VOLUME

Hexapoda: Insecta: Orders

FIGURE 1.4 Example of an adult aquatic beetle (Coleoptera, Cneo-

FIGURE 1.9 Lateral view of the larva of the Cheumatopsyche

FIGURE 1.10 Dorsal view of a larval Ilyobius sp. (Megaloptera,

Chapter Outline Collembola: Poduromorpha: Onychiuridae: Genera 48

INTRODUCTION to a surface and used Latin/Greek terminology to system-

Thorp and Covich’s Freshwater Invertebrates, Fourth Edition. https://doi.org/10.1016/B978-0-12-804223-6.00002-0 11

T HERE was a feeling of constraint when Gaunt led Edward Drake

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