An Introduction to the Aquatic Insects of North America [5 ed.] 1524968544, 9781524968540

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An Introduction to the

AQUATIC INSECTS of NORTH AMERICA

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Edited by

I R.W. Merritt

K.W. Cummins

M.B. Berg Ui:

Front Cover Photo:

Roaring Fork Creek, Great Smokey Mountain National Park, Tennessee Photo by Keith Kennedy, Raleigh, North Carolina

Insect on Back Cover:

Odonata: Calopterygidae (Calopteryx maculata), Rose Lake, MI

Photo by F. William Ravlin, Okemos, Michigan

Back Cover Editors Photo:

Cordova, Alaska

Photo by Gary A. Lamberti, Notre Dame, Indiana

Kendall Hunt pub l i sh i ng

company

www.kendallhunt.com

Send all inquiries to: 4050 Westmark Drive

Dubuque,lA 52004-1840

Copyright © 1978, 1984,1996, 2008, 2019 by Kendall Hunt Publishing Company ISBN 978-1-5249-6854-0

All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording, or otherwise, without the prior written permission of the copyright owner. Published in the United States of America

DEDICATION We would like to dedicate this 5th edition of our book to the contributors who have

passed since the 4th edition was published in 2008. These individuals were not just authors, but many were close friends of ours and their hard work and dedication to the field of Aquatic Entomology have helped make this book a success over the past 40 years. We have listed their names in alphabetical order: Norman H. Anderson, George W.Byers, Kenneth A. Christiansen, William P. Coffman, Clyde H. Eriksen, Oliver S. Flint, John T. Polhemus, Robert E. Roughley, Kenneth W. Stewart, and Glenn B. Wiggins. These indi viduals will not be forgotten.

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CONTENTS Preface xi Acknowled^iments xiii List of Contributors xvii CHAPTER 1 Introduction

1

R. W. Merritt, K. W. Cummins, and M. B. Berg CHAPTER 2

General Morphology of Aquatic Insects 9 M. B. Berg, K. W. Cummins,and R. W. Merritt CHAPTER 3

Sampling Aquatic Insects: Collection Devices, Statistical Considerations, and Rearing Procedmes 17 J. K. Jackson, V. H. Resh, D. P. Batzer, R. W. Merritt and K. W. Cummins CHAPTER 4

Aquatic Insect Respiration 43 D. B, Buchwalter, V. H. Resh, G. A. Lamberti, and W.C.E.P. Verberk CHAPTER 5

Habitat, Life History, Secondary Production, and Behavioral Adaptations of Aquatic Insects 65 A. D. Huryn and J. B. Wallace CHAPTER 6

Ecology and Distribution of Aquatic Insects 117 K. W. Cummins, R. W. Merritt, and M. B. Berg

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Contents

CHAPTER 7

Use of Aquatic Insects in Bioassessment 141 R. D. Mazor, D. M. Rosenberg and V. H. Resh CHAPTER 8

An Overview of the Aquatic Insect Ecological Tables 165 M. E. Benbow, J. P. Receveur, and S. Nowak CHAPTER 9

Adaptations and Phylogeny of Aquatic Insects 175 K. W. Will and V. H. Resh

CHAPTER 10

Aquatic Insects of North America: A Photographic Overview 193 G. W. Courtney and S. A. Marshall CHAPTER 11

General Classification and Key to the Orders of Aquatic and Semiaquatic Insects 231 G. L. Parsons

CHAPTER 12

Aquatic Collembola 245 R. J. Snider CHAPTER 13

Ephemeroptera 263 S. K. Burian

CHAPTER 14 Odonata

341

K. J. Tennessen CHAPTER 15

Semiaquatic Orthoptera 411 H. Song

Contents

CHAPTER 16

Plecoptera 429 R. E. DeWalt and B. C. Kondratieff

CHAPTER 17

Aquatic and Semiaquatic Hemiptera 521 D. A. Polhemus

CHAPTER 18

Megaloptera and Aquatic Neuroptera 569 D. E. Bowles and A. Contreras-Ramos

CHAPTER 19

Trichoptera 585 J. C. Morse, R. W. Holzenthal, D. R. Robertson, A. K. Rasmussen, and D. C. Currie CHAPTER 20

Aquatic and Semiaquatic Lepidoptera 765 M. A. Soils

CHAPTER 21

Aquatic Coleoptera 791 A. E. Z. Short and D. S. White

CHAPTER 22

Aquatic Hymenoptera 909 A. M. R. Bennett

CHAPTER 23

Aquatic Diptera 925 G. W. Courtney CHAPTER 24

Tipuloidea 1023 J. K. Gelhaus and V. Podeniene

ix

Contents

CHAPTER 25

~'iliidd^*1071 J. R. Wallace CHAPTER 26 Simixliidae

1097

P. H. Adler and D. C. Currie

CHAPTER 27

Chironomidae

1119

L. C. Ferrington, Jr. and M. B. Berg

Glossary 1275 B. W. Merritt

Bibliography 1289 Index

1455

PREFACE Another decade has passed, and it is time for the revised 5th edition of An Introduction to the Aquatic Insects of North America. Ken, Marty, and I are excited about the new and revised additions to this

new edition. When first published in 1978, the book had 22 authors, keys only to the family level, and 1,712 references. This 5th edition has 45 authors, expanded generic level keys,and over 7,000 references.

Sadly, nine authors have passed since the last edition and are listed in the Dedication. We have added 18

new authors in this 5th edition. As with previous editions, this new one is intended to serve as a standard

guide to the aquatic and semiaquatic insects of North America,including keys to the immatures and in most cases adults, with pupal keys to the Trichoptera, Diptera, Culicidae, Simuliidae, and Chironomidae.

There have been substantial additions and expanded coverage to some of the introductory chapters, especially the Bioassessment, Respiration, Habitat, and Life History chapters. A separate chapter on Ecological Tables of Aquatic Insects, the hallmark of the first four editions, has been added to summarize

and elaborate on the ecological information for each taxon,as well to update,revise and expand the content.

In addition,a new chapter on A Photographic Overview ofAquatic Insects of North America has been added to the book. This chapter includes outstanding color

photographs of the majority of aquatic insect families as a supplement to identification by two of the best insect photographers in North America (Courtney and Marshall). Important changes have been made, including revision and expansion of keys, along with new figures added to the taxonomic chapters. Figures have also been added to the General Classification and Key to Orders chapter so that students do not need to refer to figures in other parts of the book when keying out aquatic insects to Order level. There have been particularly significant revisions to the chapters on Ephemeroptera, Plecoptera, Trichoptera, Coleoptera, Diptera,and Tipuloidea.The Trichoptera and Diptera chapters have each been combined into one chapter for each order. Larvae, pupae,and adults together are treated as one chapter for each order. As with any comprehensive treatment on aquatic insects, coverage ofthe literature can only be partial because it continues to grow exponentially. This is a true measure of the popularity of the subject matter. We added to the references submitted by the authors by surveying other literature sources. As before, we strongly encourage users of the book to continually update material in their own areas of interest. Also, we hope this new edition will be of even greater use to both professional and lay groups interested in aquatic insects.

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ACKNOWLEDGMENTS The editors would like to thank all the contribu

tors, both old and new, for their cooperation during the production of the 5th edition of this book. This new edition would have not been possible without their help and expertise. We also would like to thank our publisher, Kendall/Hunt Publishing Company, Dubuque, Iowa for their cooperation and patience with this endeavor. They have been our publisher since the first edition in 1978 and we have developed a good working relationship over the years to help make this book a success. At MSU, we would like to

Acknowledgments, thanks, and credits by con tributors and the editors for specific chapters are as follows:

Chapter 7: Biomonitoring We thank Marcus Beck, Dave Buchwalter, Joyce Chou, Matthew Cover, Cody Fees, David Gillett, Charles Hawkins, Ryan King, Katrina Krievins, Jason May, Alvina Mehinto, Patina Mendez, Peter Ode, Alison O'Dowd, Ashley Park, Andrew Rehn, Eric Stein, Stephanie Strachan, Susanna Theroux,

thank Dr. Bill Ravlin, Chairperson of Entomology, MSU,for his continued support of this project over the past 2.5 years. We would especially like to thank

and Rebecca Willison.

Mr. Scooter Nowak for assistance with the computer program to merge old and new references and all

Overview

other issues dealing with computer programming in this edition.

I(RWM)would like to thank my wife Pam for

her continued patience during another edition of this book and my close friends and colleagues, Gary Lamberti, Eric Benbow, Marty Berg, and Johnny Wallace who continually harassed me along the way with their texting! I (KWC) Over 60 years as an aquatic ecologist there have been so many who have helped, collabo rated and inspired me along the way. The list is too long to include here, but I would single out Noel Hynes, the two other editors of this book. Rich and Marty, and Margaret Wilzbach, Clyde Eriksen, Bill Coffman, and Bob Peterson.

I (MBB), first and foremost, thank my wife Pat

for all of her support and for enduring yet another edition of the book. I also thank my children, Juliana and Ethan, for their understanding, patience, and for agreeing not to ask "Is it done yet?" Finally, I thank Rich and Ken for the opportunity to join them on this and the previous edition, and Gary Lamberti, my good friend and valued colleague for his sage advice over the years.

Chapter 10: Aquatic Insects of N. A.: A Photographic We are grateful to numerous colleagues for shar ing their knowledge and advice on certain families of Ephemeroptera (B.C. Kondratieff, D. Lemkuhl, and J. Webb) and Trichoptera (N.H. Anderson, D.E.Bowles,J. Giersch,L.Myers,and R.W.Wisseman). Without their generous assistance, we would have

been unable to locate and photograph many fami lies included in this chapter. Several individuals graciously helped with identification of our images: Ephemeroptera (S. Burian), Odonata (Ken Tennessen), Plecoptera (B.C. Kondratieff and R.E. DeWalt), Coleoptera(D.R. Maddison and A. Short), Trichop tera (J.C. Morse, A.K. Rasmussen, D. Ruiter, and R.W. Wisseman), and selected Diptera (A. Fasbender and B.J. Sinclair). Jon K. Gelhaus kindly provided specimens of larval Phalacrocera (Cylindrotomidae)

for us to photograph. We thank the following indi viduals for allowing use of their images: C.R. Nelson (Ephemeroptera: Euthyplociidae), M. Garrison (Odonata: Corduliidae), Ken Tennessen (Odonata: Lestidae), and U.G. Neiss(Odonata: Platystictidae), and J.C. (Skip) Hodges, Jr. (Trichoptera: Dipseudopsidae, Ecnomidae, and Xiphocentronidae). Field work associated with this chapter was supported in part by a National Science Foundation grant

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Acknowledgments

(DEB-0933218)to G.W.Courtney and the National Institute of Food and Agriculture, Project No's. 6693 and 5473.

Chapter 13: Ephemeroptera

The keys to nymphs and adults in this chapter were originally developed based on those presented by Edmunds et al. (1976) and first appeared in the second edition of this book. Since then the keys have changed considerably with each edition ofthe book as did our knowledge of the taxonomy of North Ameri can mayflies. 1 am indebted to George Edmunds, Jr. and Robert Waltz for all of their great work on pre vious editions of this chapter. The keys in this edition also have benefited from the generous contributions by bench taxonomists and aquatic biologists that use keys as part of their jobs and discovered problems or observed variations in characters that created stum

bling blocks in parts ofthe previous keys. I also want to thank all my mayfly colleagues that were willing to share their ideas and time discussing some of the per sistent problems still plaguing mayfly taxonomy rele vant to the keys presented here. In this edition there are several new figures, but most of the figures from the previous edition are retained. I grant permission to use figures from my publications; Fig. 13.229 (Burian 2001) and Fig. 13.234 (Burian 1995). I am grateful to the University of Min nesota Press for the continued use of figures from Edmunds et al. (1976). I greatly appreciate the permis sion of Dr. R.D. Waltz to continue to use illustrations

from his publications: Fig. 13.255 (Waltz and McCafferty 1989); Fig. 13.263(Waltz and McCafferty 1987a), Fig. 13.108(Waltz eta/. 1985); Fig. 13.49(Waltz 2002); Fig. 13.124-13.125(Waltz and McCafferty 1985); Figs. 13.116 and 13.261 (Waltz and McCafferty 1999). The late Dr. R.K. Allen allowed us to republish Figures 13.79 (Allen 1974), 13.90, 13.92 (Allen 1973), 13.91 (Allen 1976), and 13.246 (Allen 1965). Figures 13.80, 13.223-13.227, 13.230, and 13.267 are from Burks(1953)and are published with permission of the Illinois Natural History Survey. Figure 13.44 is from Needham et al. (1935) and is published courtesy of Cornell University Press. Figure 13.264 is from Provonsha and McCafferty(1982),courtesy ofthe authors. Figures 13.119, 13.269, and 13.271 are from Davis (1987). Figures 13.120, 13.122 are from Peters (1971) with permission ofthe author. Figures 13.66-13.68 are from Bednarik and McCafferty (1979), courtesy of the Canadian Bulletin of Fisheries and Aquatic Sciences. The late Dr. G.F. Edmunds, Jr. provided Figs. 13.83-13.84, 13.240 (Allen and Edmunds (1962); Fig. 13.243 (Allen and Edmunds 1965);

Figs. 13.244-13.245 (Allen and Edmunds 1963a); Fig. 13.246 (Allen and Edmunds 1963b); Fig. 13.72 (Bednarik and Edmunds 1980); and Fig. 13.76(Traver and Edmunds 1967). Figures 13.93-13.105 and Figs. 13.197-13.198, 13.202-13.207 are taken from Wiersema and McCafferty (2000); Figs. 13.265-13.266, Figs. 13.268, 13.270 are taken from McCafferty and Provonsha (1985), courtesy of the authors; Fig 13.259-13.260 are taken from McCafferty and Lugo-Ortiz (1998); Fig 13.123 is taken from LugoOrtiz and McCafferty (1998c); Figs. 13.179-13.180 are taken from Lehmkuhl (1976); Figs. 13.177-13.178, 13.152 are taken from Kluge (2004); Figs. 13.77 and 13.205 are taken from Flowers and Dominguez(1992). Chapter 14: Odonata

John C. Abbott (University of Alabama, Tuscaloosa, Alabama)for providing a wing scan of Leptobasis and proofreading the draft for the 4th edition; Maria C. Garrison (McHenry County College, Illinois) for proofreading and critiquing the entire chapter for the 5th edition, as her comments and questions resulted in many corrections and improve ments in the draft manuscript.

Chapter 16: Plecoptera We graciously acknowledge that the following keys to adult and nymphal stoneflies are based upon the groundbreaking work of Harper and Stewart (1984), Stewart and Harper(1996), Stewart and Stark (2002), and Stewart and Stark (2008). Their contribu tions were influenced by regional keys to stoneflies by Jewett (1956) and Hitchcock (1974). We have incor porated the new stonefly genera proposed since the last edition and made other modifications suggested by colleagues and students. The update of this chapter has built a solid foun dation for its future improvement. Repeated photo copy reproduction of previous nymphal illustrations degraded image quality. Therefore, we replaced most of the nymphal illustrations of Stewart and Stark (2008) with those scanned from Stewart and Stark (2002). Adult images from the 4th edition were near pristine, requiring only scanning with minor editing. All images are now saved as tiffs and deposited for safe keeping. Many new figures were added from var ious sources and we extensively edited the keys to improve their usefulness. New references have been added and some older ones retired.

Chapter 19: Trichoptera Larval and pupal keys are based on the work of Wiggins and Currie(2008). Assistancefrom D.E.Ruiter

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is gratefully acknowledged and we would like to thank James C.(Skip) Hodges, Jr. for allowing us to use his excellent caddisfly case photographs.

Acknowledgments

J.K. Moulton (Dixidae), and B.J. Sinclair (Empidoidea). 1 am especially grateful to Brad Sinclair for his revision of all couplets pertaining to empidoid flies. I would also like to thank the scientists at

Chapter 21: Coleoptera

Rob Roughley contributed extensively as the senior author of the 4th edition. We thank Stephen Baca (Noteridae), Grey Gustafson (Gyrinidae), Crys tal Maier (Lutrochidae), and Phil Perkins (Hydraenidae)for reviewing various portions ofthe text and key for the 5th edition.

Chapter 23: Diptera

I wish to acknowledge numerous colleagues for sharing their knowledge and advice on selected Dip tera, especially A. Borkent (Ceratopogonidae), G.R. Curler (Psychodidae), J.K. Gelhaus (Tipuloidea),

Rhithron Associates, Inc. for beta-testing the keys. This work was supported in part by the National Science Foundation (grants DEB-0933218 and EF-1115156) and the National Institute of Food and Agriculture, Project No's. 6693 and 5473. Chapter 24: Tipuloidea

We acknowledge a great debt to the late Prof. George Byers, University of Kansas. His extensive studies ofthe North American Tipuloidea, spanning more than 50 years, and his development of the first comprehensive keys to aquatic crane fly larvae, provide much of the basis for the present keys(Gelhaus et al. 2018).

LIST OF CONTRIBUTORS P. H.ADLER Department of Plant and Environmental Sciences, Clemson University, Clemson, SC 29634 D. P. BATZER Department of Entomology, Univer sity of Georgia, Athens, GA 30602 M. E. BENBOW Department of Entomology and Department of Osteopathic Medical Specialties, Michigan State University, 243 Nat. Sci. Bldg. 288 Farm Lane, East Lansing, MI 48824 A. M. R. BENNETT Canadian National Collection

of Insects, Arachnids and Nematodes, Agricul ture and Agri-Food Canada,960 Carling Avenue Ottawa, Ontario Canada KIA 0C6

M.B.BERG Department of Biology,Loyola University Chicago, 1032 W. Sheridan Rd., Chicago, II60660 D. E. BOWLES Department of Biology, Missouri State University, Springfield, MO 65897 D. B. BUCHWALTER Department of Environmen tal & Molecular Toxicology, Campus Box 7633, NC State University, Raleigh, NC 27695-7633 S. K. BURIAN Department of Biology, Southern Connecticut State University, 501 Crescent St., New Haven, CT 06515 A. CONTRERAS-RAMOS Instituto de Biologia,

UNAM,Depto. de Zoologia,Apdo Postal 70-153, 04510 Ciudad de Mexico, Mexico

G. W.COURTNEY Department of Entomology,Iowa State University, 401 Science II, Ames,IA 50011 K W. CUMMINS California Cooperative Fisheries Research Unit,Humboldt State University, Arcata, CA 95521

R. W. HOLZENTHAL Department of Entomology, Hodson Hall, 1980 Folwell Ave., University of Minnesota, St. Paul, MN 55108

A. D. HURYN Department of Biology, University of Alabama, 2107 Bevill Building, Box 870206, Tuscaloosa, AL 35487 J. K. JACKSON Stroud Water Research Center, 970

Spencer Road, Avondale, PA 19311 B. C. KONDRATIEFF Colorado State University, Department of Bioagricultural Sciences and Pest Management, 1177 Campus Delivery, Fort Collins, CO 80523

G. A. LAMBERTI Department of Biological Sci ences, University of Notre Dame, Notre Dame, IN 46556-0369

S. A. MARSHALL University of Guelph Insect Col lection and Insect Systematics Laboratory, School of Environmental Sciences (Bovey), 1216 Edmund C. Bovey Building, University of Guelph, Guelph, ON,Canada NIG 2W1 R. D. MAZOR Southern California Coastal Water

Research Project, 3535 Harbor Blvd, Suite 110, Costa Mesa, CA 92626 R. W. MERRITT 1005 Cormorant Terrace, The Villages, EE 32162

B. W. MERRITT Via dei Gilardi 17,6926 Montagnola, Switzerland

J. C. MORSE Department of Plant and Environ mental Sciences, Clemson University, Clemson, SC 29634

C. CURRIE Department of Natural History, Royal Ontario Museum, 100 Queen's Park Toronto, ON, Canada M5S 2C6 R.E. DEWALT University of Illinois Prairie Research

S. NOWAK School of Informatics, Computing and

Institute, Illinois Natural History Survey, 1816 S. Oak St., Champaign, IE 61820 L. C. FERRINGTON,JR. Department of Entomol ogy, Hodson Hall, 1980 Folwell Avenue, Univer sity of Minnesota, St. Paul, MN 55108 J. K. GELHAUS Department of Entomology, The Academy of Natural Sciences of Drexel University, 1900 Benjamin Franklin Parkway, Philadelphia,

G. E. PARSONS Department of Entomology, Michigan State University, 243 Nat. Sci. Bldg., 288 Farm Lane, East Lansing, MI 48824 V. PODENIENE Institute of Biology, Life Sciences Center, Vilnius University, Sauletekio str. 7,

D

PA 19103

Cyber Systems, Northern Arizona University, Building 90, 1295 S. Knoles Dr., Flagstaff, AZ 86011

LT-10257 Vilnius, Lithuania

D. A.POLHEMUS Department of Natural Sciences, Bishop Museum, 1525 Bernice St., Honolulu, HI, 96817

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List of Contributors

o A.K.RASMUSSEN Center for Water Resources, 113 S. Perry-Paige Building, 1740 S. Martin Luther King Jr. Blvd., Florida A&M University, Tallahassee, FL 32307 J. P. RFCFVFUR Department of Entomology,

Michigan State University, 243 Nat. Sci. Bldg., 288 Farm Lane, Fast Lansing, MI 48824 V. H. RFSH University of California, FSPM Department, Organisms & Environment Divi sion, Berkeley, CA 94720 D. R. ROBERTSON Integrative Research Center, Field Museum of Natural History, 1400 S. Lake Shore Drive, Chicago,IL 60605 D. M. ROSENBERG 280 Waverley St., Winnipeg, MB,Canada R3M 3L3

A. E. Z. SHORT Department of Ecology and Evolu tionary Biology, University of Kansas, 6002 Haworth Hall, Lawrence, KS 66045

R. J. SNIDER Department of Integrative Biology, Michigan State University, 203 Nat. Sci. Bldg., 288 Farm Lane, East Lansing, ML 48824

M.A.SOLIS SEL,USDA,Smithsonian Institution, P.O. Box 37012, National Museum Natural History, E-517, MRC 168, Washington, DC 20013-7012

H. SONG Department of Entomology, Texas A&M University, College Station, TX 77843-2475 K. J. TENNESSEN P.O. Box 585, Wautoma, WI 54982

J. B. WALLACE Department of Entomology and Odum School of Ecology, University of Georgia, Athens, GA 30602

J. R. WALLACE Department of Biology, Millersville University, Millersville, PA 17551 D.S. WHITE Hancock Biological Station, Murray State University, 561 Emma Drive, Murray, KY 42071 W. C. E. P. VERBERK Department of Animal Ecology and Physiology, Radboud University, Nijmegen, The Netherlands K. W. WILL University of California, ESPM

Department, Organisms & Environment Division, Berkeley, CA 94720

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INTRODUCTION Richard W. Merritt

Michigan State University, East Lansing

Martin B. Berg Loyola University Chicago, Illinois

Kenneth W. Cummins

Humboldt State University, Arcata

The emphasis on aquatic insect studies, which has expanded exponentially in the last five decades, has been largely ecological. This interest in aquatic insects has grown from early limnological roots (e.g., Forbes 1887) and sport fishery-related investigations of the '30s and '40s (e.g., Needham 1934), to the use of aquatic insects as indicators of water quality during the '50s and '60s (e.g., Kuehne 1962; Bartsch and Ingram 1966; Wilhm and Dorris 1968; Warren 1971; Cairns and Pratt 1993). In the '70s and '80s, aquatic insects became the dominant forms used in freshwater

investigations of basic ecological questions (e.g., Barnes and Minshall 1983). During the past 10 years (2008-2018), emerging studies have dealt with inva sions of alien invertebrate species, and forecasting responses of benthic insect community structure and function to anthropogenic climate change (Poff et al. 2010; Strayer 2010; Ricciardi 2015; Fenoglio et al. 2016). Ecological applications of observed habitat affinities and traits associated with the physical tem plate ofstreams will become increasingly important in predicting how aquatic insects respond to changing hydroclimate and flow regime (Pyne and Poff 2017; Flerbst et al. 2018). In addition, DNA barcoding also will likely become more widely used in the identifica tion of aquatic insects (e.g., DeWalt 2010). Our expanding knowledge of biodiversity and the role that different aquatic insects play in water quality assess ment is the only way to sustainably manage ecosys tems in an ever changing global environment(Foottit and Adler 2009; Dijkstra et al. 2014). The work on aquatic insects has embraced most major areas of ecological inquiry including population dynamics, predator-prey interactions, physiological and trophic ecology,competition(Resh and Rosenberg 1984; Allan 1995), and management applications of this basic research (Wright eta/. 1991; Rosenberg and Resh 1993a; Dodds 2002; Benke and Gushing 2005).

In addition, fly anglers have enthusiastically sought more knowledge about aquatic insects, both as fish foods to be imitated and as interesting cohabitants with their quarry (e.g., Swisher and Richards, 1971, 1991, 2018; Schweibert 1973; Caucci and Nastasi 1975,2004;

Borger 1980, 1995; LaFontaine 1981; Whitlock 1982, 2014; Hafele and Roederer 1995; Knopp and Cormier 1997; Ames 2005; Fauceglia 2005; Weamer 2017). More recently, aquatic entomology and its applications have experienced many improved and/or new methods, methodologies, and coupled technologies (DeWalt 2010; Hauer and Lamberti 2017; Lamberti and Hauer 2017).

Initially, the primaryjustification for this book was that the systematics ofaquatic insects had lagged behind the needs ofaquatic ecologists and water managers,and the inquisitiveness of anglers. This is still true, but to a lesser extent, although the sophistication of scientists, managers, and anglers perpetuates the need for ever better taxonomic and ecological treatments. Aquatic insects also are of concern to those involved in teaching (e.g., Resh and Rosenberg 1979; Li and Barbour 2011; Hauer and Lamberti 2017; Lamberti and Hauer 2017; Merritteta/. 2017; Gushing

2016), and in outdoor recreation activities because certain groups (e.g., mosquitoes, black flies, horse flies) are frequently pests of humans and other ani mals in water-based environments (Kim and Merritt 1987; Malmqvist et al. 2004; Lemelin 2013). Identifi cation is the first step toward a basic understanding of the biology and ecology of aquatic insects that even tually allows for the development of proper manage

ment strategies. The amateur naturalist and primary, secondary, and post-secondary school educators also require basic identification as an important initial step in familiarization. Thus, for all concerned, iden tification and basic ecological and life history infor mation is important for categorizing the aquatic 1

Chapter 1

Introduction

insects collected. This 5th edition continues to offer

information on functional adaptations of aquatic insects that allows an additional tool for categorizing aquatic insects (e.g., Chapter 6). A number of well-known general works(Usinger 1956a; Edmondson 1959; Klots 1966; Pennak 2001) and specific studies(Ross 1944; Burks 1953)are taxonomically and ecologically, at least for the most part, out-of-date. More recent comprehensive treatments of the Odonata (Westfall and May 1996; Needham et al. 2000), Plecoptera (Stewart and Stark 2002), and Trichoptera(Wiggins 1996)currently are available, as are some general works (e.g., Stehr 1987,1991; Thorp and Covich 2010; Thorp and Rogers 2016). However, at the present time only this 5th edition gathers together comprehensive, updated, generic treatments of immature and adult stages of aquatic and semiaquatic insects of North America. This edition is intended, as were previous editions, to serve as a stan dard reference on the biology and ecology of aquatic

insects with updated keys to separate life stages of all major taxonomic groupings. To this aim, we have provided additional color photographs of most all families of immature aquatic insects to assist the stu dent and professional with correct identifications. The taxonomic coverage is coupled with summa ries of related information on aquatic insect phylogeny,ecology,water quality,bioassessment,respiration, sampling, rearing, life history, and behavior. Generic keys to immatures and adults are provided for all but a few groups of Diptera. Further, pupal keys are now

provided for most holometabolous orders. All the keys have been revised, and very significant revisions have been made to many groups, such as the Ephemeroptera, Odonata, Plecoptera, Trichoptera, aquatic Coleoptera, and Diptera. Because of the size of the

order Diptera,separate chapters have been devoted to individual families or superfamilies (i.e., Chironomidae, Simuliidae, Culicidae, Tipuloidea). The distinction between aquatic or semi-aquatic and terrestrial insects is arbitrary. In this 5th edition, those orders and families with one or more life stages associated with aquatic habitats and frequently encoun tered in collections made from aquatic environments are covered. This includes the Collembola,Orthoptera, and Hymenoptera, even though they are only margin ally associated with aquatic habitats. Because terrestrial insects frequently become trapped in the surface film of aquatic systems (e.g., Collembola), a wide range of

terrestrial species are encountered with varying fre quency. A specimen not fitting the keys in this edition probably belongs to a terrestrial taxon and generally can be identified using Triplehom and Johnson (2005). An annotated list of general references to works dealing with aquatic insect taxonomy and ecology is given in Table 1. As indicated above, many of the taxonomic works are outdated; however,they include a great deal of useful biological information on the groups. More specific references are given at the end of the appropriate order (or family) chapter. Various combinations of taxa can be categorized so as to permit ecological questions to be addressed at the functional level. For example, some groups are based on morpho-behavioral adaptations for food gathering, habitat selection, or habits of attachment, concealment, and movement(Chapter 6, and ecolog ical tables at the end of each taxonomic chapter). Different levels of taxonomic identification are

required to functionally classify aquatic insects. For example, the ordinal level may be sufficient to define

functional trophic relations for the Odonata, but even the generic level may be insufficient in some of the Chironomidae (Diptera). Fcologists also have resorted to "habitat taxonomy" of a single aquatic system (e.g., Coffman et al. 1971) or regionalized keys (e.g., Brigham et al. 1982; Peckarsky et al. 1990; Ward and Kondratieff 1992; Bouchard 2004; Hudson

et al. 2012; Morse et al. 2017), where the fauna of a given system or region is studied for an extended period in sufficient detail to permit such system-spe cific keys to be written. Although this allows for sig nificant simplification in such keys by excluding taxa from other systems or regions, changes in species composition, the key element in disturbance or intro duction of exotics can be masked by the restricted nature of this approach. This means that such keys must be used cautiously and verified continually against the full range of taxonomic information available.

The particular emphasis on ecology and field techniques in all previous four editions of this book has reflected our conviction that the most critical task

at hand is the integration oftaxonomic and ecological approaches. This integrated approach will permit important questions concerning environmental qual ity and management to be addressed. It is our hope that this expanded 5th edition will provide another significant step toward this goal.

Chapter 1

Introduction

Table 1 Selected North American literature dealing with general aquatic insect identification and ecology. Taxonomic GENERAL COVERAGE

Treatments Source

Biology

Immatures Adults

Ward and Whipple

General Comments

x

Generic

Contains much information on biology.

Chu(1949)

X

Family

Generalized treatment of immatures.

Peterson (1951)

x

Family

Limited to hoiometabolous groups; descriptive in nature.

Usinger (1956a)

x

(1918)

x^

Primarily generic level,

x

with keys to Calif, species Edmondson (1959)

x

X*

Considerable information on West Coast

species. A standard reference on freshwater

Generic

invertebrates; some keys outdated. Eddy and Hodson (1961)

x

Needham and Needham

x

X*

X

X*

Order

Keys to common animals, including water

Generic

Keys to many genera; field manual.

mites, of the North Central states. (1962) Klots(1966)

Primarily family level, with

x^

Field manual; some keys based on ecology and behavior of group.

keys to some genera Borror and White (1970)

x^

Swisher and Richards

X

Orders, some keys to families

x

Comprehensive field manual on insects; primarily based on examination of insects in the hand; coior plates.

X

Generic (mayflies only)

x

Anglers' guide, primarily to mayflies; color photographs; seasonal data.

None

X

Extensive treatment of immature aquatic insects for anglers; color plates; seasonal

None

x

(1971, 2018) Schweibert(1973)

and distributional data. Caucci and Nastasi

X

(1975)

Anglers' guide to mayflies and stonefiies; color photographs; seasonal and distributional data.

Parrish (1975)

Keys only to Southeastern United States; limited to water quality indicator organisms.

Generic

x

Smith and Carlton

Primarily family level, with

x^

keys to some species

(1975) Tarter (1976)

x*

Merritt and Cummins

x

Generic

x^

Keys only to West Virginia genera and occasionally species.

Orders, families,

x

Chapters on morphology, ecology, phylogeny, life history, behavior, biomonitoring, and sampling; summary tables on ecology and N.A. distribution with

and genera of North American aquatic insects

(1978, 1984, 1996), Merritt ef a/. (2008)

Keys only to intertidal insects of the central California coast.

references.

Pennak (1978, 1989, 2001)

x

X*

Generic

x

Extensive treatment on biology of many freshwater invertebrates. Editions 1989 and

2001 do not treat aquatic insects. Lehmkuhl (1979a)

X*

Borger(1980, 1995) Hilsenhoff (1981)

X*

Families

x^

Field guide to aquatic insects.

Orders and families

x'^

An angler's guide to the major food organisms of trout and their presentations.

Generic

x^

Keys only to Wisconsin genera, but generally applicable to Great Lakes region.

*Only adult keys to Hemiptera and Coleoptera.

^Contains notes on biology or ecology. ^Covers adults of some groups. ^Primarily adult coverage, brief treatment of immatures of some groups.

(continued)

Chapter 1

Table 1

Introduction

Continued

Taxonomic GENERAL COVERAGE

Treatments Source

Biology

Immatures Adults

LaFontaine (1981)

McCafferty (1981)

General Comments

None

An angler's guide to the caddisflies; good biological section on caddis.

Pictorial keys to aquatic

A thorough scientific introduction to aquatic insects for the fly anglers; excellent

insect families

illustrations.

Brigham etal.(1982)

A thorough treatment of the aquatic insects and oligochaetes of the Carolines.

Families and genera for eastern North America;

species for the Carolines

Leiser and Boyle (1982)

None

Good information on stonefly biology and emergence patterns for anglers.

Whitlock(1982, 2014)

None

Good practical books on fly-fishing entomology.

Orders and families of

A thorough treatment of terrestrial and aquatic immatures with notes on relationships. Literature sources provided.

Stehr(1987, 1991)

terrestrial and aquatic insect immatures

Intended as text for classes on immature insects. Excellent illustrations.

Arbona(1989)

X

Good section on mayfly biology for anglers.

None

Borror etal.(1989)

X

Families

Generalized treatment of adults.

Guthrie (1989)

X

Family keys to animals

Practical guide to animals found at the

found at the water surface

surface of freshwaters, including some excellent photos and drawings: Good biological information.

Families and genera of

Good regional treatment of aquatic

Peckarsky etal.(1990)

Pobst(1990)

immature freshwater

macroinvertebrates of northeastern North

macroinvertebrates

America.

A streamside guide for anglers to the major

None

trout-stream insects with excellent color

photos. Clifford (1991)

Swisher and Richards

Pictorial keys to families and

x"*^

Written for aquatic invertebrates of

genera of immature aquatic

Alberta, Canada, but much broader

insects and other invertebrates

coverage. Excellent drawings and color photographs.

None

A good presentation on the emergence of mayflies, caddisflies, and stoneflies for anglers.

(1991)

Thorp and Covich

Orders and families of

Emphasis mainly on freshwater

(1991, 2001, 2010)

aquatic insects and

invertebrates, other than insects.

differential taxonomic

Comprehensive treatment.

treatment of other

invertebrates Ward and Kondratieff

Orders, families and some

Very useful guide with illustrated keys to

(1992)

genera of selected aquatic

mountain stream insects of Colorado.

insects

*Oniy adult keys to Hemiptera and Coleoptera. ^Contains notes on biology or ecology. *Covers adults of some groups. (continued)

Chapter 1

Table 1

Introduction

Continued

Taxonomic

GENERAL COVERAGE

Treatments Source

Biology

Immatures Adults

General Comments

Hafele and Roederer (1995)

x

x*

Order

Introductory angler's guide; seasonal and distributional data; fishing strategies based on insect emergence patterns.

Knopp and Cormier (1997)

x

X*

None

Excellent descriptions and drawings, no photos, of nymphs to the spinner stage of mayflies. Good biological information on life cycle, behavior, imitative patterns; geared for the serious angler interested in all aspects of mayfly biology and

X*

Common families

Covers temperate Australia but applicable to temperate N.A.; color photographs of live specimens.

X*

Order and families

identification. Gooderham and

Tsyrlin (2002) Voshell (2002)

Describes 100 most common invertebrate

groups; quality color illustrations. Wichard etal.(2002)

X

None

An overview of the numerous adaptations of aquatic insects to life in an aquatic environment with more than 900 scanning electron microscope photographs. The basic functions of an aquatic mode of life, e.g., respiration and osmoregulation, are described for all of the insect groups.

Bouchard (2004)

X*

Order and families

Useful for identification in Upper Midwest of N.A.; includes feeding behaviors, tolerance values, and primary habitat preference.

Fauceglia (2005)

X

None

Biological information on Eastern and Midwestern US mayfly hatches with excellent photographs.

Triplehorn and

Families of terrestrial

Johnson (2005)

and aquatics

Mostly adult coverage; newly described orders and families incorporated. Widely used book for terrestrial insect identification.

Izaak Walton League

A handy resource for anglers, students, and biologists spending time near rivers and streams. Gives tips for distinguishing similar species on behavior and their role in

X

Orders and some families

Ames(2008)

X

None

A fly angler's guide to families, genera and species of Eastern US caddisflies with color photographs.

Thorp and Rogers (2011)

*

Orders of aquatic insects

This handy field book complete with color photographs is designed for students and laypersons interested in general identification and ecology of inland water

of America (2006)

stream ecosystems.

invertebrates of the USA and Canada.

*Only adult keys to Hemiptera and Coleoptera.

"•"Contains notes on biology or ecology. *Covers adults of some groups. (continued)

Chapter 1

Table 1

Introduction

Continued

Taxonomic GENERAL COVERAGE

Treatments Source

Biology

Immatures Adults

Tzilkowski and

General Comments

Identification, habitat and life history information for fly fishers and fly tiers on Eastern North American nymphs of several orders of aquatic insects, along with

None

Stauffer (2011)

imitations.

Thorp and Rogers (2016)

Orders and families of

Deals with inland water invertebrates

aquatic insects and

of the Nearctic, primarily include taxonomic keys supplemented by an introduction to the group and sections on limitations to taxonomy of the

differential taxonomic treatment of other invertebrates

group, information on critical structural terms used in the keys, and recommendations for preparation and preservation of specimens. Morse etal.(2017)

Families, genera, species

Keys to larvae of the Southeastern USA mayfly, stonefly, and caddisfly species, excellent photographs.

Weamer (2017)

None

Guide to aquatic entomology written for new anglers who want a basic understanding of aquatic insects or more seasoned fly fishers who want to take their skills to the next level.

Ecological Treatments

GENERAL COVERAGE

Source

Hynes(1972) Ward and Stanford (1979) Lock and Williams(1981)

The "classic" on the ecology of running waters. A "must have" for every student and researcher. Wide coverage of topics on biology of rivers and streams with emphasis on aquatic invertebrates. Comprehensive treatment on the ecology of stream regulation, with an emphasis on downstream effects on biotic (especially aquatic insects) and abiotic components. This text was written in honor of the retirement of H.B.N Hynes by former graduate students. Contains chapters on migrations, distributions, hydrodynamics, and ecology of aquatic Insects.

Barnes and Minshall (1982) The first attempt for the application and testing of general ecological theory to stream ecology. Several

chapters discuss the ways In which aquatic insects can be used to empirically test ecological theory. Resh and Rosenberg (1984) A very good overview of aquatic insect ecology, highlighting research needs and suggested avenues of Williams(1987) Ward (1992)

Williams and Feltmate

(1992, 2017)

Investigation. Good reviews on several current topics in aquatic ecology. Widely used reference source. An introduction to the ecology of temporary aquatic habitats, with a discussion of the abiotic features of these environments and the biology of invertebrates colonizing these habitats. A treatment of the evolutionary considerations, habitat occurrences of aquatic insect communities, and the relationship of aquatic insects to environmental variables. Good treatment of physical aspects of aquatic insect biology and their habitat.

An introductory text to the study of aquatic insects, with background information on the aquatic insect orders and good coverage of life histories, adaptations, population biology, trophic relationships and experimental design and sampling methods.

Rosenberg and Resh (1993) A very thorough reference source dealing with many different approaches for using benthic macroinvertebrates in biological monitoring programs. *Only adult keys to Hemiptera and Coleoptera. ^Contains notes on biology or ecology. ^Covers adults of some groups. (continued)

Chapter 1

Table 1

Introduction

Continued

Ecological Treatments

Wotton (1994)

GENERAL COVERAGE

This book takes a functional approach In reviewing the role of partlculate and dissolved matter in a wide range of marine and freshwater ecosystems. Specific chapters deal the food of aquatic Insects and the manner In which they capture particles In their environment. A good reference source for students and researchers alike.

A beginning text In stream ecology, with good overall coverage of biotic and abiotic factors influencing aquatic insect distributions and abundance. Good coverage on subjects such as drift, predation, competition, feeding ecology of fish, and the modification of running waters by humankind. Giller and Malmqvist(1998) Introductory text provides an overview of physical processes and chemical dynamics In structuring lotic communities. Discusses the physiological and physical adaptations of organisms and lotic food webs, along with a discussion on water pollution and conservation. Provides examples of global lotic habitats.

Allan (1995)

Batzer etal.(1999)

This text synthesizes research regarding the ecology and management of invertebrates found In N.A. freshwater coastal and Inland wetlands.

Karrand Chu (1999)

Gushing and Allan (2001)

Mackle (2001)

Dodds (2002)

Resh and Garde

(2003, 2009)

Benke and Gushing (2005)

Discusses biological monitoring and assessment of freshwater ecosystems in the U.S. and describes the use of biological communities for diagnosing environmental degradation. Examines the use of multimetric Indices and how they can be Incorporated into environmental policy and management. A practical bock for students, researchers, and managers. Gomprehenslve book covering the fundamentals of stream ecology. Provides a discussion on a wide range of river types and the diverse biota that comprise stream food webs. Geared toward conservation groups, adopt-a-stream programs, and Individual citizens. Summarizes fundamental limnologlcal and water management concepts. Applications of concepts are provided In each chapter. Includes descriptions of aquatic organisms, especially macrolnvertebrates, and their use to assess water quality. A treatment covering basic and applied concepts of freshwater ecosystems. Includes chapters on physical processes, chemical cycles, and a diversity of organisms, such as microbes, plants. Invertebrates, and fish. An Ideal text for students and managers.

This encyclopedia of Insects contains subject area coverage of many aquatic topics (e.g., aquatic habitats, growth, marine Insects, metamorphosis, respiratory system) and several aquatic Insect orders. It is geared for the beginning and advanced student, as well as professionals. Excellent photographs and illustrations. This comprehensive treatise on North American rivers was written for scientists, students, river conservationists, and lay persons. It contains a detailed examination of N.A. rivers that provides a regional framework for comparing the physical, chemical, and biological properties of rivers. For each river, it

Brdnmark and

often Includes a section on aquatic Invertebrates, their diversity, abundance, and ecology. A thorough overview of lake and pond ecology. Discusses the structure and function of lentic

Hansson (2005) Williams (2005)

ecosystems, emphasizing the importance of abiotic factors and blotic Interactions. Examines the ecology of temporary waters In natural and human environments. Synthesizes the diverse

global literature and applied aspects of these systems, discussing the ecological Importance and need for conservation. A relevant text for graduate students and researchers. Hauer and Lamberti

(2006, 2017), Lamberti and Hauer (2017)

Marshall (2006)

Lancaster and

Downes(2013)

Thorp and Rogers (2015)

A detailed description of field and laboratory methods commonly employed in the study of physical, chemical, and biological components of stream ecosystem structure. Reflects latest advances in the technology associated with ecological assessment. Includes data sheets and links to downloadable spreadsheets for conducting stream ecology. Also Includes keys to common stream macrolnvertebrates and functional group keys. A great Introduction to insect diversity and natural history with basic Information (characteristics, habitat, behavior) about all major Insect families with comprehensive picture keys. Goverage in aquatics Include chapters on mayflies, dragonflles, damselflies, stoneflles and caddisflies. Photographs are excellent. A biological approach to aquatic entomology structured around four sections; distribution patterns and environmental gradients, dispersal and movement, population dynamics and persistence, and trophic relationships. This first of 10-12 projected volumes In this series provides ecological, morphological, and general biological coverage of Inland water Invertebrates of the world, and Is meant as a companion volume for all subsequent volumes focused on identification of invertebrates.

^

m

4^..V

'Vfy^ f"

S&s'> -

'Sfes*/^ ■l'.^

GENERAL MORPHOLOGY

OF AQUATIC INSECTS Martin B. Berg Loyola University Chicago, Illinois

Richard W. Merritt

Miehi^an State University, East Lansing

Kenneth W. Cummins

Humboldt State University, Areata

OVERVIEW

A stonefly (order Plecoptera, family Pteronarcyidae) serves to illustrate the general external morpho logical features of aquatic insects used in taxonomic determinations. This prototypical insect exhibits basic morphological features in a relatively unmodi fied or nonspecialized form. However, modifications of the general morphological plan are found in each insect order having aquatic representatives. These modifications and associated terminology are pre

sented with the introductory material for each group and should be carefully studied before attempting to use the keys in the following chapters. The insect body represents the fusion and modifi cation of the basic segmentation plan characteristic of the Annelida-Arthropoda evolutionary line (e.g., Snodgrass 1935; Manton and Anderson 1979). Each segment of the body can be compared to a box, with the dorsal (top) portion, the tergum or notum, joined to the ventral (bottom) portion, the sternum, and to the sides or lateral portions, the pleura, by mem branes. The legs and wings are hinged (articulated) on the pleura of the mid-body region, the thorax. The body regions, head, thorax, and abdomen, and asso ciated appendages of a stonefly nymph are shown in Figs. 2.1 and 2.2. The life cycle of stoneflies is repre sentative of those orders characterized by simple {incomplete or hemimetabolous by some authors) metamorphosis, consisting of egg, nymph (imma ture), and adult stages; more advanced orders exhibit complete (holometabolous) metamorphosis, consist ing of egg, larva (immature), pupa, and adult stages. Distinction between the terms "nymph" for immature hemimetabolous insects and "larva" for immature

holometabolous insects is supported by endocrine

and developmental data (Truman and Riddiford 2002; Chapter 11 in this volume). HEAD

The generalized insect head represents the evolu tionary fusion of six or seven anterior segments in the ancestral Annelida-Arthropoda line (e.g., Snodgrass 1935; Rempel 1975). Two or three preoral (procephalic) segments, or somites, were fused and now bear import ant sensory structures used by present-day insects to monitor their environment—the compound eyes,

light-sensitive ocelli (simple eyes), and the antennae (Figs. 2.1 and 2.2). The labrum, which forms the upper lip, is joined at its base to the clypeus, which in turn is fused to the frons, or face. The margins of the clypeus and frons are bounded by the anterior portion of the Y-shaped epicranial suture (in Fig. 2.1, the line ofjoining between the clypeus and frons, termed a suture [sulcus], is not externally visible so the structure is referred to as the frontoclypeus [Nelson and Hanson 1971]). Three postoral (gnathocephalic) segments are fused in modern insects to form the posterior portion of the head and bear the remaining structures of the feeding apparatus (Snodgrass 1935). As described above, the labrum forms the upper lip and the paired mandibles and maxillae form the mouth region laterally (Figs. 2.2 and 2.3). The bottom of the mouth is set by the labium or lower lip (Figs. 2.2 and 2.3). The maxillae and the labium bear palps (palpi), which are sensory in function (Figs. 2.2 and 2.3). The mandibles are used for chewing or crushing food or may be modified for pierc ing (piercing herbivores or predators) or scraping (scraping herbivores that graze on attached algae). The maxillae and labium are variously used for tearing and manipulating food, or they may be highly modified as

10

Chapter 2 General Morphology of Aquatic Insects

frontociypeus pedicel

lobrum

scape

antenna

oceiii

arm of frontal suture

compound eye

cervix

epicranlal suture occiput

pronotum (pronotal stiield; notum) of prothorax

foretibia

foreleg branched gili

mesonotum (mesonotal shield; notuni of mesothorax

midleg forewing pad

metanotum (metonotol shield; notum) of

metathorax

femur of hind leg hind leg

tarsus of hind leg

hind tarsal claw

^ terga of abdomen hind wing pad tergum

paraproct

(subanai lobe) epiproct

(supraanal process) cercus

& Gun>

Figure 2.1 Dorsal view of Pteronarcys sp. nymph (Plecoptera: Pteronarcyidae).

Chapter 2 General Morphology of Aquatic Insects

antenna

labrum

mandible

pedicel maxillary palp

scape

compound eye coxa of foreleg maxilla

trochonter

'a'''^^^]^^__^pronotum (pronotal shield; tergum) of prothorox

femur tarsus

tibia

prosternum (sternum) of prothorox branched gill

mesosternum (sternum) of mesothorox

forewing pad

metasternum (sternum) of metathorax

hind wing pad

8th obdominal sternum

epiproct (supraonol

paraproct (subanal lobe)

process)

cercus

Figure 2.2 Ventral view of Pteronarcys sp. nymph (Plecoptera: Pteronarcyidae). Gills on left side of thorax and first two abdominal segments removed to show underlying structures.

11

12

Chapter 2 General Morphology of Aquatic Insects

1 labrum (upper lip) right mandible hypopharynx

^terminal (distal) incisor lobe of teeth

basal

(proximal) molar lobe of teeth

points of articulation

maxillary palp (palpus) showing palpal segments

lacinia

palpifer

cardo

(base of maxilla) ventral

right maxilla

glossa

labial palp (palpus)

paraglossa

showing

palpal

femur

prementum

segments

(lablostipites)

trochanter

postmentum submentum

B

labiuffl (lower lip)

Figure 2.3

2

tarsus (tarsal segments) .tarsal claws

Figure 2.4

Figure 2.3 Ventral view of head and mouthparts of Pteronarcys sp. (Plecoptera; Pteronarcyidae): A. ventral view of head; B. lablum; C. right maxliia; D. right mandible; E. hypopharynx; F. labrum.

Figure 2.4 Foreleg of Pteronarcys sp.(Plecoptera; Pteronarcyidae) showing segments.

Chapter 2 General Morphology of Aquatic Insects

in the Hemiptera, adult Lepidoptera, Hymenoptera, and Diptera. The hypopharynx or insect "tongue," located just anterior to the labium,is a small inconspic uous lobe in some larval forms,but is subject to extreme modification in some orders (e.g., Diptera). The sides of the head are referred to as genae (singular, gena; Fig. 2.2)and the top ofthe head as the vertex. Immediately behind the vertex is a large area called the occiput(Fig. 2.1). The head is joined to the thorax by a membranous neck region or cervix (Fig. 2.1). If the head is joined to the thorax so that the mouthparts are directed downward (ventrally), the condition is termed hypognathous(e.g., many caddisfly larvae). Mouthparts directed forward (anteri orly) are prognathous (e.g., beetle larvae) and those directed backward (posteriorly) are opisthognathous (e.g., some true bugs). In aquatic insects that are dorsoventrally flat tened, such as some stoneflies and mayflies, the sen sory structures (eyes, ocelli, and antennae) are dorsal and the food-gathering apparatus is ventral. These modifications allow certain groups to move through interstices of coarse sediments and cling to exposed surfaces in rapidly flowing streams.

THORAX

The midregion of the body, or thorax, bears the jointed legs (Fig. 2.4) and the wings, and is divided into three segments (Figs. 2.1, 2.2, 2.5, and 2.6). The prothorax bears the forelegs, the mesothorax the midlegs and forewings, and the metathorax the hind legs and hind wings (if wings are present). The jointed legs are five-segmented: the coxa, trochanter, femur, tibia, and the three- to five-seg mented tarsus, which terminates in one or two tarsal

13

types of wing venation are shown in Figs. 2.5 and 2.6. The prototypical stonefly wings have many branches of the major veins with many crossveins between them. The highly evolved wing of a dipteran Tipulidae (Tipula sp.) is characterized by the fusion of veins and the loss of branches and crossveins.

The general venation pattern (Figs. 2.5 and 2.6) consists of: a costal vein (C), the anterior marginal vein; a subcostal vein (Sc)just behind the costal vein and often with two branches near the wing tip; a radial vein (R), often the heaviest vein of the wing, which forks near the middle of the wing, with the main part forming the radial sector vein (Rs) that typically divides into two branches, each of which may divide into two or more branches near the wing margin; a medial vein (M)(the fourth major vein), which has a maximum offour major branches (typi cally two or three); a cubital vein(Cu), which has two major branches, the anterior of which usually forks into two branches; and an anal (vannal) vein (A), which has a maximum of three major branches with considerable secondary branching, particularly in more ancestral forms. Although crossveins are highly variable, certain ones are usually present. There are generally at least one humeral crossvein(h) between the base of the wing and the apex (tip) of the subcosta; a radial crossvein (r) between the radius and the first branch of the radial sector; a radial-me

dial crossvein (r-m) between the lower first fork of the radial sector and the upper first fork of the medial vein; and a medial-cubital crossvein (m-cu) between the lower first fork of the medial and the

upper first fork of the cubital (see Snodgrass 1935; Daly eta/. 1978; Bonoretal. 1981; and discussion of taxonomically significant wing veins given in the order and family chapters below).

claws (Fig. 2.4). In aquatic insects, modifications of the hind legs for swimming (e.g., a fringe of tibial hairs) are common in certain adult Coleoptera, some larval and adult Hemiptera,and a few larval Trichoptera. The forelegs are modified for burrowing in Ephemeridae(Ephemeroptera), Gomphidae(Odonata), and some semiaquatic Orthoptera. Most adult forms of aquatic insects bear two pairs of wings(mesothoracic and metathoracic);some mayflies and all Diptera have only one pair. The sec ond pair of wings (metathoracic) in Diptera is modi fied into balancing organs (halteres. Fig. 2.6) that function somewhat as gyroscopes. Collembola are wingless (apterous), as are females of certain species of Trichoptera and Diptera. The structures that extend into the wings are termed veins. The form and location of these veins are

used extensively in insect taxonomy. Two extreme

ABDOMEN

The prototypical insect abdomen is composed of eleven segments, although in most adults fusion ofthe last two makes them difficult to distinguish. In some immature forms (notably Ephemeroptera and Mega-

loptera), gills arise from the pleural regions—being extensions of the tracheal system borne in variously shaped plates or filaments (finger-like gills). In the stonefly shown in Fig. 2.2, the branched filamentous gills are attached to the sterna of the thorax and the first two abdominal segments. The end of the abdomen of hemimetabolous

insects (i.e., Hemiptera, Orthoptera, Ephemerop tera, Odonata, and Plecoptera) bears the reproduc tive structures (Figs. 2.1, 2.2, and 2.7-2.9). The terminal segment bears the anus at its apex and the

2

1

anal veins

3

anal cell

\

vein

cubital

pronotum

told lines

anal veins

crossvems

medial

subcostal vein

subcostal vein

costal vein

anal (vannal) region

vein

costal

tiumeral

radial vein

cubital vein

branches of

forewing

branches of medial vein

branches of radial sector

posterior cubital vein

hind wing

anterior cubital vein

- medial vein

- radial sector vein

radial sector

radial vein

) ) ) )))) ) ) ) )) 1 ) ) ) ) ) ) i ))) ) ) ) )

Figure 2.5 Adult Pteronarcys sp.(Plecoptera; Pteronarcyidae)showing head, thorax, basal portion of abdomen, and fore and hind wings.

tergum

1st abdominal

metanotum-

mesonotum

pronotum

compound eye

ocelli

scape

pedicel

.antenna (terminal portion cut away)

Chapter 2 General Morphology of Aquatic Insects

15

- antenna

.compound eye

subcostal radial costal

radial

stigma

sector

humeral crossvein

axillary region

halter meson otum

metanotum

cubital-anal crossvein

1st abdominal tergum

anal veins

medial vein cubital vein

Figure 2.6

median

supraanal lobe

hemitergal anterior

epiproct (supraanal process) cercus (terminal portion cut away)

cercus

posterior lobe /

hemitergal (terminal portion

hemitergal

lobe

cut away)

lobe

tergum

9th tergum

remnant ot 11th tergum

epiproct

posterior hemitergal lobe

10th tergum

(supraanal process)

8th abdominal

median hemitergal lobe

r^ 9th tergum

tergum

supraanal lobe paraproct

anterior hemitergal lobe

(subanalprocess) pleuron

toth sternum

8th tergum 9th sternum Bth abdominal sternum

Figure 2.8

Figure 2.7

epiproct cercus

(terminal portion cut away) paraproct (subanai lobe) 11th abdominal sternum toth sternum

9th abdominal tergum 9th sternum

vaginal protection

genital opening (gonopore)

8th sternum

8th abdominal tergum imcKsrr

Figure 2.9

Figure 2.6 Dorsal view of adult Tipula sp.(Diptera: Tipulidae) showing head, thorax, basal portion of abdomen, forewing and halter. Figure 2.7 Dorsal view of terminal male abdominal segments of Pteronarcys sp.(Plecoptera: Pteronarcyidae); terminology after Snodgrass(1935) and Nelson and Hanson (1971).

Figure 2.8 Lateral view of terminal male abdominal segments of Pteronarcys sp.(Plecoptera: Pteronarcyidae). Figure 2.9 Ventral view of terminal female abdominal segments of Pteronarcys sp.(Plecoptera: Pteronarcyidae).

16

Chapter 2 General Morphology of Aquatic Insects

cerci laterally. The dorsal surface is covered by a triangular or shield-shaped tergal plate,the epiproct, and the ventral surface bears two lobes, the paraprocts. In males, the ninth sternum often bears two

lateral styli or claspers (harpagones). These acces sory structures bound the phallobase and aedeagus that comprise the main reproductive organ, the penis or phallus. The terminal segments of adult females, in addi tion to the dorsal epiproct and lateral paraprocts below the cerci, generally consist of three pairs of lobes or valvae (valves), which form the visible por tion of the ovipositor and arise from the eighth and ninth sterna. The bases of the valvae are usually cov ered by the projecting eighth sternum (Fig. 2.9).

Specific morphological modifications in each of the orders(or families receiving special treatment)are detailed in the introductory material covering the respective groups.The modifications usually represent fusion or specialization of the basic structures discussed above. However, some of the terms used in naming the various segments of the genitalia have restricted meanings, and homology with primitive forms is not always possible (Tuxen 1970; Scudder 1971a). For a more complete treatment of insect morphology, the student should consult Snodgrass (1935), DuForte (1959), Matsuda (1965, 1970, 1976), Chapman (2013), Resh and Garde(2009), and Gullan and Cranston(2005). Consult Torre-Bueno(1937)for further explanation of terms.

iMkmm

^"^'4

^

mr '%

SAMPLING AQUATIC INSECTS Collection Devices, Statistical Considerations, AND Rearing Procedures John K. Jackson Stroud Water Research Center, Avondale,

Pennsylvania Vincent H. Resh

Richard W. Merritt

Michigan State University, East Lansing Kenneth W, Cummins

Humboldt State University, Arcata

University of California, Berkeley Darold P. Batzer

University of Georgia, Athens

The study of aquatic insects depends on our abil ity to collect these organisms, which leads to the need to make decisions about appropriate sampling

devices and/or laboratory processing procedures to separate the animals of interest from the abiotic material they reside in. However, the first step in planning any scientific study, before any sampling decision can be made, is to answer the WHY

question—why are we collecting samples? At the core of this question is the need to have a clearly defined scientific hypothesis or goal that underlies the objec tives ofthe study. A clearly defined scientific question then leads to other inquiries about what types of data are needed (e.g., qualitative vs. quantitative, abun

dance vs. biomass, population vs. community), where to sample (e.g.,the habitat such as a wadeable stream, deep river, wooded wetland, or stormwater pond, and more specific locations such as upstream vs. downstream or littoral vs. profundal), and when to sample (e.g., once per year [e.g., spring vs. summer], quarterly, monthly, according to flow regimes). Answers to those questions are important when developing a plan that includes decisions about sam pling devices or laboratory processing procedures to be used. It is also through this process that other data such as water chemistry, algal biomass, benthic CPOM (coarse particulate organic matter), etc., are identified as essential and can be added to the sam

pling plan.

COLLECTING AND SAMPLING DEVICES

A variety of approaches and devices have been used to collect aquatic insects or provide quantitative information on their richness, abundance, or biomass (Table 3A). The table provided is not a complete list ing of the references on sampling methods, and stu dents and researchers should continually check the current literature for methods suited to their specific objectives. The bibliographies of the Society for Freshwater Science (formerly the North American Benthological Society) contain scores of other papers describing devices for collecting aquatic insects, as does the extensive bibliography of Elliott eta/.(1993). As noted by Cummins(1962)long ago,the number of different samplers used for benthic macroinvertebrates is nearly equal to the number of benthic investigations! The classification system used in Table 3A is based primarily on the habitat and community being sampled. Substrate composition, although not out lined in detail, also is an important consideration when sampling the benthos (e.g., Minshall and Minshall 1977; Rabeni and Minshall 1977; Lamberti and Resh

1978; Reice 1980; Minshall 1984), and the equipment and techniques listed below may require modification depending on the substrate type. For example, an Ekman grab is listed as an appropriate device for littoral benthos(IV. A.1. a. in Table 3A); however,the presence of sticks or even small stones could prevent

17

Rivers, and Springs

1. Shallow Streams,

LOTIC HABITATS

Habitat

Major Sampling Figure Reference(s)

Electroshocking

Riffle sampler

(subterranean)

Standpipe corer

3.42

3.41

3.29

3174, 235, 6214

3.9

Ellis-Rutter Stream

sampler

725

3.40

Suction samplers (air-lift and water pump)

1733, 6583, 6595, 6597

517, 1086, 2125, 2834, 4144, 4781

1077

623, 4384, 6698, 2262,

3.5

126, 5690, 6409, 6556

5378, 2753, 6768, 6214, 3928

Wilding or stovepipe sampler: box-type sampler

outcrop sampler

Individual stone, bedrock, or rock-

3655

3.8

T-sampler

2571, 2902, 6363, 879,

4808

1673

3.6

Mess or modified Mess

2125

6897, 6906

Canister sampler

Pump sampling

296, 4197, 6099

5752, 665, 4837, 664, 3579

3.41

3.33

1246, 1247

Freeze-core samplers (including electrofreezing)

Graded sieves

Photographic methods

4025, 4655, 5394, 624, 1450, 6896

Leaf packs

6596

2004, 2829, 4773, 3631, 5775, 3486, 6892

1460, 6768, 3928

2764, 3613, 3765, 5084, 5115, 5823, 954, 6416, 3371, 896, 2901, 6889, 6903

6099

6099, 3631, 3721

References

3613, 6099 3.25

3.23

3.3

3.2

Figure

Hand collection

Recolonization

Kick sampling

Individual stone sampler

samplers

Artificial substrate

Hand screen collector

1031, 2764, 3613, 4090, Aquatic net 4289, 5823, 954, 5775,

sampler

3.7

Surber sampler

b, Hyporheic area Implants

a. Sediments

1. Benthos

A. Riffles (erosional zones)

Sampler

or Semi-

quantitative Sampler

Quantitative

Ecological Community

Qualitative

and

Subhabitat

Table 3A Collecting and sampling methods for aquatic macroinvertebrates based on the habitat and community being sampled.

) ) ))) ) ) ) ) ) ) )j ) ) ) ) I ) ) ) ) ) :i

00

^

a. Sediments

1. Benthos

B. Pools depositional zones)

2. Emerging Adults

(surface)

Neuston

d. Drift and

c. Plants

Figure Reference(s)

4193

Mundle pyramid trap

3.12 3.40

3.14

Suction samplers (air-lift and water-pump) Mark-recapture

3.5

Single corerwith pole

3.16

Ekman grab with pole Wilding or stovepipe sampler

1089, 2032, 6227 623, 725

Graded sieves

Aquatic net

3.33

6099

299

3501, 3615,4130, 1352

Floating emergence Traps

2833

5582

Light traps 6343

Enclosed channels

3.9

984, 5837

Window traps

2342, 2844, 3773

3.2

6099

6099

2246, 3335

3.3

3.2

Pan traps

Hand screen collector

Aquatic net

1395

6099

Aquatic net Snag sampler 3.2

6099

(continued)

References

Needham apron net

Figure

Stationary screen trap

3.26

1074

Surface film sampler

1074

1624, 2833

1288, 4195

1627, 2833

4808

2514

3.11

Cushing-Mundle drift sampler Hardy plankton indicator type sampler Surface film sampler Colonization cages

Plankton net

208, 1627, 6355, 6596, 6638, 91, 1673, 3822,

3.10

Drift net

126, 6393 2587

3.5

Stovepipe sampler

2833,6409

Lambourn sampler

3.7

Surber sampler

Bag sampler

Sampler

or Semi-

quantitative Sampler

Quantitative

Ecological Community

Qualitative

and

Subhabltat

Continued

SAMPLING

Habitat

Major Sampling

Table 3A

) ) J ) ) ) 3 ) )))) ) )) ) ) ) 3 ) ) ) ) ) ) ) )

2, Emerging Adults

a. Sediments

1. Benthos

B. Pools

2. Emerging Adults

Neuston

b. Drift and

a. Sediments

1. Benthos

A. Riffles

Figure Reference(s)

1627,4196

3.17

3.19 3.16

3.12

Ponar grab Petersen-type grabs Ekman grab Core sampler

1648, 1846, 2032, 2118

1617, 1645, 1648, 2833, 3517

1645, 3517, 4674

1645, 1648, 3517, 4801

Floating drift trap (with floats)

3399, 4130, 1352 4196

Insect emergence traps

Hardy plankton indicator-type sampler

1624, 2833

82, 1645, 1648

floats)

3.40

Suction samplers (air-lift and water pump)

SCUBA diving

Basket-type artificial substrate samplers

Drag-type samplers

samplers

3.24

3.23

3.24

Figure

Basket or cylindricaltype artificial substrate samplers

Section I.A.2

See Section I.A.I.d.

Needham apron net

2034, 2035, 2587, 1725 Single or multiple-plate

1648, 2036, 2285, 6175, 1516, 2380, 4384, 1517

Grab samplers

3.40

2833

Floating drift trap (with

SCUBA diving

Suction samplers (air-lift and water pump)

See Section I.A.2

See Section I.A.I ,d

c. Drift and Neuston

Bag sampler

b. Plants

Sampler

or Semi-

quantitative Sampler

Quantitative

Ecological Community

Qualitative

and

Subhabitat

Continued

) ) ) )) ) ) ) ) ) ) ) V ) ) ) ) ) ) ) )

II. Large Rivers

Habitat

Major Sampling

Table 3A

) ) ))

1648, 2833

1847, 2034, 3768, 5123

1646, 1648, 6104

1423, 1847, 2424, 2574, 5123

1516, 4532, 3651

147, 174, 371, 1219, 1847, 2034, 2337, 2587, 2597, 3768, 3771, 4489, 5123, 6395

6099

References

Ki

SAMPLING

Figure 82

Reference(s)

3.20

b. Rooted Plants, Macan sampler Periphyton

2092, 153, 685

3.22

Gerking sampler Modified Gerking sampler

2059, 3054, 4086

2587

3.21

Lambourn sampler

1493

Minto sampler

1493, 1497, 2736, 3043, 3611

725, 2901

3.40

Suction sampler

3096, 6350

Activity traps

traps

Drop traps and pull-up

Modified KUG sampler

Artificial substrates

Aquatic or sweep net

3.44

3.2

(continued)

4218, 5036, 685, 6066, 5426, 2370, 4231, 6890, 6891, 6899

291, 1968

328

3621, 5494, 5495, 685,

3054, 6200, 993, 6066, 328,2522, 6890, 6891, 6895, 6901, 6907

3395, 5935, 6099 3.33

2571, 2587, 6556, 2522, Graded sieves 6902

2522

6099, 993, 6066, 328,

References

3.2

3469, 5840, 153

3.13

3.5,3.6

Wilding or Hess-type sampler

Kellen grab

3.12

Core sampler

See Section II.A.2.

Figure

368, 1089, 2032,6227, Aquatic net 1717, 685,6066, 328,2522

Section III.A.1 .b

Section III.A.1 .b.

See Section II.A.1 .b.

See Lentic Habitats,

See Lentic Habitats,

3.18

Water column sampler

a. Sediments

1. Benthos

A. VEGETATED

2. Emerging Adults

Neuston

c. Drift and

b. Plants

HABITATS

Also, see LENTIC

Allan hand-operated grab

Sampler

or Semi-

quantitative Sampler

Quantitative

Ecological Community

Qualitative

and

Subhabitat

Continued

III. Wetlands(ponds, swamps, marshes, bogs, rice fields, etc.)

LENTIC HABITATS

Habitat

Major Sampling

Table 3A

Habitat

Major Sampling

Table 3A

Figure Reference(s)

a. Sediments

1. Benthos

B. Nonvegetated

2. Emerging Adults

c. Neuston

Suction sampler (air-lift and water pump)

Petite Ponar grab

Kellen grab

pole

Ekman with or without

that float)

3.40

3.13

3.16

2531

Funnel trap

623, 590

3096

Activity traps

Aquatic net

1497, 1617, 2736, 3043 Dredges

3,44

3.2

3.28

4218, 5036, 685, 6066, 5426, 2370,4231

6099, 328, 2522

3043

30, 194, 571, 902, 903, 1709, 2789, 2806, 6350, 6871

5818, 6099, 6200 3.4

Hand dipper

328

328, 6894

3.11

Throw-net

Plankton tow net

30, 916, 903, 2789, 2806, 6350, 6871

2522

5818, 6099, 6200, 328,

2610, 2623, 5036, 685, 6066, 5426, 2370, 4231

328

References

2226

3.28

3.2

3.44

Figure

Telescope method

Subaquatic light traps

Aquatic net

Activity traps

Throw-net

1128, 1294, 1352, 1673, Subaquatic light traps 1716, 2298, 3017, 3142, 3375, 3501, 3532,3924,4130, 4131, 4193, 4198, 5229, 5118, 6769

2044

D-vac vacuum sampler

Emergence traps (primarily surface traps

410, 3469, 153

Water column sampler

3.27

1490

Douglas method Hess sampler

1717, 5495

Plexiglas or metal tubes

3.12

5495

Removal of natural substrates

1493, 153

Quadrat clipping

Sampler

or Semi-

quantitative Sampler

Quantitative

Ecological Community

Qualitative

and

Subhabitat

Continued

) ) ))) ) ) ) ) ) ) ) ) ) )! ) ) > ) ) ))) ))

K> Ul

OJ

2. Emerging Adults

b. Vegetated (plant zone)

a. Nonvegetated (wave swept)

1. Benthos

A. Littoral

2. Emerging Adults

b. Neuston

Figure

3.12

3.16

Core sampler with or without pole Ekman or similar type grab with pole

substrate by SCUBA

Removal of natural

attachment rod

Macan sampler with

Kornijow hinged-box sampler

Gillespie and Brown sampler

Wilding or stovepipe sampler

III.B.1 .a.

Also, see Section

SCUBA diving with sampling gear

Reference(s)

6440

3043, 4434, 6440

6898

2117

6393, 6556

1737, 2033, 1497

178, 1645, 1846, 2735, 3043, 4801

1387, 1645,2735, 4434, 6697, 1497

368, 1089, 1846, 2032, 6227, 1497

1825, 2285, 1497

3043, 6409, 6556

5840

368, 1089, 2032, 3043, 328, 2522

See Section III.A.2.

3.20

3.5

3.17

3.40

Suction samplers (air-lift and water pump)

Ponar grab

3.5

3.12

Wilding or stovepipe sampler

See Section III.A.2

See Section III.A.I.c

Water column sampler

Core sampler

Sampler

Modified KUG sampler

Photographic methods

Dredges and grabs

Graded sieves

Basket-type artificial substrate samplers*

See Section III.A.2

See Section III.A.1 .c

or Semi-

quantitative Sampler

Quantitative

Ecological Community

Qualitative

and

Subhabitat

Continued

SAMPLING

IV. Lakes

Habitat

Major Sampling

Table 3A

3.33

3.24

Figure

727

6393

1497

6099

(continued)

178, 209, 371, 1847, 4111, 5123, 6209, 6395

References

Margins, Dry

Figure Reference(s)

A. Flying Aquatic or Semiaquatic Insects Window traps

malaise traps

Emergence traps or

Emergence traps (mainly funnel and submerged traps)

SCUBA diving with sampling gear

Petersen-type grab

Ponar grab

3.29

3.27

3026

3685, 5402, 5516, 6025, 6027

755, 1759, 5121, 2298, 3017,4130,4131,4193, 4198, 5118,6471, 1352

1737, 3043, 6209

3043, 4674, 5540, 6409

3.19

1617, 1846, 3043

1846, 3043, 4801, 1497

3.16

Ekman or modified

1846, 2344, 3043, 4050

Sieve sampler splitter

Graded sieves

1237, 3769

2422,6905 6099

2731,4182, 5402, 5582, 5815

3043

Sticky traps

3.33

3.30, 3.31

3.13

1497

References

Aspirator

Light traps

Activity traps

Photographic methods

683, 1846, 2699, 3043, Dredges 3050, 3064, 4050, 6409, 6471, 1497, 78, 2841

3.17

3.14

Multiple core sampler

Ekman grab

3.13

Single core sampler

Figure

) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ))) )

Wetlands

2. Emerging Adults

a. Sediments

1, Benthos

B. Profundal

Sampler

or Semi-

quantitative Sampler

Quantitative

Ecological Community

Qualitative

Subhabitat and

Continued

V. Stream and Lake

HABITATS

TERRESTRIAL

Habitat

Major Sampling

Table 3A

'Ji

Substrates

Figure

Reference(s)

1209, 1237, 1238, 3433, 3446, 6482, 6470

Eiutriation and flotation

1743

6695

5582, 6099

4774

3.45

3.32

Field washing

procedure

Behavioral extraction

Emergence chamber

funnels

B. Aquatic Insects in Dry Beriese or Tullgren

Sampier

or Semi-

quantitative Sampler

Quantitative

Ecological Community

Qualitative

and

Subhabitat

Continued

SAMPLING

Habitat

Major Sampling

Table 3A

Figure

References

))))) ) )) ) ))))))))) 3 )))))) )

26

Chapter 3 Sampling Aquatic Insects

the jaws from closing, resulting in the collected mate rial being washed through the slightly opened jaws as the sample is pulled to the surface. Resh et al. (1990) have prepared a videotape of the operation of twenty of the devices listed in Table 3A that is available online

(https://nature.berkeley.edu/reshlab/samplingvideos. htm). Comprehensive reviews and discussions of differ ent collecting and sampling methods for macroinvertebrates have been presented by Welch (1948), Macan (1958), Albrecht (1959), Cummins (1962), Hrbacek (1962), Hynes (1970a), Edmondson and Winberg (1971), Weber (1973), Hellawell (1978, 1986), Southwood (1978), Downing and Rigler(1984), Batzer etal.

(2001), Carter and Resh (2001, 2013), and Blaustein and Spencer (2005). Additional information on sam pling devices is available in Peckarsky (1984), Winterbourn (1985), ASTM (1987), Britton and Greeson (1987), APHA (1989), Klemm et al. (1990), Williams and Feltmate (1992), Cuffney et al. (1993a), Murkin et al.(1994), and Keiper et al.(2002).

CHOOSING AN APPROPRIATE SAMPLING DEVICE

Ultimately, the decision to use a specific sampler or collecting device should depend on the objectives of the study (e.g., Andre et al. 1981) and a thorough

Table 3B Selected examples of factors that affect benthic sampling devices and may result In sampling bias (modified from Resh 1979a).

Factor

Examples of Samplers Affected

Problems Created

Remedy

A. Factors related to characteristics of the samplers Backwash created in

Loss of benthos aound sides of

Increase the net's surface area and/or

sampler

decrease size of net opening; use enclosed double netted sampler (4197); alternatively use a hand-operated Ekman grab or cylinder box sampler (2833)

Turbulence scours substrate surface

Use permeable sides

Corer and Grab

Loss of small organisms and surface

samplers

dwellers

Modify Ekman grab by removing screens and Incorporating heavier materials in design; alternatively use a pneumatic grab

Netted and kick

netted samplers by water samplers not being able to pass through net Washout of surface

Hess sampler

organisms upon placement of sampler Disruption of substrate surface by Shockwave when sampler strikes bottom

Disturbance of biota

Variable depth of penetration into substrate by sampler

(4225)or a modified corer (683, 3042, 3043) Surber sampler and Underestimation of biota caused by Add screened openings on top Allan grab disruption when sampler is set In place

Shovel sampler

Loss of motile organisms

Grabs

Inconsistent volume of sediment

Surber

sampled: loss caused by overfilling or incomplete closure Failure to consider stream hyporheic zone

Sampler mesh size too

Netted samplers

coarse

Sampler mesh too fine

Netted samplers

Sampler dimension too large

All samplers

Sampler dimension too

All samplers

Early Instars, small and slender organisms missed May cause backwash (see above)

Add screened openings on top Leave 5-cm space above substrate; use a corer whenever possible (3042)

Use two stage sampling for surface and hyporheic zone Use finer mesh; preferably use a double bag sampler (296, 5776, 5493, 6213, 1067) Use coarser mesh as In a double bag sampler (296, 1067, 5493)

May increase sorting time; may reduce Take smaller samples or subsample number of sample units that can be taken

small

Inconsistency or bias of operators

All samplers

Variability increases because of edge effect

Use nested sampler to determine optimal sampler dimension

Repeated, systematic error In taking samples

Use a single operator; or develop correction factor for each operator (1716, 1068) (continued)

Chapter 3 Sampling Aquatic Insects

Table 3B

27

Continued

Factor

Examples of Samplers Affected

Remedy

Problems Created

3. Factors related to characteristics of the environment

From 0.5 to 4 m depth, use an airlift sampler (Fig. 3.40); 0.4 to 10 m deep, use SCUBA and dome suction sampler (2036) or modified Hess sampler (4850, 1673); or use a modified Allan hand-operated grab (82) or

Water depth limitations in lotic environments

Surber and Hess samplers

Because surber sampler limited to 30 for 95% probability level of D In Student's t-distrlbutlon.

t^

(1)riest

(2)ne;

jV

General formula for sample size; if 95% confidence limits of ± 40% of

D^x^

n

_ 255^

^ 2 Ij {t,^[„] + t2(i„p)(v)}

(Sokal & Rohlf 1981, p. 263).

a

= true standard deviation

5

= difference between means expressed as a percent of Y, e.g., = 20 for a 20% difference between means

= desired probability that a difference will be found to be significant

P

= degrees of freedom of the sample standard deviation with a groups and n replications per group

V

faH 3rid

2(1-P)M

= values from a f-table with v degrees of freedom and corresponding to probabilities 1

of a and 2(1-P), respectively. Note if P = t then ta = 0.

e.g., changes along a pollution gradient. A detailed discussion of this type of sample-size determination is presented in Norris et al.(1992). Choice of location where the desired number of

sample units will be taken also is an important consid eration. Sample units could be collected randomly through the study area or they could be collected randomly only within defined strata(termed stratified random sampling), which could be defined by habitat (e.g., a riffle), certain substrate sizes, depth (usually in lakes), hydraulic features (e.g., see Statzner et al. 1988), or a variety of other factors. In practice, strat ification is used to reduce variability or to facilitate comparisons. Concordance of the true population

universe(when the organisms occur)with the sampling universe (where samples are taken) is a goal of stratified sampling approaches. However, one prob lem in using stratified sampling is that extrapolation of trends observed to those expected in other strata is difficult. For more information on stratification, see

Norris et al.(1992)and Resh and McElravy(1993). In some studies, it may be most appropriate to use a transect technique that acknowledges the existence of

known habitat gradients, such as from the margin of a stream or lake to the deepest portion near the center (e.g.. Cummins 1975). If a sample unit contains a very large number of in dividuals in a given taxon, e.g., midge(Chironomidae)

32

Chapter 3 Sampling Aquatic Insects

Table 3F Determination of the number of sample units required to estimate age-specific and total population size of the caddisfly Glossosoma nigrior Banks (Trichoptera: Glossosomatidae) in two first-order Michigan streams. See Table 3E for sample size formula.

Sample Sizes(to nearest integer)for 95% Confidence Limits Where Glossosoma Stream

Augusta Creek (August)

Spring Brook (July)

n

nigrior Age Class

44

s^

D=±40%

i20%

±10%

2.6

16.8

63

249

994

2

13.4

292.4

41

165

652

3

7.8

44.9

19

74

296

4

1.8

4.0

31

124

494

Instar 1

62

x/0.016

5

5.6

19.4

16

62

248

Prepupae Pupae

2.2

5.8

30

120

480

4.5

51.8

64

256

1024

Total

32.8

681.2

16

64

254

Instar 1

0.5

1.0

100

400

1600 459

2

4.3

21.2

29

115

3

6.7

37.2

21

83

332

4

5.3

22.1

20

79

315

5

1.7

4.0

35

139

554

Prepupae and pupae

2.1

6.3

36

143

571

Total

20.6

201.6

12

49

196

larvae, it may be necessary to subsample to obtain an estimate because the time required for a total count is prohibitive (Waters 1969; Elliott 1977; Wrona et al. 1982; Fig. 3.1). Ifthe subsample counts fulfill the criteria of randomness, then a single subsample count can be used to estimate the number in the original sample unit. To satisfy randomness, the mean of at least five subsa mple counts should fall within the 95% confidence interval, that is, a chi-square {%') value between the 5% significance levels for n—1 degrees of freedom

(2(x - 3c)^ found in a standard statistical table

should be obtained. The count should be for the cate

gory of interest, e.g., all taxa or a particular taxon such as a species, age class, or life stage of a species or func tional group. For a further discussion of subsampling, see Elliott(1977). Composite sampling can be used as part of an aquatic insect(or other biological and chemical)sam pling program when the objective is to determine if two (or more) sites are different (e.g., distinguishing stream A from stream B, or an upstream site above a pollution source from a site below the source, or changes in a stream site before and after a distur bance) and knowledge of the variance of each site alone is not critical(USEPA 20021). It can be helpful in reducing the sample processing effort when studies involve many sites, and sites are the replicates to

contrast conditions (e.g., forested versus deforested reaches, Sweeney et al. 2004). The strategy is to characterize the site well (i.e., generate a more accu rate description of average conditions) by collecting a large number of sample units per site (more than the normal 3-5 sample units per sites, e.g., 8-16) in recognition of the natural high variability (e.g., see Table 3F). Composite samples are generated when individual sample units(e.g., random Surber samples) are physically combined, thus homogenizing the variance among those sample units into a new sample (i.e., a composite sample. Fig. 3.46). Because the com posite sample contains more organisms than are needed or can be sorted and identified cost-effec

tively, the composite sample is subsampled randomly to the desired or recommended number of specimens or fraction of a sample unit (e.g., for quantitative samples). The biological or chemical analyses of interest are then performed on the random subsamples or aliquots of the composite sample. Because benthic samples often contain silt, sand, gravel, and even rocks that can damage specimens and make subsampling difficult, it may be better to create 2-4 composite samples based on 8-16 sample units per site, rather than one large composite sample. Finally, sampling other key structural and functional components (Resh et al. 1988) of aquatic insect populations and communities requires special

+

Ash free biomass of

each taxon by size class

detrital particle sb.6 fr^tion *

biomass for each particle size

1r

taxa to coiutant ^

Ash free mass of each

i

mass of eadi size fraction

Calculate, by volume, original

t

size fractions

Weigh detritus

constant mass

size fractions to

Oven dry(50*Q

Ash (5S0*C)by

length-mass regressions

sieve

53

^

Calculate mass from

Microsct^ sort

Collect on

T

1

I

- subsample

Entire sample •

Estimates of microbial

{ Qualitative data on 1 microbial components

I

I size fraction

carbon-nitrogen ratio

lipid, total nitrogen, [

I ^il Pi /g/hr fOT I

I

t each deUitus|

|

Biochemical analyses:!

microscopy

hemiceUutose, ligoin,{

e.g. ceiiuiose,

Determine respiration

I

▼

epifluoresccAce, or associated witih each size scanning electron ffK^ion (3-5 r^licaics)

i Phase contrast.

Remove animals

Resptrometnc

nested sieve

Wash through

Sort by cyc/hand lens for

I

class

of each taxon by size

Numbers and biomass

I

Weight by taxa (nearest O-Sp-g)

(tessicate 24 Ivs

i

Oveo dry size fhKtions (50*C) to constant mass,

measure lengths'

Separ^ taxa and

^ Macroinvertebrates

>- case-bearing Thchopt^a, Mollusca, and Ivge specireeiu

J ))

SAMPLING

content, peroxide digestion may be preferable to combustion. Also, see Chapter 6.

(1922)scale (modified by Cummins et al. [1973]); 2, size traction and subsample volumes should be based on the nature of the sample, especially the density of macroinvertebrates; note also that animals can be classed Into length or weight groups by sieve size (Reger et al. 1982);(Waters[1969b] has devised an efficient subsampler and subsampling is discussed by Elliott [1977]); 3, a number of computer based image analysis systems are available (e.g., Bloquant) that greatly tacilltate measuring; 4, Gllson differential respirometer or oxygen electrode In small circulatory chambers, Bilson (1963); 5, Asmus (1973); 6, long ashing times are required to achieve constant weight, particularly with large detritus samples. It sample contains significant clay

used on all samples, dashied arrows are procedures used only on selected samples or subsamples. Notations: 1, size classes are based on ttie Wentworth

Figure 3.1 A flow diagram summarizing general procedures for analyzing stream or wetland bottom samples. Solid arrows indicate procedures normally

system)'

(luciferin-Iuciferinase

ATP determination

Biochemical

Microbial

Detncal analysis

Wash h through thrc )Un sieve si 250|Un

Wash and decani animus and detritus from sediments

pump or scoop

sediments by hand and tlury of fine sediimots with hand bilge

Sediments ^aerobic zone (approx. iO-20 cm): remove course

) ) )) ) ))) ) ))) ) ))) ) ) 3 ) ) ) J

34

Chapter 3 Sampling Aquatic Insects

attention. Detailed discussions on sampling consider ations for these components are presented in the fol lowing reviews: drift (Waters 1972; Brittain and Eikeland 1988); secondary production (Benke 1984; Rigler and Downing 1984); taxonomic richness(Resh and McElravy 1993); structural, functional, and tro phic diversity (Chapter 6; Cummins and King 1979); nutrient cycling(Webster and Benfield 1986); life-his tory patterns (Butler 1984); size spectra (Clifford and Zelt 1972); biotic interactions (Peckarsky 1984; Pow ers et al. 1988); and biomonitoring (Carter and Resh 2013, 2017, and a variety of papers in Chapter 7). Hauer and Resh (2017) provide detailed exercises of methods and study designs for different types of stud ies, such as distribution and habitat relationships, watershed scale distributions, and population dynamics and movements. They also list materials and supplies needed to conduct these studies. One advantage of studies on aquatic insect dynamics is that many times the choice of study organism can fit the desired study objectives. This opportunity is quite different from the plight of economic entomologists that select their study organ ism based on its pest status. Aquatic entomologists don't always have this freedom (e.g., in the case of

endangered species) but when it is possible to choose a study organism there are several life history features, such as the presence of a single cohort or a univoltine life cycle, that can make study design easier. Likewise, the choice ofa species that is identifiable to species-level for all its life history stages, is abundant (or can be sampled at a scale that makes it abundant), and that shows a positive or negative response to the stressor or other change being studied is a major advantage to any study.

SORTING AND SAMPLE PRESERVATION

Once the sample has been collected, it must be treated according to the nature ofthe substrate materials and the types ofanalyses to be made. We have presented a generalized flow diagram summarizing some general procedures that might be used in analyzing stream bot tom samples (Fig. 3.1). Sorting can be a time-consum ing and, consequently, a costly procedure; the use of elutriation or flotation techniques,sieves,and stains can greatly reduce the time required (see Table 3G, Resh and McElravy[1993],for details). Sorting must be done with care and the advantages gained from adequate sampling designs and appropriate numbers of samples

Table 3G Selected examples of factors related to benthic sorting procedures that may result in sampling bias (modified from Resh 1979a).

Procedure

Potential Problems

Preservation

Alcohoi preservative may cause a weight loss resulting in erroneous biomass and secondary

Live sorting by eiectroshocking

Selective for large organisms (3510)

production estimates(3510, 6502, 1471)

Sieves too coarse

Remedy or Comment Weight loss stabilizes over time so correction factors may be used; alternatively use other preservatives (1471, 3510)or other weighing methods (6502). Useful technique when large amounts of organic matter present

Underestimation of numbers; misinterpretation of

Use finer mesh

life histories

Sieving

Physical damage to specimens; too much material to sieve

Flotation

Flotation time varies with preservative used, taxa present, and instar within same taxa; animals remain

in same fraction as organic detritus Selective against case-bearing caddisflies, molluscs and microbenthos

Sieve under water, without spraying directly on top; elutriation may be preferable (3446, 6482, 3433); use sieve sample-splitter in field (3769)

Live flotation or formalin and sugar solution maximize flotation time; try phase separation technique (304) or centrifugal flotation (1209) Repeated flotation with freshwater washes and examination of remaining material may be necessary

Subsampling

Rare organisms or life history stages may be missed

Subsample size should be adjusted so that smallest taxon counted > 100; subsample by weight (5392)

Counting

Smaller specimens may be missed; too many specimens to count(5509)

Maximum counting efficiency at 25X magnification with transmitted light; try biovolume calculations

Sampling and/or processing time

Costs prohibitive, takes too much time

Use smaller sampler or subsample (1036, 5409); consider redefining study objectives

and costs

Chapter 3 Sampling Aquatic Insects

taken can be obliterated by introduced bias during sorting (Table 3G). If the analysis is to be on a functional feeding group(FFG)basis, sorting of live specimens in the field is most desirable, if practical, because sorting is greatly enhanced before nonstructural colors and behavior are lost (e.g., Chapter 6). Additional information on sorting proce dures is available in Weber (1973), Cummins (1975), Hellawell (1978), Downing (1984), Winterbourn (1985), APHA (1989), Cuffney (1993b), Resh and McElravy (1993), and Carter and Resh (2013). Subsampling of specimens is of importance when sample units or composite samples contain more specimens than are needed,or to be can be sorted and identified cost-effectively. Carter and Resh (2013) discussed how subsampling is done by state agencies. The process ofcreating and subsampling a composite sample is illustrated in Fig. 3.46. Considerations about the use ofsample and spec imen preservatives have undergone major revision because of environmental and individual health con

cerns. Although buffered 5-10% formalin was widely used in the past, ethanol now is usually substituted. For sample preservation in the field, 95% ethanol is often recommended (to account for dilution from water in samples). That field preservative should be replaced with 70-80% ethanol within 1-2 days of col lection to insure good preservation and reduce speci men brittleness; 70-80% ethanol is also recommended

for specimen storage in the laboratory. If there is a possibility that specimens are to be used for DNA analyses, 95% ethanol should be used to preserve all field samples, and the preservative should be replaced with 95% ethanol soon after collection(Sweeney et al. 2011; Stein et al. 2013, 2014; Jackson et al. 2014). Be

aware that ethanol preservation complicates esti mates of insect biomass and production because this fluid dissolves stored fats reducing specimen mass (Benke et al. 1999,Table 3G).Proper disposal ofspec imens and preservatives is essential and should be in accordance with your institution's approved hazard ous-waste disposal program. In some cases, it might be suitable to store samples frozen when preservatives are not available. However, these specimens tend to be of poor quality and careful, slow thawing is neces sary to minimize the breakage ofspecimens that often results from freezing.

TAXONOMIC IDENTIFICATIONS AND REARING METHODS

Most ecological studies of aquatic insects have dealt with their immature stages because it is the larva or nymph that normally occurs in aquatic

35

habitats and represents the major portion of insect life cycles. The identification of most aquatic immatures is difficult because: (1) taxonomic names are generally based on characteristics present in the adult stage, (2) for many North American species, immatures and adults have not been associated, and

(3) insufficient comparative analysis of congeneric (i.e., in the same genus) immatures has been com pleted to produce species-level keys. Even identify ing immature aquatic insects to the genus-level has its limitations. For example, the keys to aquatic insects presented in this book are most reliable if used on late instar individuals because of changes that occur in morphological structures as the larva or nymph matures. The required level of identification is essentially a reflection of study objectives(Resh and McElravy 1993). For stream bioassessment, a controversy has developed over the relative values of family vs. genus- or species-level identification of aquatic insects. Bailey et al.(2001) argue that in many cases species- or genus-level classification provides only a minimal amount of additional information for clas

sifying habitat impairment, and the additional costs and time investment for these detailed identifica

tions may not be warranted. Lenat and Resh (2001) counter by saying that while species- or genus-level identification initially requires more time, the time investment declines as people become familiar with the fauna. They maintain that the value of speciesor genus-identifications more than compensates for these initial costs. Species-level identifications are important in ecological studies because congeneric species do not necessarily have identical ecological requirements or water quality tolerances. Also, the inability to distinguish between coexisting species may mask population dynamics or trends, and, without species identification, comparisons with results obtained from other studies (possibly even with related species) are difficult (see individual order chapters for exceptions). The calculation of diversity indices, a technique often criticized but still widely used in aquatic insect community studies, may result in significant underestimates when generic- or family-level identifications, rather than those made at the species level, are used or applied in an uneven fashion to different higher taxa (Resh 1979b). An additional problem is what Carter and Resh (2013) refer to as unresolved taxa in calculat ing species richness from a collection. For example, an identification of Baetis tricaudatus could be done

for a fully grown nymph, but Baetis sp. or Baetidae may be the extent of the morphological identification that could be done for small and young specimens.

36

Chapter 3 Sampling Aquatic Insects

Consequently there is a need for consistency in dealing

organisms for bioassay tests (e.g., Anderson 1980;

with this issue when it arrives, which is often in

Buikema and Voshell 1993). The methods outlined by Lawrence (1981) and references in Table 3H

benthic studies.

The taxonomic problems of identifying the immature stages of aquatic insects have traditionally been solved either by rearing the larva or nymph to the adult stage, or in some groups by collecting asso ciated adult and immature stages. For example, asso ciations can be made by examining mature pupae (Milne 1938) and cast larval skins (e.g., Trichoptera), and by collecting exuviae in organic foam accumula tions, drift (streams), or windrows (lakes)(e.g., Chironomidae; Coffman 1973). Rearing techniques range from very simple to highly complex, and no single technique is suitable for all aquatic insects or even all species of a given genus or family. In Table 3H, gen eral references to techniques for obtaining adult stages of immature insects from lotic and lentic habi tats are given, as are rearing (i.e., a single generation) and culturing (i.e., rearing through subsequent gener ations) methods appropriate for the different orders of aquatic insects. Taxonomic identifications, associations, and

relationships have also benefited from genetic analy ses of immatures and/or adults, starting first with allozyme (protein) analyses (Zurwerra et al. 1986; Lees and Ward 1987; Sweeney et al. 1987; Funk et al. 1988; Funk and Sweeney 1990; Sperling and Spence 1990; Jackson and Resh 1992, 1998), and more recently molecular DNA analyses based on local (Monaghan et al. 2005; Xhou etal. 2010, 2011; Gill et al. 2013), regional(Xhou et al. 2007; Moriniere et al. 2017), or global efforts (Kjer et al. 2001; Ogden and Whiting 2005; Holzenthal et al. 2007). Taxonomic identifications based on genetic analyses are now being used in a variety of ecological studies, including biomonitoring efforts (Hajibabaei et al. 2011; Swee ney et al. 2011; Jackson et al. 2014; Stein et al. 2014; Macher et al. 2016).

Published reports of rearing techniques gener ally fall into three categories: (1) descriptions of various running-water systems (artificial streams; see discussion in Vogel and LaBarbera 1978), (2) methods of maintaining larvae and pupae until emergence occurs, and (3) methods of obtaining eggs from adult females and then rearing the newly hatched larvae as in (2). Because lotic insects are often more difficult to rear than lentic ones,a greater number of techniques has been published on the former. In recent years, culturing methods (i.e., for continuous generations) have been improved, largely in response to the need for maintaining

should be consulted for more detailed information

on the approaches used. The subject of culturing of invertebrates for bioassays has been covered by Buikema and Cairns (1980). Field rearings are generally more successful than laboratory rearings, but are often impractical because of the time or frequency required to be on-site. The choice of mass(many species per con tainer) versus individual rearings is based on the degree of similarity among immatures being reared. Collection of adults in the vicinity of the immature aquatic habitat with sweep nets or light traps can give some idea of the presence of system atically related species whose immature stages may not be distinguished easily from those under examination. In the River Continuum studies and

those at Oregon State, large screen cages were used to collect adults emerging from a set area of stream bottom. The adults were collected by vacuuming the inside of the cages (K. W. Cummins, pers. comm.). This information may help avoid the situation in which two or more species of adults emerge from a presumed single-species rearing. An alternative approach is to obtain eggs from known females and rear these to maturity (e.g., Resh 1972). One of the most common problems encoun tered in rearing aquatic insects involves mortality during transport from the field to the laboratory because of inadequate oxygen supply and/or tem perature control. Agitation during transport will

maintain oxygen levels but may damage delicate specimens. Alternative methods include transport ing the animals in damp moss (excess water removed), burlap, or paper towels; using small "bait bucket" aerators; or attaching tubing to an exterior funnel that can pick up a "wind stream" while a vehicle is moving. To maintain cool tem peratures, thermos containers or ice coolers should be used.

Laboratory rearings can be maintained at field temperatures using an immersible refrigeration unit or by recirculating water through a cooling reservoir. If laboratory temperatures do not match those in the field, mortality can be reduced by allowing tempera tures to equilibrate slowly. To maintain water quality in the laboratory, tap water should be dechlorinated and distilled or spring or stream water added to replace evaporative loss.

Chapter 3 Sampling Aquatic Insects

Table 3H

37

Selected references on aquatic insect rearing methods.

Laboratory Culture

Immature to Adult Rearing Methods

Methods

Laboratory Figure(s)

Field Order

References

Figure(s)

Lentic insects in most

References

References 209*

3.37

521, 1178*, 1180, 3432, 2380, 3770, 5342, 3088

3.38-3.39

1376, 1977, 4251,4595

3.37-3,39

411, 1977, 1978, 4222,

365*, 851,3656*, 4596,

3.38

2236, 3316, 3656, 5290,

orders^

Rheophiiic (current-loving)

3.35

insects in ail orders

5602

Coilembola

Ephemeroptera

865, 1032*, 2659, 3.34

5190

2944

Odonata

2639

3.35

6448

6448

Plecoptera

1262, 1787

3.34

Hemiptera Trichoptera

2198

3.34-3.35

674, 1987, 1988, 2067, 2827*, 3060, 4610, 6770

3.37-3.39

521, 3060

365*, 3263, 3385, 3967, 3386, 3969, 3970, 4487

3.37

1653, 2679, 2953

157, 158, 159, 166, 2586, 3.37-3.39 4685, 4934, 5524, 6512

Neuroptera

731, 4439

Megaloptera

4822*, 5515*

Lepidoptera Hymenoptera

3.35

161, 521, 4934

3396

See methods for rearing specific hosts

Coleoptera Lentic species

36, 149, 365*, 6806

Lotic species

737, 6458*

737, 6458

Diptera 3127, 3545, 5970

Ceratopogonidae Chironomidae

3.36

365, 654, 1599

2322, 3543 3.36

505, 1199, 1494, 3989, 5873

Dixidae

2177

Culicidae

1989, 2090*

Sciomyzidae Simuliidae

1768,2090, 5514*

4292 2933

904, 2329, 2443, 5879*, 5880, 5962

662*, 1811, 1959,4180, 4181,4884, 5454, 5455, 6708, 6709, 467*, 1582, 6755

3393, 3719, 5072, 5073, 5365, 5936

Tabanidae

Tipulidae

2933

Parasitic mites on aquatic insects

*Recommended techniques.

"•"Hemimetabolous or with.aquatic pupai stage.

2519*, 5105 1220, 1221, 4957

Figure 3.3

Figure 3.6

Figure 3.5

Figure 3.4

Figure 3.2

Figure 3.8

Figure 3.7

t

y

Figure 3.10

Figure 3.12

Figure 3.13

Figure 3.15 Figure 3.14

Figure 3.11

Figure 3.2 Figure 3.3 Figure 3.4 Figure 3.5 Figure 3.6 Figure 3.7 Figure 3.8 Figure 3.9 38

D-frame aquatic net. Hand screen collector. Hand dipper. Wilding or stovepipe sampler. Modified Hess sampler. Surber sampler. Stream bottom T-sampler. Ellls-Rutter Stream sampler.

Figure 3.10 Drift net. Figure 3.11 Plankton tow net. Figure 3.12 Core sampler with pole. Figure 3.13 Kellen grab. Figure 3.14 Multiple core sampler. Figure 3.15 Single core sampler.

Chapter 3 Sampling Aquatic Insects

39

mm

Figure 3.19 Figure 3.24

Figure 3.22

Figure 3.16

Figure 3.25 Figure 3.20

Figure 3.23

Figure 3.17

Figure 3.26

Figure 3.18

Figure 3.16 attachment). Figure 3.17 Figure 3.18 Figure 3.19 Figure 3.20 Figure 3.21 Figure 3.22

Figure 3.21

Ekman grab (with and without pole Ponargrab. Allan grab. Petersen grab. Macan sampler. Gerking sampler. Modified Gerking sampler.

Figure 3.27

Figure 3.23 Multiple-plate artificial substrate sampler. Figure 3.24 Basket-type artificial substrate sampler. Figure 3.25 Leaf pack sampler. Figure 3.26 Mundie pyramid trap. Figure 3.27 Emergence traps: A, submerged; B, floating pyramid; C, staked box.

40

Chapter 3 Sampling Aquatic Insects

plastic cup witti lid

nylon mesh

styrotoam

netting

Figure 3.34 mosquito netting wood stick

giess

■\

wire screen

blocks

Figure 3.31

Figure 3.36

Figure 3.28

to air pump

mosquito netting

stones

Figure 3.37

mosquito netting

to air pump

Figure 3.29

Mason iar- -^--_

Figure 3.32

Figure 3.38 recirculetion

pump

Figure 3.30

Figure 3.33

aquarium

Figure 3.39

Figure 3.28

Subaquatic light trap.

Figure 3.29

Malaise trap.

Figure 3.30 New Jersey light trap. Figure 3.31 CDC light trap. Figure 3.32 Beriese-Tuiigren funnel. Figure 3.33 Graded sieves. Figure 3.34 Floating cages (drawn after Edmunds etal. [1976]). A mesh lining attached along the Inside wall of the cups will allow the subimagos to cling to the side and not slip back into the water.

Figure 3.35 Pillow cage (drawn after Peterson [1934]). The inclusion of larger stones to serve as ballast may prevent the pillow cage from being washed away. The top portion of the cage must be above the water surface.

Figure 3.36 Vial rearings (drawn after Peterson [1934]). Fungal growth will be retarded if distilled water is used and the temperature is kept 16°C or lower. Figure 3.37 Aquarium rearing method. Figure 3.38 Quart jar rearing method. Figure 3.39 Artificial stream rearing design.

Chapter 3 Sampling Aquatic Insects

41

30

IV ■

B"

"U '

Figure 3.42

Figure 3.41

Figure 3.40

Figure 3.43

Figure 3.44

/=

_ Emergence Chamber

Rehydraied Substrate

Figure 3.45

Figure 3.40 Modified air-lift sampler (redrawn from Norris [1980]). Figure 3.41 Hyporheic canister (implant) sampler (redrawn from Gilpin and Brusven [1976]). Figure 3.42 Hyporheic standpipe corer (redrawn from Williams and Hynes [1974]). Figure 3.43 Benthic invertebrate elutriation apparatus (redrawn from Worswick and Barbour [1974]).

Figure 3.44 Activity trap (from Batzer et al. 2001, with permission of John Wiiey and Sons). Figure 3.45 Insect emergence chamber for rehydrated wetland soil (from Wissinger and Gallagher 1999, with permission of John Wiley and Sons).

42

Chapter 3 Sampling Aquatic Insects

1

3

5

6

7

Composite

Algal and detrital food supplies are often best maintained by periodic replenishment from the field. Detritivores that eat leaf litter (shredders) require conditioned material, i.e., leaves colonized by aquatic hyphomycete fungi and bacteria. Wheat and other grains can be used to supplement the diets of detritivores, and scrapers can fre quently be fed on spinach leaves. The addition of

Subsamples brought back from field

Subsamples to be

analyzed I Figure 3.46 Illustration of combining four sample units (e.g., Surber samples) to form a composite sample that is first subsampled in the field, and then subsampled again in the laboratory. The subsample size (proportion) is determined by the number of insects present.

OC

enchytraeid worms or wheat grains to a detri tus-based diet not only reduced development time but also increased the weight of individuals in a limnephilid caddisfly culture (Anderson 1976; Hanson et al. 1983). Larvae of Drosophila, house flies, mosquitoes, and tubificid or enchytraeid worms can serve as food for predators. The most critical problem in rearing aquatic insects that require a highly specific food (e.g., freshwater sponges, bryozoans) may be in the culture of the food source itself. Mortality can be reduced by choosing only immatures close to emergence or larvae about to pupate. Adjustments of photoperiod (using a light/dark regime similar to that in the field during emergence)or temperature may be required to break the arrested development of spe cies that undergo diapause.

AQUATIC INSECT RESPIRATION David B. Buchwalter

North Carolina State University, Raleigh

Gary A. Lamberti University of Notre Dame,Indiana

Vincent H. Resh

Wilco C.E.P. Verberk

University of California, Berkeley

Radboud University Nijmegen, The Netherlands

INTRODUCTION

Biologists have been fascinated for centuries by the diversity of ways in which organisms have solved life's most fundamental challenges. Respiration (the process of converting nutrients to energy) is one such challenge. The overwhelming majority of animal life requires oxygen (O2) in cellular respiration, because

serve as the structural plan for their aquatic respira tory system as well, which we will explore further in this chapter.

An incredible array of respiratory strategies and morphologies evolved within and across the aquatic insects in response to the challenges associated with life in water. This book contains numerous examples

O2 is the final electron acceptor in a series of mito-

of the morphological diversity of aquatic insect respi

chondrial reactions that release energy from organic molecules obtained from digested food. Although most if not all insects are also capable of generating

ratory systems, many of which are important diag nostic features used by taxonomists to differentiate

some energy without O2, such anaerobic metabolism

lenge of breathing in water, provide an overview of

releases about 15-fold less energy and often generates toxic metabolites. When O2 is limiting, animals may suffer asphyxiation but equally, too much O2 can be toxic. Toxic effects of O? arise as radicals are pro duced in the mitochondria in an 02-dependent man

the various respiratory modes that exist in aquatic insects, and discuss important functional aspects of

between taxa. In this chapter, we highlight the chal

respiration in relation to the diversity of thermal and

O2 environments found in freshwater ecosystems.

ner. Thus, an animal must be able to reduce the risk

of O2 toxicity, while at the same time retaining suffi

OXYGEN: FROM SOURCE TO CELL

cient scope for O2 uptake across different levels of

Oxygen in the Environment When water and air are in contact,an equilibrium

activity to avoid asphyxiation. As a result, aquatic insects and indeed almost all animals depend on a continuous and adequate supply of O2 to meet energy demands associated with locomotion,feeding,growth,

is established between the gaseous components of each. Some air will dissolve in the water, with the

amount depending on the partial pressure of the gas

and reproduction.

eous component and its solubility in water. The gases

All aquatic organisms that rely upon dissolved O2 to respire share a common challenge—namely that there is substantially less O2 in water than in air (see

and thus their proportions and absolute amounts in water and air are quite different. For example, air

that make up air have different solubilities in water,

below). However, aquatic insects differ from the vast

contains approximately 21% O2 and 78% nitrogen

majority ofother aquatic organisms(e.g., fish, crusta ceans, bivalves) in that they are secondarily aquatic (Bradley et al. 2009; Misof et al. 2014). Insects origi nally diversified on land where they developed a gas-

(N2), but the water solubility of O2 is greater than that

of N2. Consequently, O2 typically makes up -33% of the gases normally dissolved in water, while N2 is less than 67%. Carbon dioxide(CO2)is highly water solu

filled (tracheal) respiratory system (see Chapter 9).

ble and although only 0.03% of air, it can be almost

Over evolutionary time, several invasions offreshwa ter habitats occurred,sometimes even more than once

3% of the gases dissolved in water. (Note that the

in a given insect lineage. As insects adapted to aquatic environments, their air-filled tracheal system had to

perature, atmospheric pressure, and the buffering capacity and hardness of the water.) However, even

exact percentages of dissolved gases depend on tem

43

44

Chapter 4 Aquatic Insect Respiration

though O2 is more soluble in water compared to some

other gases (e.g., N2), its absolute amount in water is small compared with that in an equal volume of air. On a volumetric basis, O2 is more plentiful in air than in water. For example, 1 L ofair contains 209.5 mL

of O2, compared to 9.1 mL (equivalent to 12.8 mg)of O2 in a fully saturated liter of water at 5°C—which is a 23-fold difference. On a mass basis, air contains four

orders of magnitude more O2 than water. This lower amount of O2 in water (relative to air) requires that

However,photosynthesis by phytoplankton, periphyton (attached algae), and macrophytes may cause

warm, algae-rich ponds to become supersaturated in O2 during daylight hours. At night, however, O2 lost to community respiration is no longer replenished by photosynthesis and O2 levels in the pond may decline drastically. Thus, the normally higher daytime and lower nighttime dissolved O2 levels clearly reflect the daily cycle in photosynthesis and its relation to the continual respiration of the aquatic community. The ratio ofdaily gross photosynthesis to daily respiration

aquatic insects move much more water across their respiratory surfaces to extract the same amount of O2. Therefore, as a result of the higher density and

(P/R) often is used as an index of aquatic community metabolism (Cummins 1974; see also Kosten et al.

viscosity of water, aquatic insects require much larger

2014).

effort to ventilate their respiratory surfaces com

Oxygen often is nearly absent in groundwater because ofthe bacterial respiration that occurs during its slow movement through the soil and typically long residence time in the aquifer. Once groundwater comes to the surface, O2 is replaced at a rate deter mined by local conditions, especially current, turbu lence, and primary productivity. Cold groundwater

pared to their terrestrial counterparts. In addition, processes that consume (respiration) or produce (photosynthesis) O2 can result in large fluctuations in the availability of the aquatic Ojsupply and in some habitats O2 may be low (hypoxic) or even totally lacking (anoxic). Environmental conditions can strongly affect the

solubility of O2 in water. Increasing temperature and salinity decrease the solubility of O2 and thus reduce levels of dissolved O2. At high altitudes, where the total atmospheric pressure is reduced, the partial

pressure of O2 is lower. However, dissolved O2 levels tend to be similar across altitudinal gradients. This is

because at high altitudes, the lower partial pressure tends to be counterbalanced by the higher solubility of cold water (Jacobsen 2000). Water turbulence

entering streams is usually quickly re-aerated but in ponds or stratified lakes, groundwater may remain depleted in O2 until an equilibrium with the surround ing water is established. More-detailed treatments of O2 in aquatic environments are given in Hutchinson (1957), Hynes (1970a), Wetzel (2001), Cole (1994), Lampert (1997), Kalff (2002), Dodds (2003), and Verberk et al.(2011).

Obtaining O2 from the Environment

(mixing flow in which velocity and direction vary unpredictably over very short distances)enhances the exchange of O2 by increasing the water surface area, forcing aeration, and moving water with lower O2

Because the insect respiratory system evolved to obtain and transport the abundant atmospheric O2,

concentration to the surface. In highly turbulent

encounter O2 limitations (though they have to make

waters, dissolved O2 concentrations may even exceed those expected in an equilibrium (which results in a situation called supersaturation) as a result of vigor

that rely on O2 dissolved in water face a number of difficult problems and, as described below, have

ous aeration. Once dissolved,ifO2 were distributed by

evolved multiple ways of coping with these issues.

aquatic insects that use air directly do not typically air contact to be able to breathe). In contrast, insects

diffusion alone in water that was devoid of this gas,

Whatever the environmental conditions that

years would be required for even traces of O2 to reach

insects inhabit, the process of diffusion ultimately

several meters in depth, as a result of the low diffusiv-

moves O2 and other gases both to and through respi ratory surfaces. Gases diffuse down the gradient of partial pressure, and these partial pressure gradients frequently coincide with concentration gradients.

ity of O2 in water. Thus, wind-generated currents and turbulence are vital to the mixing of gases within the water column. In fact, gas exchange and mixing are

significantly reduced by anything that inhibits the effect of wind on water, such as low water surface area, pro tective vegetation, or ice cover.

Typically, dissolved O2 levels are higher in flow

ing water than in still aquatic environments, such as

However, this does not need to be the case when the solubility of O2 also differs across the gradient. For

example,in insect species that use respiratory proteins such as hemoglobin to increase the solubility of O2, the hemolymph may contain more O2 than the sur

water turbulence (although organic matter or sewage

rounding water. Despite the high concentration of O2, the partial pressure of O2 in the hemolymph can still

pollution may reduce O2 concentrations in either).

be lower than that of the surrounding water; in this

lakes, because ofthe enhanced mixing associated with

Chapter 4 Aquatic Insect Respiration

45

case O2 still diffuses down the gradient of partial pressure but against the concentration gradient. The diffusion speed is described by Pick's law, which considers the solubility, the pressure gradient, a diffusivity coefficient, and the surface ofthe respiratory area, and is then divided by the distance (i.e., the

flattened shape or small size, a number of aquatic insects can dwell within the boundary layer that is

combined thickness of the cuticle and boundary

water flow to obtain adequate O2 for respiration.

layer). The diffusivity coefficient depends on the molecular weight of the gas and the permeability of the medium through which it must pass. In water, the diffusivity coefficient of O2 increases with tem perature. As a consequence, maximum rates of O2 diffusion may increase slightly with temperature, even though the O2 solubility decreases (see Verberk

O2 Environments

et al. 2011).

O2 diffuses rapidly through air but not in water. In fact, the diffusion coefficient is ~10,000 times smaller and the solubility is 20-30 times lower, result

ing in diffusion rates that are about 300,000 times slower! If that is not problem enough, insect cuticle reduces the O, diffusion rate even more, slowing its

created as stream water flows over rock surfaces.

Within the boundary layer, an insect is removed from significant water movement and abrasion by small particles but it remains close enough to turbulent

Respiratory Mechanisms and The evolution of insects from a terrestrial to an

aquatic existence has resulted in a broad range of adaptations of the original, terrestrial-insect respira tory system (see also Chapter 9). Aquatic insects have developed major structural and behavioral adapta tions to the various habitats with seemingly endless variations. Among these adaptations, respiratory structures and behaviors have evolved to allow insects

to occupy virtually every aquatic habitat regardless of O2 concentration. Pools in flowing water and other

speed by almost 850,000 times compared to the rate of

still-water habitats(Chapter 5) possess such a tremen

diffusion of O2 in air(Miller 1964a). The distance over which the diffusion gradient exists includes not only the cuticle-tissue thickness, but also the thickness of the adjacent layer of water (Fig. 4.1). This adjacent layer of water is termed the boundary layer and is defined as the layer in which water movement is

dous variety of microenvironments that insects with most of the aquatic respiratory adaptations can be

restricted as a result of the frictional resistance that

results from the viscosity of water. As water moves

over a respiratory surface (or any surface), frictional resistance slows the adjacent water molecules until flow ceases at the surface. The thickness ofthe bound

ary layer is dependent on the flow speed. At very slow flow, viscosity effects are more important and increas ingly thicker boundary layers develop. Boundary layers can significantly impede gas exchange, as gases must move through it by diffusion alone (Ambuhl 1959; Feldmeth 1968; Vogel 1988), whereas outside the boundary layer, movement of O2 by convection also takes place. Because the thickness of the boundary layer decreases as the rate of flow in the adjacent water increases, both the water velocity and an organism's self-generated ventilation currents (e.g., moving its abdomen or gills) can make the boundary layer thinner. In the true bug Aphelocheirus

found there.

The deep parts of lakes lack aquatic insects that must return to the surface to obtain O2 from the air because of the dual problem of having to travel to the surface to renew their O2 supplies and losing O2

through diffusion to the surrounding 02-poor water when they return to the lake bottom. Those insects that do occur in the depths oflakes commonly possess hemoglobin and have strong ventilatory abilities(e.g., chironomids) or perform diurnal migrations (e.g., chaoborid midges) and often have metabolic adapta tions for managing the byproducts of anaerobic metabolism. Earlier and smaller life stages, along with the few species that inhabit the open water (away from the shore and bottom)of ponds and lakes, often have a high surface-to-volume ratio and consequently use cutaneous respiration. The constantly high-Oj habitat of rapidly flowing water is suitable for species using plastrons, spiracular

gills, cutaneous respiration, or tracheal gills (see below for descriptions). Those insects that require

aestivalis, which is common in European streams,

direct air contact cannot live in rapid flow because stable connection with the atmosphere is usually

such body movement was found to appreciatively thin the boundary layer, enabling approximately a fivefold greater rate of O2 uptake in active compared to inactive animals (Seymour et al. 2015). Neverthe less, even in the most rapid flows that occur in streams or that are produced by the movement of the organ ism, a thin boundary layer persists. Because of their

impossible given the turbulence of the water. Some diving beetles (Dytiscidae) that typically rely on sur facing for gas exchange have solved this problem by having dense setae (Kehl and Dettner 2009). These hollow setae on their elytra interconnect with the bee tle's tracheal system, providing a gas exchange surface that allows O2 to be extracted directly from the water.

46

Chapter 4 Aquatic Insect Respiration

Setal gills allow the beetles to circumvent the diffusion barrier inherent to their thick exoskeleton (see above) and enables them to perform underwater gas exchange that is functionally similar to the cutaneous respira tion seen in many other aquatic insects larvae. If flow is sufficiently rapid (leading to low diffusion resis tance due to a thin boundary layer), these dytiscid beetles can stay submerged indefinitely, which offers distinct advantages (e.g., increased feeding and mat ing, and reduced risks of predation and being swept away by fast water currents). More specific general izations cannot be made because the major habitats that aquatic insects occupy encompass so many dif ferent O2 microhabitats. Chapman et al. (2004) dis cuss respiratory adaptations in tropical environments.

Tracheal System and Respiratory Surfaces In most insects, either terrestrial or aquatic, gas distribution takes place through a network of inter nal, air-filled tubes known as the tracheal system (Chapman 1982). The larger tubes, or tracheae, exchange respiratory gases with the atmosphere through segmentally arranged lateral pores called spiracles. Tracheae, which are cuticular ingrowths, branch internally from the spiracles and become pro gressively smaller (to 2-5 pm in diameter). Further branching forms capillaries called tracheoles, which are generally less than 1 pm across and terminate where they contact individual cells or cell clusters. Gases are transported by a combination of diffusion and convection within the tracheal system but, at the

tracheoles, O2 enters the cells by diffusion. To aid convection in the air-filled tracheal system, some large or highly active insects can advantageously ven tilate at least the outer portion of their tracheal sys tem. In such species, contraction of abdominal muscles compresses flexible air-sacs that are located

along the longitudinal tracheal trunks (Fig. 4.2). Compression empties the air-sacs and flushes their contents through the tracheal trunks and out the spir acles. When abdominal muscles relax, higher external atmospheric pressure forces air back through the spir acles and tracheal trunks to refill the air-sacs.

consumption that appears to be compensatory—i.e., repaying the O2 debt incurred during the molt(Camp et al. 2014). Most terrestrial and some aquatic insects have

multiple pairs of spiracles(usually 8-10)that open on the body surface, which are referred to as polypneustic systems (Fig. 4.2A and B). This tracheal respira tory system is regarded as the ancestral design because insects evolved terrestrially and later was modified by evolution in various ways as insects adapted to an aquatic lifestyle. Oligopneustic systems, which devel oped from the ancestral, polypneustic design have only one or two pairs of functional spiracles, often located at the posterior end (Fig. 4.2C). Both designs are often referred to as open tracheal systems because of the presence of functioning spiracles. In contrast, tracheal systems with no functional spiracles are referred to as closed or apneustic (Fig. 4.2D-F) and, although otherwise complete, lack direct contact with the outside water and rely on gases diffusing through the cuticle for respiratory exchange. The rate of O2 exchange is partially determined by the amount of surface area through which gas molecules can pass. In water, where rates of O2 diffusion are low, larger respiratory surfaces are generally required for O2 sup ply to meet demand. The body surface of a small, elongate organism (e.g., early instar chironomid midges. Fig. 27.5) may be large enough to allow suffi cient O2 diffusion to meet the organism's metabolic needs. However, as that animal increases in size, its

body volume (and hence O2 demand) will increase more rapidly than its surface area and this may limit O2intake. However,long before such limitations, ani mals tend to compensate by behavior, morphology, and physiology, to prevent O2 intake from becoming surface-limited. Some insects employ additional gas exchange surfaces, such as large, thin, tracheated body outgrowths called gills (Fig. 4.2E and F) that serve to counter this trend. Other insects possess air bubbles that can be replenished by O2 diffusion (Fig. 4.3). As an insect grows, these surfaces often grow disproportionally large in order to maintain a suitable surface-to-volume ratio that can meet the

insect's respiratory needs.

One of the more remarkable features ofthe insect

tracheal system is the fact that when the insect molts, the lining of the tracheae is shed along with the old exoskeleton(Tower 1906; Snelling 2011). This process can be accompanied by a significant disruption in O2 consumption. For example, Cloeon dipterum and other mayfly nymphs experience a transient(~46-60 minutes) but marked reduction in O2 consumption rates during the molt. This bout of reduced O2 con sumption is immediately followed by a sharp spike in O2

RESPIRATORY OPTIONS WITH AN OPEN TRACHEAL SYSTEM

Open tracheal systems are characteristic ofinsects

that breathe air. Aquatic insects with open tracheal systems must therefore establish direct spiracle-to-air contact, either by connecting directly with a station ary air source (e.g., the atmosphere) or by carrying a store of air when they dive (Table 4A).

Chapter 4 Aquatic Insect Respiration

47

O2 concentration gradient TURBULENT FLOW

dependent upon turbulence of the water

' Effective limit of

boundary layer

O2 concentration gradient LAMINAR FLOW

dependent upon

O2 diffusion rate

O2 CONCENTRATION Figure 4.1

Figure 4.2

Figure 4.1 The boundary layer and its effect on O2 reaching an organism's respiratory surface. The dashed line demarcates the boundary layer's outer limit. An O2 pressure gradient (solid line) is established because O2 is consumed at the organism's surface and is replaced within the boundary layer only by diffusion. The diffusion rate is dependent on the thickness(D) of the boundary iayer and the steepness of the O2 gradient (modified from Feldmeth 1968).

Figure 4.2 A. Open polypneustic tracheai system; B. Polypneustic tracheai system with air-sacs for ventiiation; C. Oligopneustic tracheai system in which the terminal spiracles alone are functional; D. Closed tracheai system allowing cutaneous respiration oniy; E. Closed tracheai system with abdominal tracheai gills; F. Closed tracheai system with rectal tracheai gills (modified from Wigglesworth 1972).

48

Chapter 4 Aquatic Insect Respiration

Figure 4.3

Figure 4.4

9%

Figure 4.5

Figure 4.6

Figure 4.3 Ventral air bubble, which also serves as a temporary physical gill, of the pleld backswimmer Neoplea (after GIttelman 1975). Figure 4.4 Longitudinal section of postabdominal respiratory siphon of a Taeniorhynchus (Culicidae) larva. Barbed hooks allow the larva to maintain contact

between the spiracle and the plant air stores (modified from Keilin 1944).

Figure 4.5 Hydrofuge hairs comprising the plastron of the aphelocheirid bug Aphelocheirus (after Hinton 1976a).

Figure 4.6 Hydrofuge cuticular network found in the spiracular gill of the tipulid Dicranomyia (after Hinton 1968).

Chapter 4 Aquatic Insect Respiration

49

iili Table 4A Respiratory options with open and closed tracheal systems. The life stages known or inferred to use a particular respiratory option are indicated by the following: L=larvae: N=nymphs; P=pupae: A=adults. Wichard eta/. 2002 provide many excellent examples, including illustrations of respiratory adaptations in various aquatic insects.

Respiratory Option

Atmospheric Breathers

Trachea!

Oxygen

System

Source

open

atmosphere

Examples

Selected

References

DIptera: Culicldae (L, P), Dollchopodidae (L), 2661, 2839, 4159 Ephydrldae (L, P), Psychodldae (L), Stratiomyidae (L, P), Syrphldae (L, P), Tabanldae (L, P), TIpulldae (L, P), Ptychopterldae (L, P) Coleoptera: Amphlzoldae (L), Dytiscidae (L, A), Hydrophllldae'(L, A) Hemiptera: Nepldae (N, A)

Plant Breathers

open

plants

Coleoptera: Chrysomelidae (L, P, A),

2345, 2346, 2347,

Curculionidae (L)

5291

DIptera: Culicldae (L, P), Ephydridae (L, P), Syrphldae (L) Temporary Air Store

open

atmosphere and dissolved

Coleoptera: Dytiscidae (A), Gyrlnldae (A), Hallplldae (A), Helodidae (A), Hydraenldae (A), Hydrophllldae (A)

1388, 3907, 3908, 4122, 4196

Hemiptera: Belostomatldae (N, A), Corlxidae (N, A), Naucorldae (N, A), Notonectldae (N, A), Pleldae (N, A) Permanent Air Store Plastrons

open

dissolved

Coleoptera: Curculionidae (A), Dryopidae (A), Elmidae (A), Hydraenldae (A), Hydrophllldae (A) Hemiptera: Naucorldae (N, A)

2066, 2067, 2286, 5145, 5146, 5147, 5148, 5149

Lepldoptera: Pyralidae (L, P) Spiracular Gills

open

dissolved

Coleoptera: Hydroscaphldae (L), Psephenldae (P), Sphaerlidae (L), Torrldincolidae (L, P)

2282

DIptera: Blepharicerldae (P), Canacldae (P), Deuterophleblldae (P), Dollchopodidae (P), Empldidae (P), Simullidae (P), Tanyderidae (P), TIpulldae (P) Tracheal Gills

closed

dissolved

Ephemeroptera (N), Odonata (N), Plecoptera (N), Megaloptera (L), Neuroptera: SIsyrldae (L), Coleoptera (several families), DIptera (several families), Trichoptera (L), Lepldoptera: Pyralidae (L)

Cutaneous

closed

dissolved

Diptera: Ceratopogonldae (L, P), Chaoborldae (L, P), 1247, 1670, 2911, Chlronomldae (L, P), Simullidae (L), TIpulidae (L) 4159, 3901, 5455

1456, 1457, 1458, 2911, 3556, 5596, 5743, 1460, 1462, 3780, 4690

Lepldoptera Plecoptera (gill-less N) Trichoptera (gill-less L) Hemoglobin f

open or

atmosphere

closed

or dissolved

Hemiptera: Notonectldae (N, A) Diptera: Chlronomldae (L, P)

closed

dissolved

Plecoptera (N)

3502, 5419, 5420, 5421

1

Hemocyanin

Stationary Air Sources Aquatic insects that connect with a stationary air source have an oligopneustic tracheal system, with the functional spiracles located at the end ofthe abdomen of larvae or on the thorax of pupae. The submerged insect obtains O2 either by placing its spiracles above

the water surface (atmospheric breathers) or by forc ing them into plant air stores (plant breathers). Atmospheric breathers seldom maintain a continu ous connection with their air source. Therefore, spira cles must be adapted to prevent flooding both when the insect submerges and when it comes to the surface and

50

Chapter 4 Aquatic Insect Respiration

to contact the air. Spiracles are commonly surrounded by a water-repellent (hydrofuge) cuticle or by waterrepellent hairs (Figs. 23.51 and 23.54). Upon submer

gence, flooding is prevented by these hydrofuge hairs, by retractable fleshy lobes that seal the spiracular open ings, or by holding an air bubble over the openings. Many aquatic insect families contain atmospheric breathers (Table 4A). Undoubtedly, culicid larvae, commonly called mosquito wrigglers, are far and away the most familiar of the oligopneustic atmo spheric breathers. Their common occurrence in pools and puddles has allowed many of us to view them hanging from the water's surface, or wriggling down into their watery home to escape perceived surface threats.Other commonly observed atmospheric breath ers are the larvae of dytiscid and hydrophilid beetles, which have functional spiracles at the end of their abdomen (Figs. 21.113 and 21.114). The majority of dipteran larvae also have functional spiracles located posteriorly but they are at the end of a tube called the respiratory siphon (e.g., Figs. 23.22, 23.25, 23.65, 23.70, 23.84, and 23.113). In Eristalis (Syrphidae), this siphon can extend to six times the body length (Fig. 23.84) giving rise to its common name of rattailed maggot. Larvae of species with siphons often are restricted to shallow seeps (e.g., ephydrid shore flies, ptychopterid phantom crane flies), to living near the surface in algal mats (e.g., dolichopodid longlegged flies, ptychopterids) or along pond and stream margins (e.g., tabanid horse flies), or to swimming short distances away from and back to the water sur face (e.g., ephydrids, culicids). Plant breathers (Table 4A) have spiracles modi fied to pierce submerged portions of aquatic plants and tap the plants' specialized air channels called aerenchyma (Houlihan 1969b, 1970). The sharp, barbed respiratory siphons of mosquitoes in the genera Mansonia!Coquillettidia and Taeniorhynchus (Culicidae; Fig. 4.4) can pierce the roots and stems of plants in open water, allowing larvae to remain sub merged until adult emergence and thus reduce their predation risk (Keilin 1944). Because these larvae have a thin cuticle and inhabit open water, the O2 obtained from plant stores can be supplemented by cutaneous respiration. Nonetheless, such larvae move and feed slowly and therefore appear unable to gain sufficient O2 for an active existence, perhaps as also reflected in their extended life cycle (Gillett 1972).

Transportable Air Stores Aquatic insects that rely solely on stationary O2 sources (e.g., the atmosphere)can leave those sources for only brief periods oftime or must remain relatively

inactive while separated from them. In contrast, aquatic insects that carry their own air supply can stay submerged longer and be more active. Many diving insects use air-sac flushing while surfacing. While airsacs in most terrestrial insects seldom compress more than 10-20%(Miller 1964a), Dytiscus sp.(Dytiscidae) and Eristalis sp.(Syrphidae) can flush about 65% of their entire system per compression (Krogh 1920, 1943), and Hydrocyrius giant water bugs(Belostomatidae) may completely collapse parts of their air-sac system (Miller 1961). High-volume ventilation con veys a substantial advantage to divers that only briefly contact the atmosphere. When a transportable air supply is exposed to the water, it can serve not only as an air reserve but also as a physical gill. Two types of physical gill are distinguished: (1)compressible physical gills, whereby diving time is extended but eventually the insect has to surface again to replenish the depleted O2, and (2) incompressible physical gills (sometimes also termed plastrons) that can last indefinitely. Compressible physical gills. When the air bubble is in contact with the water, O2 can diffuse from the water into the bubble and the physical gill is able to supply more O2 than it contained originally. At the start of the insect's dive, gases in the atmosphere, the bubble, and the water are in equilibrium (assuming that the water is fully saturated with O2). As the insect consumes O2 from its bubble, CO2 produced in cellu lar respiration, replaces the O2. However, because CO2 diffuses rapidly out of the bubble and into the

surrounding water (recall that the solubility of CO2 in water is very high), it contributes little to the gas com position of the temporary air store and therefore we can focus on O2 and N2. As a result of respiration, the partial pressure of O2 in the bubble decreases, and the partial pressure of N2 increases. This will result in an inward diffusion of O2 from the surrounding 02-rich water and an outward diffusion of N2, which will

eventually decrease the gas bubble size and force the animal to replenish the air store. However, because O2 diffuses into the bubble two to three times faster than

N2 diffuses out, this compressible physical gill can theoretically supply up to eight times more O2 than the original air store contained before the air store is depleted. However, animals tend to surface before the bubble is exhausted (Seymour and Matthews 2013). The length of time that a bubble can act as a gill decreases when the animal has a high rate of O2 con sumption (leading to a rapid depletion of O2 in the bubble) and increases with the bubble-water surface area, leading to a rapid inward diffusion of O2(Rahn and Paganelli 1968). Large insects that have high O2 demands must refill their air stores often because they

Chapter 4 Aquatic Insect Respiration

carry bubbles with relatively less surface exposed. For example, the air stores of adult giant water bugs (Belostomatidae), creeping water bugs Hydrous sp. (Naucoridae), and predaceous diving beetles(Dytiscidae) serve as effective physical gills only when water temperatures are low (e.g., winter) and O2 consump tion of the insects is minimal (Ege 1915; de Ruiter et al. 1952; Popham 1962). Other small diving insects that have relatively more bubble area exposed, such as corixid water boatmen, use their physical gill con tinuously (Ege 1915; Gittelman 1975). Consequently, these insects can swim some distance away from the surface. In fact, corixids have essentially become bot tom dwellers and thereby avoid competition for food at the water surface(Popham 1960). Incompressible physical gills. A number ofaquatic insects have a permanent gas film that acts as a phys ical gill (Table 4A). This permanent gas film is some times also called a plastron and functions in much the same way as the compressible physical gill described above. A key difference is that the gas film is held in place by tightly packed hydrofuge hairs(Fig. 4.5) or a cuticular meshwork (Fig. 4.6). This prevents the col lapse of the gas film and thus a decrease in volume. Consequently, O2 consumption results in a decrease in both partial pressure of O2 and also total pressure, as the volume of gas remains constant. Thus, the par tial pressure of N2 does not increase and there is no outward diffusion of N2 to the water so the bubble can be maintained indefinitely (Seymour and Mat thews 2013). O2 consumption, and thus metabolic rate, are determined by the rate of O2 diffusion through the fixed and limited surface area of the plastron. As a consequence, most insects with plastrons are slow-moving and are limited to habitats with high dissolved O2 and rapid flow as this helps in thinning the boundary layer. For example, Elmidae, which are commonly called riffle beetles because they occur in fast-flowing streams, use plastrons in their adult stage. Those insects that use plastrons in still water must either be good swimmers (e.g., hydrophilid beetles) or capable of crawling out of the water (e.g., curculionid weevils) to avoid low O2 conditions (Hinton 1976a). The ability of these still-water insects to detect and avoid low O2 is absolutely essential because O2 will diffuse away from the insect if the partial pressure of O? is higher in the plastron than in the surrounding water. Plastrons are quite variable in structure. Hydro fuge hair systems evolved in a wide variety of taxa, including lepidopterans (e.g., Acentropus caterpil lars), several kinds of beetles (e.g., the weevil Phytobius and the elmid Stenelmis), and the true bugs (e.g..

51

Aphelocheirus). The latter has one of the most effi cient plastrons known, consisting of a dense mat of

hydrofuge hairs(estimated to be 4.3 x 10^ hairs/mm^) that are bent at the tips (Fig. 4.5). This mat covers most of the ventral surface of the insect and most of

the dorsal surface as well (Hinton 1976a). The exten sive nature of its plastron allows this insect to spend its entire life cycle under water. In contrast, elmid beetles have a plastron formed by short, dense hairs located on the lateral and ventrolateral body surfaces and on the dorsum of the thorax. Their plastron is overlain by a second, temporary air store, some times referred to as a macroplastron, which is formed by longer, less dense hairs. The macroplastron air store is used when the O2 demands of the beetle are high (Thorpe 1950). Another way in which a plastron may be held in place is by means of a hydrofuge cuticular network. These structures are always associated with out growths of the area around the spiracular opening and often arise as columns from the body surface that divide at the top to form an open canopy (Fig. 4.6). An air film held beneath the canopy serves as an incompressible physical gill. These spiracular gills are found in pupae of many Coleoptera and Diptera, and in larvae of beetles in the families Torridincolidae,

Sphaeriidae, and Hydroscaphidae (Table 4A; Hinton 1968). Such insects often inhabit streams with highly fluctuating water levels. In these habitats, spiracular gills serve in both O2 acquisition when the insect is submerged and water retention when the insect is exposed to air.

Black flies (Simuliidae) are among the most com monly encountered insects that possess spiracular gills. Their pupae have a filamentous gill tuft on each anterolateral corner ofthe thorax(Fig. 26.24). The fact that these gill tufts are associated with spiral-shaped vortices in the flow that moves over them suggests their aquatic respiratory function (Eymann 1991). Black fly pupae live on stones and plants in running water, with the open end of their sac-like cocoon facing downstream. The spiracular gills project out ward, and the water circulating around them elicits a drop in pressure. As a result, the gill takes up O2 and collects air bubbles that are carried in the water,

which are constantly added to their plastron(Wichard et al. 2002). Spiracular gill dimensions in the pupae of both Simulium monticola and S. argyreatum dif fered in spring versus summer cohorts and both body size and sex also played a role in determining gill dimensions in these species (Kudela and Jedlicka 2002). Intraspecific variation in gill dimensions resulting from seasonality, body size, and sex likely can be found in other species as well.

52

Chapter 4 Aquatic Insect Respiration

Several factors reduce the effectiveness of physi cal gills. Deeper dives increase hydrostatic pressure, which causes gases to diffuse out of the bubble faster and may induce the collapse ofthe plastron. Lower O2 concentrations in the surrounding water decrease the diffusion gradient and therefore reduce the inward rate ofO,diffusion. Finally,increased water tempera ture increases O2 consumption by the insect, while also lowering the solubility of O2 in water; this combi nation results in a more rapid depletion ofO2from the surrounding water in the boundary layer. Insect size and shape also matters because small insects will have more efficient physical gills because of their relatively low rate of O2 consumption and the large surface area of the gas-water interface. Hutchinson (1981, 1993) has suggested that the relationship between gill effi ciency and water temperature may explain the pre dominance, within corixid water boatmen and other

aquatic groups using temporary air stores, whereby smaller species occur in warmer climates.

RESPIRATORY OPTIONS WITH A CLOSED TRACHEAL SYSTEM

Closed tracheal systems have no functional spira cles, so gas exchange must occur by diffusion through the cuticle. As aquatic insects typically do not have to deal with the stress of water shortage, some species may have thinner and more permeable cuticles(to both gases and water), and gas exchange may be further enhanced by a dense network of tracheoles just below the cuticle that provides a large exchange surface for gas diffusion. Because of either the surface-to-volume considerations mentioned previously or the presence of thicker, 02-impermeable cuticular surfaces, most insects with closed tracheal systems cannot fulfill their O2 requirements solely by diffusion through the gen eral body surface (i.e., cutaneous respiration). There fore,cutaneous respiration is commonly supplemented with O2 diffusion across highly tracheated, thin, body wall outgrowths called tracheal gills.

Cutaneous Respiration Because the amount of gas exchanged is propor tional to surface area, only aquatic insects that have a high surface-to-volume ratio can rely on cutaneous respiration alone (see Table 4A). This requires insects to be small, because small insects inherently have a high surface-to-volume ratio. Such a situation is demonstrated by the smallest of the aquatic Hemiptera, the non-North American Idiocoris and Paskia (Helotrephidae), which are the only known adult,

free-living, apneustic(no functional spiracles) insects

(Esaki and China 1927, cited by Hutchinson 1981). Alternatively, insects have to be flattened or elon gated. Most of the strictly cutaneously respiring spe cies are small, worm-shaped larvae such as chironomids (Fig. 23.42; Fox 1920; Parkinson and Ring 1982), ceratopogonids (Fig. 23.48; Ward 1991), chaoborids(Fig. 23.32), some tipulids (Pritchard and Stewart 1982), simuliids, and gill-less plecopterans and trichopterans. However,gas exchange in the early life stages of a large number of aquatic insects proba bly is through cutaneous respiration as well. Typi cally, these larvae are far less sclerotized than later stages. Young Trichoptera larvae respire exclusively through the cuticle, and gill filaments develop and become important only in later instars(Wiggins 1977) where their addition maintains a high surface-to-volume ratio. A similar situation is evident in some Plecoptera (Shepard and Stewart 1983). Even for those insects with tracheal gills, cutaneous respiration probably accounts for a significant but variable portion of total O2 intake. For example, Eriksen and Moeur(1990)demonstrated that even the may fly Siphlonurus occidentalis, with its proportionately very large tracheal gills, uses its abdominal surface for about 30% of its O2 intake. The damselfly Lestes disjunctus, whose gills are "...so large that one may suppose it would be an embarrassment to them if they grew any bigger..." (MacNeill 1960), normally uses cutaneous respiration to meet 70-80% of its needs at high O2 concentrations (Eriksen 1986), with gill surfaces providing the remaining 20-30%. The extent to which animals can meet their meta

bolic O2 requirements by cutaneous respiration likely decreases when their O2 demand increases (because of temperature or activity) and they will become more reliant on gill surfaces for O2 uptake.

TRACHEAL GILLS

Tracheal gills are present in the immature stages of at least some species in every aquatic insect order except the Hemiptera (see Table 4A). Tracheal gills have been shown to serve in ventilation, protection, hydraulic streamlining, swimming, and ion exchange. However, whether or not they are really used in respi ration has been the subject ofconsiderable debate. For example, removal of tracheal gills from larvae of the caddisfly Macronema resulted in no difference in O2 intake between normal and gill-less individuals(Mor gan and O'Neil 1931) In fact, gill-less larvae generally behaved normally and eventually pupated. However, such experiments were typically conducted under the favorable conditions of high dissolved O2 concentra tion and low temperature. When respiratory studies

Chapter 4 Aquatic Insect Respiration

were conducted under varying O2 and temperature conditions with the mayfly Cloeon dipterum, nymphs maintained a high metabolic rate down to 1 ppm O2 (equivalent to 1.8 mg/L) when gills were intact, but gill-less individuals experienced O2 stress below 3 ppm (equivalent to 5.3 mg/L)(Wingfield 1939). Gills ofthe damselfly Lestes disjunctus performed no respiratory role at 7°C and 5.5 ppm O2 (equivalent to 9.8 mg/L) but became increasingly important as temperature rose and O2 decreased until they accounted for up to 80% of total O2 intake (Eriksen 1986). Similarly, in the damselfly Coenagrion puella, gill autotomization (i.e., the spontaneous release of a body part) reduced their ability to tolerate extreme heat (Janssen et al. 2018). In general, tracheal gills apparently are not very important as O2 intake sites under high environ mental O2 conditions but become progressively more important as O2 concentration decreases or tempera ture increases.

Segmental pairs of lateral abdominal gills are found on at least some species in a number of orders containing aquatic insects (e.g., Megaloptera, Coleoptera, Zygoptera, Neuroptera) but they show the greatest structural diversity in the Ephemeroptera. Gills of mayflies vary from leaf-like (Fig. 13.66) to two-branched structures with single (e.g., Paraleptophlebia; Fig. 13.6) or multiple filaments (e.g., Habrophlebia; Fig. 13.28). Combinations of leaf-like and filamentous gills occur in a number of mayfly genera including Ephoron (Fig. 13.5) and Caenis (Fig. 13.16b). In Tricorythodes and Caenis, the first gill pair is enlarged to cover the posterior pairs (Fig. 13.15), presumably to shield them from being covered by fine sediments in the depositional habitats where nymphs are found. In a number of ephemeropterans that occur in rapidly flowing water (e.g., Iron), the gills overlap each other and are held against the substrate to provide a flattened shape to improve hydrodynam ics, thereby serving a dual function. The abdominal gills oftrichopteran larvae appear as scattered single(Fig. 19.92)or clustered(Fig. 19.63) filaments, which may or may not be branched. Their number and size tend to increase with increasing body

size. Wichard (1978) reported that gill number in the European species Molanna angustata was inversely related to the average environmental O2 concentra tion, a phenomenon suggested much earlier by Dodds and Hisaw (1924b). Although two families of damselfly nymphs (Odonata: Zygoptera) have paired lateral abdominal gills that are used in respiration (Norling 1982), this group is noted for its terminally placed abdominal gills. These caudal gills are usually leaf-like structures (Fig. 14.15), with two ofthem placed laterally and one

53

medially (Fig. 14.12). MacNeill(1960) described two gill types among Zygoptera: (1) the simplex type, which increases in size uniformly as the nymph grows and is typically found in the Lestidae and (2) the duplex gill, which consists of a thick proximal area and a thin distal zone that becomes disproportion ately larger at each molt. Duplex gills are typical of the Coenagrionidae. Internal placement of tracheal gills is found in dragonflies(Odonata: Anisoptera), where six longitu dinal rows of gills are located in an enlarged, anterior portion of the rectum called the branchial chamber. Nymphs respire by exchanging water through their anus, a behavior that ventilates their rectal gills. As a result, those species that burrow in mud must pro trude their anus above the sediment surface to prevent fouling of their branchial chamber and gills (Corbet et al. 1960). Although uncommon, gills(or what appear to be gills) can be found on the head and thorax of some Plecoptera, Diptera, and Lepidoptera. In stoneflies, care is required to determine whether or not these gills are truly respiratory organs (i.e., tracheated struc tures). The submental gills (Fig. 16.60) of Perlodidae, the so-called cervical gills(Fig. 16.25)of Nemouridae, and the coxal gills of Taeniopteryx (Taeniopterygidae)are nottracheated butrather are hemolymph-filled evaginations of the membranes between sclerites that function primarily as osmobranchiae (Shepard and Stewart 1983). Apparently, no tracheal gills occur on the head or cervical region of stoneflies, and the only thoracic gills in the Plecoptera that serve primarily in O2 intake are found in the Perlidae and Pteronarcyidae. Tozer's (1979) suggestion that the cervical gills retained in adult Zapada cinctipes (Nemouridae) probably function in respiration when adults enter the water to avoid subzero air temperatures appears to be incorrect.

RESPIRATORY PIGMENTS

One of the most vivid images in aquatic entomol ogy is the discovery of bright red "bloodworms" in dark, anoxic sediments of lakes and ponds. The red color of these chironomid larvae is caused by their respiratory pigment, which is hemoglobin. Although hemoglobin is characteristic of vertebrate blood, it occurs in some species ofmost animal phyla(Terwilliger 1980). Among insects, hemoglobin appears to be restricted to true bugs(e.g., Notonectidae)and dipterans (Chironomidae, and the terrestrial dipterans such as Drosophila and Gasterophilidae; the latter being bot flies that parasitize a variety of mammals; Table 4A) (Burmester 2015). Flowever, the respiratory pigment

54

Chapter 4 Aquatic Insect Respiration

hemocyanin, which is typically used by crustaceans, is also present in various insect orders including stoneflies, testifying to their phylogenetic links as the pan-

to maintain neutral buoyancy(Matthews and Seymour 2006). By doing so, these notonectids have exploited resources and avoided competition in the sparsely col

crustacea. These hemocyanins possibly also aid in O2

onized, midwater habitat in ponds.

storage and supply, and may work in tandem with the tracheal system (Burmester 2015). Insect hemoglobin differs from that ofvertebrates

by containing two (instead of four) heme groups. Chironomus possesses a high-affinity hemoglobin, which means that the pigment only releases O2 at low external O2 pressures, thus contributing to O2 uptake only in a I0W-O2 habitat such as that found in water and mud. By contrast, vertebrate and other insect hemoglobins tend to be low-affinity pigments. These hemoglobins release their O2 in the high O2conditions found in air, thus aiding insects having such a pigment in a terrestrial environment. In such low-affinity pig ments, the Bohr effect(whereby high CO2 concentra tions promote release of hemoglobin-bound O2) may confer a significant advantage for organisms that obtain O2 from a high-02 environment (i.e., lungs) and release it in a high-C02 environment(i.e., tissues). However, in environments where the O2 concentra tion is always low and CO2 is plentiful (e.g., lake muds), low-affinity hemoglobin would be inefficient. When chironomid larvae undulate their bodies in

their mud burrows to bring in water of higher O2 content (and hence higher partial pressure), their hemoglobin becomes 02-saturated. In between peri ods of undulation, hemoglobin gradually releases O2 to the tissues and the hemoglobin thus functions in O2 storage. If the hemoglobin's approximate 9-minute supply of O2 (as determined by Walshe [1950] for a species of chironomid) is less than is needed in the interval between undulations, the undulation fre

quency increases or anaerobic respiration becomes necessary. When undulations resume, hemoglobin enables a larva to recover rapidly from these anaero bic periods because the pigment facilitates O2 uptake (Walshe 1950). Two genera of Notonectidae {Anisops and Buenoa) have a low-affinity hemoglobin that performs a very different function than just described for midges. Miller (1964b, 1966) and Wells et al.(1981) observed that O2 released from hemoglobin con tained in certain richly tracheated abdominal cells markedly reduced the rate at which the temporary external air store depletes during diving. Because these hemoglobin-containing cells provide about 75% of the O2 used during a dive, the insect can carry a smaller air bubble, and does not have to fight the buoyancy associated with a larger air store. Also, gradual O2 release from the hemoglobin stabilizes the volume ofthe air bubble,helping the backswimmer

VENTILATION AND REGULATION

As mentioned previously, an organism has a much more difficult time obtaining sufficient O2from a dissolved source than it does from air. What makes

breathing under water a challenge is the much larger effort of ventilation required in water compared to air because of the higher density and viscosity of water. Greater efforts are required in water to reduce the thickness of the boundary layer and create a suffi ciently steep gradient in the partial pressure of O2 to facilitate O2 diffusion across respiratory surfaces. This challenge also means that the ability of organ isms to dynamically change and regulate O2 uptake (i.e., their regulatory ability) is inherently more lim ited in water than in air(Verberk and Atkinson 2013). One way to reduce the cost of breathing in water is the adoption of cutaneous respiration, because this pro cess requires no ventilatory effort. In actuality, virtu ally all aquatic ectotherms use cutaneous respiration to augment their O2 uptake. However, many insects also use O2 ventilation, which involves the flow of air

or water by active or passive means over respiratory surfaces or through part of the tracheal system, to thin the boundary layer. Ventilation currents may result from abdominal contractions, body undula tions, gill beating, swimming through the water, movement to a more favorable microhabitat, utiliza tion of stream flow, or a combination of these mech

anisms (Table 4B). Ventilation by an insect generally pushes water posteriorly over the gills and dorsal body surface. Eastham (1934, 1936, 1937, 1939) demonstrated that mayflies beat their gills to create respiratory currents, and the frequency of gill beat increases as O, concentration decreases (Eriksen 1963a; Eriksen and Moeur 1990). Riley(1879, as cited by Tracy and Hazelwood 1983) alludes to a similar behavior for the hellgrammite Corydalus(Megaloptera). Trichopterans, chironomids, ephemeropterans, and aquatic lepidopterans all use body undulations to pump water through their cases or tubes and burrows (Welch and Sehon 1928; Walshe 1950; Feldmeth 1970). For some Trichoptera, efficient ventilation apparently depends on the presence of a case that restricts and directs water flow (Williams et al. 1987), as some larvae removed from their cases ultimately die even though they continue to undulate. Rather than relying on external tubes or cases, dragonfly nymphs (Anisoptera) possess an especially effective

Chapter 4 Aquatic Insect Respiration

llil Table 4B

5!

Ventilation methods for aquatic insects utilizing dissolved oxygen.

System

Ventilation Method

References

Taxon

Ventilated Cutaneous

Undulation

Chironomidae

Trichoptera (gills lacking) Lepidoptera (gills lacking) Swimming Natural water flow

Trachea! Gills

Beating gills

Undulation

^

2911

Chaoboridae

*

Chironomidae

*

Trichoptera (caseless, gills lacking) Plecoptera (gills lacking)

1247, 1868 1247, 1868

Simuliidae

5860

Ephemeroptera

1346, 1347, 1348, 1349, 1456, 1462, 5743

Psephenidae Corydalidae Gyrinidae

4774

Trichoptera

1524, 2962, 4043, 4044, 4055, 4046, 5596

Lepidoptera

5526

3159 2973

Chironomidae

5421

Leg contractions that move body (push-ups)

Plecoptera

Rectal pump

Anisoptera Heptageniidae Plecoptera Zygoptera Trichoptera (caseless) Trichoptera (with case) Biephariceridae

343, 1779, 2639, 2750, 2751, 3383, 3361 1459, 3662 2641, 3490, 3491

1

Lestidae Natural water flow

Temporary and

2561,3144, 5421

Leg movements

Permanent Air Stores

Swimming

96 1247 5918 96

1524, 4045, 4046 *

Notonectidae Naucoridae Corixidae

1195

All taxa that swim with exposed

1195

1195

1195, 4122

air bubbles Natural water flow

Simuliidae (pupae) Dryopidae (adult) Lepidoptera (larvae and pupae)

1495, 2282 *

375, 376

*Eriksen, C, H. Personal observation.

ventilation mechanism for their gills, which are located in a blind sac off the rectum. Contraction, mainly of dorsoventral abdominal muscles, increases pressure in the branchial chamber and forces water

out the anus. When muscular relaxation occurs, neg ative pressure in the chamber allows 02-rich water to return. Ventilations increase in frequency as O2 decreases and temperature increases (Mantula 1911; Mill and Hughes 1966; Cofrancesco and Howell 1982). In contrast, some organisms, such as stoneflies and lestid damselflies, perform "push-up" ventilatory

movements(Knight and Gaufm 1963; Eriksen 1984).

Although helpful, these movements are initiated only during periods of respiratory stress because they are inefficient, and merely stir up the surrounding water rather than force a directed convection of oxygen ated water over the gills as is the case with tube ven tilation. Some insects are unable to accomplish any self-generated ventilation because they have adapted so completely to the ventilation provided by natural water movement(Table 4B). In the absence of water flow, these insects cannot acquire sufficient O2 and soon die (Jaag and Ambiihl 1964). Insects can enhance respiratory processes in other behavioral ways as well. Many insects possess regions

56

Chapter 4 Aquatic Insect Respiration

of trachea! expansion and compression in the head and thorax. These movements are independent of hemolymph circulation and other body movements, and likely serve to aid internal convection of air in the trachea! system, somewhat analogous to the inflation and deflation of vertebrate lungs. This phenomenon has been observed in numerous terrestrial and aquatic insect groups, including Hemiptera and Odonata (Westneat et al. 2003). Apodaca and Chapman (2004) demonstrated another behavioral adaptation, whereby the African damselfly Pwischnura subfurcatum migrates to the water surface to make contact with atmospheric air under hypoxic conditions. Gill autotomization is common in this species, and both gilled and gill-less individuals performed surface migrations under hypoxic conditions. Gill-less individuals also were found to rely more heavily on manipulation of wing sheaths (evidenced by lifting and spreading behavior) than gilled individuals. A distinction is made between an 02-regulator (or respiratory regulator) and 02-conformer (or respiratory conformer) when referring to how an organism responds to O2 changes in the surrounding environment. Some species maintain relatively stable rates of O2 intake across wide gradients of environ mental O2availability; these species are "oxyregulators." This may occur if O2 availability far exceeds demand, so changing O2 conditions do not affect O2 intake. As O2 drops further, they may,for example, compensate

by increasing ventilation behavior. If O2 availability continues to drop, a threshold called the critical point (or Pcrit) is reached where the supply of O2 is not suf ficient for the organism to maintain the same rate of O2 intake. Here, the species shifts to "oxyconforming" and likely signals the onset of anaerobic metab olism, which is not sustainable for most species. In contrast to regulators, respiratory conformers are unable to create significant respiratory currents or compensate otherwise, and as a result, their O2 intake is proportional to the O2 availability in their micro-habitat. As O2 availability in the water declines, so does their O2 uptake. Examples of oxyconformers are certain species of stoneflies and caddisflies, where O2 intake changes with O2 availability (Kapoor and Griffiths 1975; Rotvit and Jacobsen 2013). Aquatic insects that use atmospheric air are typically respira tory regulators as they can readily compensate by increasing the length of time the spiracles are open, the number of ventilation movements, or the fre

quency of surfacing. Some rheophilic species that use natural water current to ventilate their respiratory surfaces are more likely to be oxyconformers (e.g., rhyacophilid caddis flies, blepharicerid midges) and Ambuhl (1959) has

demonstrated that in these species respiration also increases with current speed of water, at least until some plateau(regulation)is reached. Feldmeth(1970) similarly found that current speed is related to the intensity of respiration in the caddisflies Pycnopsyche lepida and P. guttifer, but locomotor behavior, as influenced by current velocity, is most important in setting the respiratory rate. Such a response makes sense because many rheophilic insects cannot venti late for themselves and, therefore, they let natural water flow bathe respiratory surfaces for them. These cases represent two extremes of a contin uum and studies of aquatic insect respiration have revealed a variety of abilities ranging from absolute conformity to strict regulation. Mueller and Seymour (2011) developed an index of regulatory capacity to assess the capacity of animals to maintain function relative to a complete oxyconformer. Furthermore, Eriksen (1963a)and Nagell(1973) demonstrated that some Ephemeroptera and Plecoptera species appear to be either respiratory regulators or conformers depending on experimental conditions. Similarly, Golubkov et al.(1992) note that some species, which might otherwise be thought of as conformers, demon strate a constant level of respiration in rapidly flowing water. In contrast, a variety of experimental condi tions did not seem to change a lestid damselfly from being intermediate between regulation and confor mity (Eriksen 1986). Clearly, the ability of insects in maintaining constant rates of O2 intake to achieve some balance between the risks of asphyxiation and O2 toxicity will differ from species to species, and depend on the environmental context.

Osmoregulation and Respiration Osmoregulation is the process of regulating and maintaining the appropriate balance of water and ions in body fluids such as blood or hemolymph—a critical function in all aquatic organisms (Kirschner 1991). Respiratory and osmoregulatory processes may often be linked, at least in insects that rely on dissolved O2. For example, surfaces that are in direct contact with water may be used for both respiration and osmoregulation. The osmolarity of body fluids of aquatic insects is generally 200-400 mosm/L(Komnick 1977), and is considerably higher than that of the surrounding water, which typically ranges from 1 to 2 mosm/L in many freshwater systems. Consequently, water has a natural tendency to penetrate the integu ment and into the insect body,just as there is a ten dency for ions to diffuse out of the animal. Just as insects had to evolve respiratory mechanisms to live permanently in freshwater environments, they also

Chapter 4 Aquatic Insect Respiration

had to devise strategies for overcoming this osmotic gradient(Wichard et al. 2002). Mechanisms have evolved to promote the uptake of O2 while, at the same time, prevent osmo sis. For example, an air bubble or film of air that encloses the body more completely will enhance res piration because of the larger respiratory surface. This air also keeps water away from the body reduc

ing osmotic problems. Another example of a respira tory strategy that reduces ionic gradients between the organism and the surrounding water can be seen in the pupal cocoon of aquatic hymenopterans that parasitize larval trichopterans. The air-filled pupal cocoon pulls in dissolved O2 from the surrounding water because of the difference in O2 partial pres

sure, thus acting as a physical gill. At the same time, the cocoon protects the pupa from direct contact with the surrounding water—again reducing osmotic problems. The few aquatic beetles that pupate under water use a similar mechanism to effectively live in what is really an underwater, terrestrial environment (Wichard et al. 2002). Insects also use many other strategies to enhance respiration that have costs associated with increasing osmoregulatory demands. For example, gill surfaces are generally more water-permeable than other integ ument, and insects with larger gills tend to be more water-permeable than small-gilled or gill-less insects (Buchwalter et al. 2002). These animals must excrete excess water through the production of hypotonic urine(Chapman 1982). The loss ofions in urine through solvent drag, and the diffusive loss of ions through paracellular channels on gills and other epithelial sur faces, is unavoidable. It is therefore not surprising

that specialized ion-absorbing cells called chloride cells are often found on gill surfaces (Komnick 1977). These mitochondria-rich cells sequester ions from the water column (usually against concentration gradi ents) to help maintain salt/water balance. Trace met als can be accumulated by these cells, and species with larger numbers of chloride cells also appear

57

jet propulsion. However, in damselflies, these func tions appear to be more separated, with absorption of O2 typically associated with the body surface, includ ing the three caudal tracheal gills, and osmoregula tion typically associated with rectal ventilation whereby dissolved ions are absorbed from water that is pumped into the anus. However, Miller (1994) observed increased abdominal pumping in damsel flies under hypoxic conditions, suggesting that rectal surfaces may also be important in gas exchange. Another example of the linkage between respira tion and osmoregulation can be found when O2 sup ply in the environment is limited. Under hypoxic or anoxic conditions, when organisms shift from aerobic to anaerobic respiration, they create metabolites such as lactate and succinate, which can produce metabolic acidosis or alter the composition of the hemolymph (Scholz and Zerbst-Boroffka 1998). For example, the nonbiting midgelarvae Chironotnusthummi(Redecker and Zebe 1988)and Chironotnus gr. plumosus(Scholz and Zerbst-Boroffka 1998) often spend considerable time in hypoxic conditions and are able to ferment ethanol, which is readily excreted and does not accu mulate in the hemolymph.The mosquito Culexpipiens, on the other hand, is intolerant of hypoxia and builds up lactate in the hemolymph and this accumulation is accompanied by a decrease in hemolymph chloride concentration (Redecker and Zebe 1988).

RESPIRATION AND TOXICANTS Human activities often lead to the introduction of

toxic chemicals to aquatic systems. These contaminants can range from inorganics such as trace metals and other salts, to pesticides and industrial organic com pounds. An emerging area of concern is the introduc tion of pharmaceuticals into receiving waters from municipalities. Toxic chemicals can influence the respi ration ofaquatic insects, and conversely, the respiratory surfaces of aquatic insects can influence exposure to toxic chemicals.

to accumulate dissolved Cd and Zn at faster rates

(Buchwalter and Luoma 2005). Metals are also known to accumulate on other osmoregulatory structures such as anal papillae(Vuori 1994). Interestingly, there

Effects of Toxicants on Respiration Rates

is not always a correlation between the numbers of chloride cells and gill surface areas, because signifi cant osmoregulatory processes occur in the gastroin

increase or decrease respiration rates. For example, exposure to sublethal concentrations of copper (Kapoor 1976) or the organophosphate pesticide Dibrom (Maki et al. 1973) increased the O2 con sumption rates of plecopterans and megalopterans

testinal system.

Among the Odonata, dragonflies differ from damselflies in terms of their osmoregulation. In drag onflies, the water that is taken into the rectal gill chambers functions in both respiration and osmoreg ulation capacities,and is also used in locomotion through

Sublethal concentrations of toxic chemicals can

and reduced their tolerance to low dissolved O2 con

centrations. Similarly, the haloform byproducts of water chlorination consistently increased the respi ration rate of dragonfly nymphs in laboratory studies

58

Chapter 4 Aquatic Insect Respiration

(Correa et al. 1985a; Calabrese et al. 1987; Dominguez et al. 1988). In contrast, however, O2 con sumption by chironomid larvae declined after exposure to naphthalene (a highly toxic polycyclic

aromatic hydro-carbon), and hemoglobin-lacking Tanytarsus dissimilis larvae were more sensitive to

naphthalene than hemoglobin-possessing Chironomus attenuatus larvae (Darville and Wilhm 1984). Respiration can also be affected when metal ions, such as iron, precipitate on gill surfaces under acidic conditions or displace functional cations from the active sites of enzymes and result in respiratory failure (Gerhardt 1992). The interactive effects of environmental stressors

on respiration have received some attention, but

results of studies are not consistent. For example, when exposed to low pH and high aluminum concen tration, a variety of aquatic insects tested displayed reduced respiration rates (Rockwood et al. 1990), increased respiration (Correa et al. 1985b; Herrmann and Andersson 1986), or showed no respiration-rate effect (Correa et al. 1986). Toxicant interactions undoubtedly are common occurrences in a number of aquatic ecosystems and deserve more study with con trolled experiments. In toxicological studies, Doherty and Hummon (1980) provide a note of caution that although responses based on respirometry may indi cate physiological distress, they fail to identify the specific toxic action. Nonetheless, these studies are important in determining the levels of contamination that cause changes in insect respiration. Given the large number of contaminants of concern, and the large number of aquatic insect species, remarkably little work has been done thus far on this subject.

Respiratory Characteristics and Contaminant Accumulation

In some cases, respiratory surfaces are involved in the absorption of toxic chemicals. Tracheal gills appear to be particularly important in this regard because cell surfaces that are directly exposed to the water column are more permeable than heavily sclerotized or waxy cuticle and may be rich in chloride cells. For example, mercury can enter Hexagenia rigida (mayfly) nymphs by direct absorption across gill lamellae in amounts that exceed those obtained from their diet(Saouter et al. 1991). Once in the may fly, this metal is dispersed through the body by the tracheal system and the hemolymph. Air-breathing insects accumulated the organophosphate pesticide chlorpyrifos much more slowly than dissolved 02-breathing insects (Buchwalter et al. 2002, 2003) and growth was much less impaired by chlorpyrifos in

damselfly nymphs that had autotomized their gills and consequently had a reduced respiratory surface for absorption of toxicants(Janssen et al. 2018). Fur thermore, species with larger gills had faster accumu lation rates than would be predicted based on body size alone. Because respiratory surfaces are permeable to water, the water permeability of aquatic insect spe cies can be used as a surrogate for gill surface area (which can be extremely difficult to measure accu rately)(Buchwalter et al. 2002). The pH of the medium is another environmental

stressor that can differentially affect species based on respiratory and osmoregulatory characteristics. For example, the air-breathing beetle Dytiscus verticalis was more tolerant of low pH compared to the dis solved 02-breathing dragonfly Anax junius (Frisbie and Dunson 1988). The ultrastructure of gill tissue in Pteronarcys dorsata (stonefly) nymphs can be altered by environmentally extreme acidic(pH 10) conditions, thereby resulting in the loss of sodium ions and eventual death (Lechleitner et al. 1985). Chloride cell structure and O2 consump tion were also affected by alkaline pH in the mayfly Isonychia bicolor (Peters et al. 1985). For a more in-depth review of hemolymph acid-base regulation, see Cooper (1994). Relatively few studies of toxicant effects on

aquatic insect respiration have been conducted in nat ural aquatic ecosystems. Herrmann and Andersson

(1986) found that different species of mayflies domi nated in natural streams according to the stream pH, a situation that corroborated their laboratory find ings that the respiratory stress caused by low pH dif fered across mayfly species. In streams receiving chlorinated effluents, perlid stoneflies and hydropsychid caddisflies had atrophied or deformed gills in 62-100% ofthe specimens found at the impacted sites. In contrast, nonpolluted upstream populations had normal gill structure (Simpson 1980; Camargo 1991). Aquatic insects such as hemipterans with physical gills (transportable air stores) are probably less sus ceptible to damage by chlorine, as their mode of res piration allows for a more impermeable exoskeleton, perhaps explaining why hemipterans frequently are found occupying swimming pools! Clearly, respiratory surfaces, especially tracheal gills, are some of the most sensitive to environmental contaminants. Toxicants can result in physical dam age to the respiratory structures, and may also pass through respiratory surfaces to affect other organ systems. However, whatever their site of effect, toxi

cants can lead to respiratory stress, which in turn can lead to death. The coupling oflaboratory experiments with in situ studies is crucial to understanding both

Chapter 4 Aquatic Insect Respiration

the mechanistic effects and ecological consequences of toxicants on aquatic insect respiration.

Limits of Respiratory Function: Environmental Hypoxia Maintenance ofan adequate supply of O2 to meet the demand of their tissues can be challenging for aquatic insects when faced with environmental hypoxia (i.e., a reduced supply of O2). Unlike most terrestrial habitats, environmental hypoxia is quite common in aquatic habitats. Inherent in the low capacitance of water for O2, processes that consume or generate O2 quickly result in large changes in aquatic partial pressure of O2. At the same time, these pressure differences take much longer to equilibrate with the atmosphere because of the much slower rates of diffusion. As a result, hypoxic events tend to grow more severe (i) during the night when respiration is not counteracted by photosynthesis, (ii) in standing waters where convection and reaeration is limited,

and (iii) in microhabitats that are rich in organic mat ter, where water flow is impeded, or both. Examples of such microhabitats include leaf litter packs in streams, benthic habitats in standing water, and the interior of dense macrophyte stands. During daytime, the reverse happens and insects may have to deal with hyperoxia (i.e., an overabun dance of O2). Fluctuations in O2 are especially strong in small, nutrient-rich water bodies with a high pri mary productivity. As levels of dissolved O2 vary

59

was the temperature dependence of lethal concentra tions. In warmer water, higher O2 concentrations were required to ensure survival. One can assess the capacity of aquatic insects to supply adequate O2 to meet tissue demand by measur ing their ability to maintain respiration rates in the face of declining supply. This metric delineates respiration regulators from respiration conformers. Manipulation of the O2 demand is another way to assess the capacity of organisms to supply sufficient O2, frequently done by comparing the rate of O2 uptake of animals that are active with their rate when at rest, or by comparing rates of fed and postabsorptive animals. The difference between these rates of O2 intake gives the aerobic scope of an animal (or the factorial aerobic scope, when rates are divided rather than subtracted), which expresses the animals"excess" capacity for O2 delivery under the conditions mea sured. These two concepts of aerobic scope and respi ratory regulation are likely related, but tests of this idea are scarce, probably because it is difficult to force insects to exercise and concurrently obtain aerobic scope measurements. Kim et al. (2017) used a meta bolic de-coupler to stimulate maximum O2 consump tion for the assessment ofaerobic scope, but it remains unclear whether these pharmacologically altered max imum consumption rates are similar to ecologically relevant rates associated with exercise. Still, both con

cepts predict anaerobic metabolism results from O2 limitation (i.e., when aerobic scope approaches zero or Pcrit is reached).

between microhabitats and with time, relevant mea

sures of the oxygenation of the habitat to explain differences in assemblages of aquatic invertebrates are frequently lacking. This is one reason why the biochemical oxygen demand (BOD) which expresses how much O2 is consumed via microbial respiration, is regularly monitored. This measure likely better approximates the O2 conditions that insects experi ence in their microhabitat compared to point mea surements oflevels ofdissolved O2in the water column (Verberk et al. 2016a). Moreover, documented effects

of flow, nutrients, effluent discharge, altitude, and temperature on aquatic insects are readily explained from an O2 perspective. These various other factors are strongly correlated with dissolved O2 or have strong repercussions for the balance between O2 demand and O2 supply (e.g., Lowell and Gulp 1999; Jacobsen et al. 2003; Verdonschot et al. 2015; Pardo and Garcia 2016; Verberk et al. 2016a). For example, in a laboratory experiment, Nebeker (1972) used a bioassay approach to show that aquatic insect species vary broadly in their dissolved O2 median lethal con centrations. One important finding from that research

Limits of Respiratory Function: Temperature Marine invertebrate studies have suggested that the thermal limits of many species coincide with the loss of aerobic scope and a shift from aerobic to anaer obic pathways. Energy deficits arise at the colder limit of the thermal performance window because mitochondrial function is impaired. At the hotter end ofthe thermal performance window, the organism's meta bolic needs can outpace the capacity to take up and transport O2 to the required tissues (Portner 2002). Research on the thermal dependency of respiratory function has given rise to the hypothesis that O2 limita tion sets thermal tolerance limits, because of the mis

match between O2 supply capacity and O2 demand (Portner 2001). The (exponential) increase in tissue O2 demand with temperature is probably more import ant for the mismatch to manifest itself, as the availabil

ity of dissolved O2 is much less temperature sensitive (Verberk et al. 2011). Still, the efficiency of ventilation

likely decreases with increasing temperature, mainly

60

Chapter 4 Aquatic Insect Respiration

because warmer water holds less dissolved O2,requiring more water to be displaced to reach similar rates of O2 uptake. For example,Philipson and Moorhouse(1976) found that for the caddisfly PolycentropusJlavomaculatus, O2 uptake efficiency (expressed as O2 uptake per body undulation) dropped by more than 50% when temperatures increased from 10 to 25°C. The O2 limita tion hypothesis has been criticized and appears to be better supported in aquatic arthropods, whereas evidence for terrestrial arthropods is more limited (Verberk etal. 2016b). Others(Kim etal. 2017; Sweeney et al. 2018) suggested that O2 limitation hypothesis might be more appropriate for acute thermal limits than chronic thermal limits in baetid mayflies. One approach to test this hypothesis of O2 limita

performance such as exercise ability, growth, and reproduction may be impaired and that these factors may be more sensitive to the interactive effects of hypoxia and warming. In contrast, Kim et al.(2017) studied nymphs of the mayfly Neocloeon triangulifer held at chronically lethal temperatures under normal O2 conditions. Aer obic scope was not reduced at temperatures associ ated with chronic thermal lethality, and genes responsive to hypoxia were not stimulated by chron ically lethal temperatures. Thus, no evidence of O2 limitation was found at temperatures that are chron ically lethal, although evidence for O2 limitation was found at acute, intense, heat-stress temperatures unlikely to be encountered in nature. The discrepancy

tion and thermal tolerance is to assess whether ther

between acute and chronic heat stress could be resolved

mal limits depend on ambient O2 conditions. For example, Verberk and Bilton(2013)assessed the acute thermal limits of eight species of aquatic insects from four different insect orders by rapidly ramping up the water temperature (0.25°C/min) and noting the tem perature at which animals became moribund. They then similarly assessed thermal limits in hypoxic water to test the prediction that thermal limits should be reduced in hypoxic waters if the animals were O2 lim ited. Almost all animals displayed reduced thermal tolerance under hypoxia. Moreover, differences in the sensitivity of species to hypoxia could be related to differences in their mode of gas exchange (Fig. 4.7), with animals relying on gas exchange across their cuticle or via a plastron being consistently more vul nerable to the synergistic effects of warming and hypoxia, relative to those that actively ventilated their (enclosed) gills or to air breathers. Potential issues with the experimental approach

by assuming that when O2 conditions are normal(normoxia), O2 limitation can occur via increased meta

described above are that the thermal limits obtained

during these acute thermal exposure trials and the thermal ramping rates used are unlikely to reflect thermal regimes that aquatic insects experience in nature. In natural environments, animals are com monly exposed to less extreme temperatures and

slower rates of thermal change, although they are exposed for much longer periods. At present it is unclear how O2 modulates tolerance to heat stress of lower intensity and greater duration. Verberk et al. (2016a)compared the effects of heat and hypoxia for aquatic nymphs of two species of mayfly between an experimental setting and the field situation. They ana lyzed a large data set with tandem measurements of water temperature, O2 levels, and the presence of mayflies across more than 2,600 field locations. Their study indicated stronger interactive effects of hypoxia and warming in the field compared to a lab setting. It is possible that over longer time scales, sublethal

bolic demand

under acute and intense thermal

challenge, but energetic issues become more import ant under chronic heat stress. For example, chronic heat stress under normoxia may impair fitness through the increased costs of maintenance and ener getic deficits(Chou et al. 2018). However, in nature, environmental hypoxia can occur episodically in the field and aquatic insects may have 02-sensitive peri ods,such as when molting(Camp et al. 2015), that are exacerbated by warmer temperatures. More research on the interactive effects of warming and hypoxia on longer timescales in a range of species with different modes of respiration is needed to disentangle cause and effect.

CONCLUDING REMARKS

This chapter has highlighted the fascinating diversity in morphological, behavioral, structural, and physiological adaptations that insects possess that help them meet their O2 needs under the great range ofconditions that aquatic insects inhabit. Water temperature, oxygenation, and presence of pollutants all interact with the respiratory biology of aquatic insects. Interpretations of the dynamics of aquatic insect populations, assemblages, and communities would benefit from a better understanding of how aquatic organisms function at the physiological level. Understanding the differential responses of taxa to environmental change requires that we more thor oughly consider the advantages, limitations, and tradeoffs for dealing with O2 availability in the aquatic environment. A thorough understanding ofthese pro cesses occurring throughout the long evolution of these ancient and diverse lineages of aquatic insects is thereby fundamental to aquatic entomology.

Chapter 4 Aquatic Insect Respiration

llyocorus cimicoides

A 42

Agabus bipustulatus

(air breather)

B 42-1

(air breather)

33

33

Limnius volckmari

(plastron breather) 24-

24

Aphelocheirus aestivalis (plastron breather)

u

X

15, n

1

15'

r

D 42-1

C 42n

1

r

-|

1

Cordulegaster boltonii (enclosed gill surface) •*

Ecdyonurus insignis (beating gills) 33-

33

24

24-

Calopteryx virgo (outer gill lamellae)

Yf^hitrogena semicolorata (immovable gills) 15

-*i

1

-|

11

\

\

1 1

1520

20

Oxygen (kPa)

Figure 4.7 Thermal tolerance limits of species pairs belonging to four insect orders: beetles (a), bugs (b), mayflies (c), and odonates (d). Hypoxia generally decreases thermal limits tolerance in each Insect order. Although species from different orders have different capacities for O2 uptake, within each order, species reflected pairwise contrasts in respiratory regulation, delineating species with poor respiratory regulation (shown in black)from those with good respiratory regulation (shown in blue). The beetles Agabus bipustulatus (Linnaeus 1767) and Limnius volckmari(Panzer 1793) are surface exchanging and plastron breathing adults, respectively, as are the bugs liyocoris cimicoides (Linnaeus 1758) and Aphelocheirus aestivalis (Fabricius 1794). The mayfly and dragonfly nymphs all have gas exchange across their cuticle and tracheal gills. The mayfly species differ in their ability to move their gills and hence their degree of respiratory regulation; Ecdyonurus insignis (Eaton 1870) is able to beat its giils; Rhithrogena semicoiorata (Curtis 1834) is not. Within the odonates, the dragonfly Cordulegaster boitonii(Donovan 1807) has the rectum modified into a heavily tracheated branchial chamber whose surface acts as a gill. Being able to force water across the respiratory surface through abdominal movement provides greater respiratory regulation relative to the damselfly Calopteryx virgo (Linnaeus 1758), which has instead external gill lamellae. © Kendall Hunt Publishing Company.

61

62

Chapter 4 Aquatic Insect Respiration

Table 4C Demonstrations of respiratory processes {superscripts refer to Section C. Useful Equipment). Table 4B contains relevant literature.

Closed Respiratory System {larvae only) 1.

Ventilation Methods and Behavior

a. Beating gills''^'® (e.g., burrowing, climbing, sprawling Ephemeroptera, Corydalidae) b. Push-ups''^ (e.g., Plecoptera, Lestldae) c. Undulatlon^'^'^ (e.g., Trichoptera, Lepidoptera, Chlronomldae) d. Muscular rectal pump''® (Anisoptera) e. Swimming'(e.g., Chaoborldae, Chlronomldae) f. None (other than possible position change)(e.g., Blephariceridae, Simuliidae, fast-water Ephemeroptera, Plecoptera) 2. Respiratory Currents Produced by Insect (Section A.l .a-d)® 3. Micro-areas from which Respiratory Water Obtained (Section A.I .a-d)® 4. Environmental Effects on Ventilation Frequency and Volume of Respiratory Flow (Section A.I .a-d)

Vary: dissolved O2 concentration'® current velocity

water temperature" Open Respiratory System (larvae and adults) 1. Ventilation Methods and Behavior

a. Leg movements"(e.g., Corlxidae, Naucoridae, Notonectldae) b. Swimming® (any species with exposed air store) 2. Environmental Effects on Diving Time (any species with temporary air store) Vary: dissolved Oj concentration®''" dissolved CO2 concentration®''" temperature®'" 3. Diving Stimulus(any species with temporary air store) Provide: air atmosphere

O2 atmosphere®''" CO2 atmosphere®''" Nj atmosphere® '" 4. Need for Surface Tension to Establish Atmospheric Connection®'"(e.g., Culicldae, Tipulldae, Syrphldae, Notonectidae, Dytiscidae) 5. Plastron (any species using plastron respiration only)

Vary: dissolved O2 concentration'''" current velocity''^ water temperature''" Useful Equipment

1. Narrow (e.g., < 3 cm) plexiglass observation aquarium.

2. U-shaped glass burrows simulating natural dimensions. Portion restricted with coarse mesh screen for containing animal but allowing current flow (Walshe 1950; Eriksen 1963a). 3. Artificial Trichoptera case: glass or plastic tubing approximating case interior diameter and length with one end restricted to a 1-mm central pore (Feldmeth 1970). 4. Vertical, clear "diving tube," 2-3 cm by about 30 cm. Vertical strip of plastic screening near surface simulating vegetation. Horizontal screening just below water level. No bottom substrate. 5. Vertical, clear "diving tube," 2-3 cm by 100-200 cm. Horizontal screening on bottom as substrate. 6. Plastic window screen cut to appropriate shapes.

(continued)

Chapter 4 Aquatic Insect Respiration

Table 4C

63

Continued

7. Water current generation:

• gravitational, from reservoir via appropriate tubing with flow control valves • air hose pump

• magnetic stirrers. Note: these create centrifugal currents; however an organism can be restricted to one area and experience essentially longitudinal current flows (e.g., Philipson 1954; Morris 1963). • water current respirometer (e.g., Eriksen and Feldmeth 1967). 8. Carmine or carbon-black suspension introduced where desired with narrow aperture eyedropper. Observe particle movement. 9. Detergent or thin oil added to water surface with eyedropper.

10. Gas concentrations: control concentration of dissolved gases in reservoir with gas mixing valves or a combination of compressed air, O2, Nj, or COj. Monitor with O2 electrode if available. 11. Temperature: many heating/cooling devices may be used to adjust reservoir temperature, or use temperature controlled environmental rooms.

APPENDIX 1: DEMONSTRATIONS OF RESPIRATORY PROCESSES

The respiratory structures and processes that have been described in this chapter are best under stood when they are seen. As a means to that end, simple experiments are summarized in Table 4C that demonstrate the structural and behavioral abil

ities, and also the limitations, of aquatic insects subjected to varying environmental conditions. When keeping aquatic insects in the laboratory, or

conducting experiments with them, always avoid stressing the organisms unless it is part of the exper imental design. Likewise, always provide suitable substrate (pebbles, plastic mesh, glass burrows,etc.) and normal environmental O2, temperature, and water flow (see Rearing Methods, Chapter 3) when

holding and using the insects in experiments and as controls. Detailed explanations of additional experiments can be found in Kalmus (1963) and Cummins et al.(1965).

-

v; j

'■Hi> ^v»«'

HABITAT, LIFE HISTORY, SECONDARY PRODUCTION, AND BEHAVIORAL ADAPTATIONS OF

AQUATIC INSECTS Alexander D. Huryn University of Alabama, Tuscaloosa

INTRODUCTION

The occurrence of insects in virtually all freshwa ter communities, and their position as the dominant class of macroinvertebrates in most of these commu

nities, provide evidence of their extraordinary evolu tionary success. In this chapter, we use the insect life history as a framework for describing different mor phological adaptations, behaviors, and life-history strategies that have enabled their unparalleled radia tion into a diversity of aquatic habitats. HISTORICAL SOURCES OF INFORMATION

There are many excellent sources of general infor mation about the natural history of aquatic insects. The first book-length treatment—Natural History of Aquatic Insects—was published over a century ago (Miall 1895). Another early work that focused on the life histories of aquatic insects, rather than their tax onomy per se, is Biologie der Susswasserinsekten (Wesenberg-Lund 1943). Although not strictly a text on insects, H.B.N. Hynes' (1970a) classic. The Ecol ogy of Running Waters, contains a thorough review of information on the life cycles and adaptations of stream insects through the late 1960s. More recent sources include Aquatic Entomology (McCafferty 1981), The Ecology of Aquatic Insects (Resh and Rosenberg 1984), Aquatic Insect Ecology (Ward 1992), Aquatic Insects (Williams and Feltmate 1992), New Zealand Stream Invertebrates: Ecology and Implications for Management (Collier and Winterbourn 2000), Biological Atlas of Aquatic Insects (Wichard et al. 2002), Aquatic Insects: Challenges to

J Bruce Wallace

University of Georgia, Athens

Populations (Lancaster and Briers 2008), and Aquatic Entomology (Lancaster and Downes 2013). The most up-to-date information will, of course, be obtained from periodicals. Since 1985 the Annual Review of Entomology has published over 20 articles on the physiology, behavior, or ecology of aquatic insects. The journal Ereshwater Science (formerly the Journal of the North American Benthological Society), first published in 1986, contains numerous papers devoted to aquatic insects, and the journal Aquatic Insects, first published in 1979, contains articles on all aspects of aquatic insect research, including systematics and taxonomy, life history, ecology, and behavior.

THE TERRESTRIAL-AQUATIC LINK

Despite their occurrence in most aquatic habitats, almost no insect species are completely aquatic. With few exceptions, terrestrial habitats are required for certain stages of their life cycle. The terrestrial stage is often the adult or egg, but even taxa with aquatic adults (e.g., Heteroptera, Coleoptera) usually require access to air [exceptions include taxa such as the Pleidae (Heteroptera) and the Elmidae and Dryopidae (Coleoptera)]. For some taxa, the pupae (e.g., Megaloptera, Neuroptera, Coleoptera) or larvae (e.g., Coleoptera: Dryopidae) are terrestrial. The only insect species known to spend their entire life cycles under water are the stonefly Capnia lacustra in Lake Tahoe (Jewett 1963) and possibly some stygobitic beetles (Spangler and Barr 1995; Balke et al. 2004). The dependence on access to terres trial habitats probably contributes to the prevalence

65

66

Chapter 5 Habitat, Life History, Secondary Production, and Behavioral Adaptations of Aquatic Insects

of complex insect communities in shallow ponds and streams, simple communities in deep rivers and lakes, and the near absence of insects from the open ocean. THE MARINE PARADOX

Although there are over 41,000 species of aquatic insects ( 1 where flows are rapid and the water depths are shallow, and 1 hr

Psephenidae Psephenus falli

1 day?

eariy May-mid Aug

In riffles, under rocks

(mature ovaries at

9 crawls down a rock and

remains submerged for life (1-3 days)

emergence) Elmidae

May-Aug

Stenelmis sexlineata

Lotic, in riffles, on sides and bottom of rocks

Submerged 9 selects depressions or cracks on rocks; deposits group of eggs usually touching each other; each egg pressed against surface for 10-20 sec to glue it down

-

LEPIDOPTERA

Crambidae

Nymph ula sp.

1 day

Juiy-Aug

Lentic, underside of

floating Potamogeton sp. leaves

9 generally does not enter water but extends tip of abdomen to attach egg mass on underside near

margin of leaf. Oviposition occurs at night HYMENOPTERA

Agriotypidae Agriotypus sp.

few days

May-July

Lentic or lotic; in cases

9crawls down a support

of goerid or odontocerid

into water and searches for

caddisflies

a host. Eggs only deposited on prepupa or pupa. 9 may stay underwater for several hr, enveloped in air bubble

DIPTERA

Tipulidae Tipula sacra

< 1 day

June-July

Lentic, in soil or algae mats near shore

Lipsothrix nigrilinea

< 12 hr

Mar-Aug: peak in

In saturated wood in

May-June

streams

9 9 emerge during the day; mate and begin ovipositing immediately

9 searches for suitable site on wood near waterline

with ovipositor. Deposits egg ca. 1 mm deep in soft wood or crack; then moves to make another insertion

-

Chapter 5 Habitat, Life History, Secondary Production, and Behavioral Adaptations of Aquatic Insects

Description of Egg or Egg Mass

Geographic

Incubation and

Number of Eggs*

Hatching Period

elongate oval, somewhat — kidney-shaped; pale yel

6 days @ 19°C; longer in field as oviposition oc

low with smooth cho-

curs at < 14°C

107

Comments

Area

Apparently a short incu- Ontario bation and hatching pe

Reference 2935

riod, as Ist-lnstar iarvae

only found for 3 wk in April

rion; 1.8 X .7 mm

Egg case floats and eggs

egg case Is yellow, turns 10-130 eggs per case; 9 brown; eggs, elongate probably matures more ellipsoid, 4.4 X I mm; than 1 batch bright yellow

Iowa

6622

do not hatch if case turns over; mast as sumed to aid in stabiliz

ing the case spherical, lemon yellow eggs, deposited in com pact, single-layered mass

ca. 500 eggs per 9; sev- 16-17 days @23°C eral may oviposit to gether, forming masses of over 2000 eggs

Apparently synchronous Southern hatching within a mass, California but extended oviposition period

4229

oblong; whitish-yellow; .55-.62 mm long

—

6-10 days @ 22-25°C

Protracted oviposition period; adults live un derwater for > 1 yr

Kentucky

6452

elliptical eggs, .45 X .6 mm; light grey or whi tish; about 20 eggs/

9 of N. badiusalis laid 441 eggs in one night

6-11 days

Direct development of eggs; synchronous hatching within a mass

Michigan

450

In lab, 5-8 days

Several eggs may be de- Japan, France posited on one host but only one larva can de velop per host

mass

elongate, .9 X .2 mm; tapered to a stalk which Is inserted into the

host's integument

X .4 mm; posterior fila

Direct development of eggs; hatching period from early July-mid

ment uncoils when wet

Aug

shining black, elongate, convex on one side; 1.0

dissected 9 9, x = 925, In lab, few days; In range, 500-1600 eggs field, < 1 mo

Alberta

1048, 2228

4812, 4819

ted as anchoring device cream colored, elongate, dissected 9 9, x = 185, About 3 wk @ 16°C smooth; no anchoring range, 106-380 eggs device

Direct development, but Oregon extended hatching pe riod because of long flight period

1530

{continued}

108

Chapter 5 Habitat, Life History, Secondary Production, and Behavioral Adaptations of Aquatic Insects

H Table 5B

Continued

Preovipositlon

Oviposltion

Ovipositlon

Period

Season

Site

Taxon

Ptychopteridae Ptychoptera

< 1 day

late May-June

Lentic, stagnant water

lenis

Oviposition Behavior

Mating and ovipositlon occur shortly after emergence. Eggs occur loose on substrate, so

probably scattered at pond surface and sink to substrate Simuliidae

Simulium spp.

variable; blood meal may be required for egg maturation

spring and summer;

Lotic; various sites (wet

Variable even within a

multivoitlne

ted vegetation, dam

species; may oviposit in flight, but more commonly

faces, debris, etc.)

on solid surface In masses

or strings, at or below Culicidae

Aedes aegypti

variable; blood meal re

nonseasonai

quired for egg development

Culex pipiens

variable; blood meal re

sprlng-iate autumn

artificial containers: cis

Eggs deposited singly, at or

terns, cans, old tires

near wateriine

quired, except In autogen

Lentic; small catch ments and pools with

ous strains; some ovenA/lnter

high organic content

as nulliparous ??

2 lands on water and

deposits eggs In raftlike masses. Oviposltion usu ally at night

Chironomidae Chironomus

2-5 days

mid May, July-Sept

plumosus

Lentic; on water or on flotsam

9 files over water

(sometimes several mi);

extrudes egg mass be tween hind tibiae and

deposits It on first surface that she touches Tabanidae Tabanus atratus

Ephydridae Notiphila

1 wk

5-15 days

(as Dichaeta) (Mathis 1979a)

June-Oct

throughout summer

On plants, near or over

9 faces head downward

water

while depositing egg mass on vertical portion of plant

Marshy areas with ac cumulation of decaying vegetation

9 scatters eggs along shore or on floating detritus. Eggs not glued to substrate but many in crevices

Sciomyzidae Sepedon spp.

4-24 days

Lentic; on emergent vegetation, from 5 cm to

position, deposits eggs in

> 1 m above water

verticai row

9 in head downward

Chapter 5 Habitat, Life History, Secondary Production, and Behavioral Adaptations of Aquatic Insects

Description of Egg or Egg Mass

elongate oval; whitish yellow; longitudinal re

Geographic

Incubation and

Number of Eggs*

dissected 9 9 contain

Hatching Period

In field, 14-20 days

530-806 eggs

109

Comments

Egg maturation occurs during pharate adult

Area

Reference

Alberta

2673

Ontario

1348

Southeastern states

2449

Holarctic

2449

Wisconsin

2596

Florida

3020

stage

ticulations on chorion;

,8-.9 mm long

oval to triangular .25 X ,14 X .13 mm; whitish,

turning brown as they mature.

elongate oval

cylindrical, tapered

300-600 eggs per 9. Eggs may occur In large aggregations (72 000/ft^)

5 days @ 23°C

Successive generations in summer; overwinter

often as diapausing eggs

average about 140 eggs Highly variable; embry onic development com when fed on humans; pleted In 2-4 days after may be 2 or more egg flooding cycles 100-400 eggs per mass; 1-3 days 9 lays 2-4 masses

Direct development in water but eggs withstand desiccation for at

least 1 yr First batch of eggs may mature without a blood meal. Size of later masses

depends on blood meals. Several genera tions per yr.

3 days @ 24°C; 14 days egg mass is dark brown, eggs per mass: X = tear-shaped; swells to 25 1676, range, 1154-2014 @ 9°C X 5 mm. Eggs, cream colored, oval, .5 X .2

egg mass is subconlcal,

Egg mass floats and lar vae remain In It for

1 day after hatching. 2 generations per yr

500-800 eggs per mass 4-12 days

oval at base, with 4-5

tiers of eggs, 5-25 mm X 2-10 mm. Eggs white when laid, then darken

egg ellipsoidal, convex on venter; longitudinally ridged; white; .9 X .3 mm

1-2 days@21-25°C

Ohio, Montana Eggs float when marsh floods and have plastron

1570

for underwater

respiration

Up to 25 eggs per row; 3-5 days eggs He horizontal touching preceding one. 9 probably deposits sev Egg elongate with eral rows coarse, longitudinal striations; white, becom

ing colored during development

USA

4292

110

Chapter 5 Habitat, Life History, Secondary Production, and Behavioral Adaptations of Aquatic Insects

eggs directly on patches of sponge, which would presumably be optimal for larval survival, because

they are unable to enter the water. Instead, they lay eggs on vegetation overhanging streams. When the eggs hatch, first-instar larvae fall to the stream where they must locate a colony of freshwater sponge— presumably by a combination of drifting and crawl ing combined with chemosensory cues. Consequently, the search for a food source with a markedly patchy distribution in a physically rigorous habitat is the province of the tiny first instar. It should thus not be surprising that the mortality of first instar larvae can be extremely high (Willis and Hendricks 1992).

Immature Survivorship and Growth Immature survivorship. Following hatching, only a small percentage of the immatures of most aquatic insects reach the adult stage. Sources of mortality include both abiotic (e.g., physical disturbances such

as floods and drought)and biotic factors(e.g., predation, cannibalism, disease, parasitism) as well as sources associated with molting. Cummins and Wilzbach (1988)found that mortality at molting of all instars in laboratory and field populations of Pycnopsyche guttifer was not significantly different and could not be explained by predation, competition, or food limitation. They proposed that this mortality was entirely due to microbial mortality at molting. In addition, the physical process of molting (Chapter 4) can significantly disrupt oxygen consumption rates of aquatic insects(Camp et al. 2014) and has the poten tial to be another source of mortality. Survivorship curves for five aquatic insects {Glossosoma, Brachycentrus, and Tallaperla and the waterlily leaf beetle Gallerucella [=Pyrrhalta] nymphaea) are consistent with the Type II exponential model of mortality,indicating a constant rate of mortality from hatching to pupation or emergence (Fig. 5.7). This conclusion is probably accurate for G. nymphaea because all stages can be precisely sampled; eggs, larvae, pupae, and adults occur on the upper surfaces of floating spatterdock (Nuphar) leaves (Otto and Wallace 1989). The conclusion of a constant rate for

the immature stages of Glossosoma, Brachycentrus, and Tallaperla, however, is probably less accurate. The data used to produce mortality curves for these taxa were obtained from benthic quadrat samples. Eggs ofstream insects are generally not sampled at all using quadrat-based methods (e.g., Surber or Hess samplers; Benke 1984), and the abundance of first instars is thought to be usually greatly underesti mated. Due to the high rate of mortality expected for first instar immatures. Type III curves are a more

realistic expectation for patterns of mortality for most aquatic insects. This conclusion is supported by the remarkably comprehensive study of the population dynamics of the caddisfly Hydropsyche slossonae by Willis and Hendricks (1992) (Fig. 5.7). Although there was essentially no egg mortality, the mortality for first-instars ofthe caddisfly Hydropsyche slossonae approached 93%,followed by constant but moderate mortality during larval instars II through V, and then high mortality in the pupal stage (Fig. 5.7). Only ~0.5% of all eggs survived to yield adults. The results of Willis and Hendricks (1992) indicates that when the mortality of all life cycle stages is studied in suffi cient detail, Type III survivorship curves may be most representative of survivorship patterns for aquatic insects. Unfortunately, such comprehensive studies are rare.

Immature growth. Growth rates for aquatic insect taxa are determined first by phylogeny and then fur

ther constrained by differences in water temperature (Vannote and Sweeney 1980; Sweeney 1984; Figs. 5.5 and 5.6), food quality (Fuller et al 1988; Thompson 1987; Sweeney 1993, 1984)food quantity (Hart 1987; Feminella and Resh 1990; reviewed by Huryn and Wallace 2000), and predation regime. A number of studies have shown that alterations in the feeding behavior of mayfly nymphs due to the mere presence of predators, e.g., predacious stoneflies and trout, also have strong consequences for their growth rates (reviewed by Huryn and Wallace 2000). Peckarsky et al (1993), for example, showed that in the absence of predators the mass of nymphs of the mayfly Baetis bicaudatus increased 50% over one week. In the pres ence of predators, however, nymphs did not grow at all due to anti-predator behaviors that disrupted their normal feeding patterns. Similarly, McPeek and Peckarsky (1998) predicted that, in the presence of predaceous stoneflies and trout, B. bicaudatus emerge at a mass 50% smaller compared to nymphs reared in

the absence of predators. Limits of immature growth rates. The highest

growth rate known for a stream insect—70% day^'— was reported for larvae of the chironomid Polypedilum in the Ogeechee River, Georgia (Benke 1998). This rate, among the highest estimated for metazoans, is more similar to growth rates of microbes than to other stream macroinvertebrates (Benke 1998)! Rapid growth rates for other macroinvertebrates, particularly the Ephemeroptera, have also been reported for the Ogeechee River as well as Sycamore

Creek, Arizona(^16-25% day'; Gray 1981; Jackson and Fisher 1986; Benke and Jacobi 1994). The midges and mayflies of these streams not only grow rapidly, but also are able to complete their life cycles rapidly

Chapter 5 Habitat, Life History, Secondary Production, and Behavioral Adaptations of Aquatic Insects

111

Brachycentrus spinae Tallaperia maria —

Glossosoma nigrior (summer cohort larvae)

liSEESS 1—I—I—I—I—I—I—r

I

50

100

150

100

200

300

400

Time in days from hatching

Time in days from hatching

Hydropsyche slossonae

Gallerucella nymphaeae

500

E ® 80)

Sweden (a c to

Llll

Georgia(A)

T

100

'

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200

'

r

300

—1

400

Time in days from oviposition

10

1

1

20

1

1

30

1

j—

40

50

Time in days from oviposition

Figure 5.7 A. Survivorship curves for the summer cohort of the caddisfiy Glossosoma nigrior (iarval instars i [L i] through V [L V] based on stream quadrat sampies (Georgian and Waiiace 1983). B. Same for the caddisfiy Hydropsyche slossonae based on egg, iarvai, pupai, and adult stages. Note the high mortaiity during the first iarvai instar (L i), foiiowed by iow mortaiity in iater iarval stages (L il-L V), and then high mortality in pupai to aduit stages (data from Wiiiis and Hendricks 1992). C. Same for iarvai stages of the caddisfiy Brachycentrus spinae (larval instars I [L I] through V [L V])(data from Ross and Wallace 1981), and the stonefly Tallaperia maria larval instars I (L I) through final instar (L F)(data from O'Hop et al. [1984]). D. Same for egg (E), three ian/ai instars (L i - L ili), and pupal stage (P) of the chrysomelid beetle, Gaiieruceiia nymphaeae,from Georgia (U.S.A.) and Sweden. Note the shorter life span and greater mortality in the Georgia population (data from Otto

and Wallace 1989). The equations for the lines are: Georgia, y = 10.1 - 0.23x, 1^=0.98; Sweden, y = 8.4 0.07x, r^ = 0.95. Larva and iarval case icons redrawn and modified from McCafferty (1981), Wiggins (1996), and Stewart and Stark (2002).

because they mature at relatively small sizes (e.g., maximum length ~4-5 mm; Benke 1998; Benke and Jacobi 1994; Gray 1981). In Sycamore Creek, for example, larvae of the chironomid midges and nymphs of the mayfly Leptohyphes complete their growth and development in 100 g DM m~- are considered exceptionally high, what levels of production should be considered exceptionally low? Based on a summary of 58 studies of production for entire macroinvertebrate communities, 40% reported

levels

Diamphipnoidae

D r+

yr

Eustheniidae

I O I

IL Q;

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cAustroperlidae

S'

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> n r+

o

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Nemouridae

n> 0)

5'

Scopuridae Pteronarcyidae

Styloperlidae Peltoperlidae Perlodidae Perlidae

Chloroperlidae Figure 9.6 Phylogeny of Plecoptera.

tenable if the correct placement of the extinct and presumed terrestrial family,"Permithonidae," is as sis ter to Neuroptera(Grimaldi and Engel 2005). Taxa in Osmylidae and Sisyridae are clearly independent deri vations of the aquatic life history. Coleoptera is an extremely species-rich order of insects that has numerous aquatic representatives. Like hemipterans, beetles have invaded the freshwater envi ronment multiple times with many beetle groups having highly modified body forms as both juveniles and adults. Some groups have only aquatic juvenile stages and still more are only semi-aquatic or hygrophilous (water-loving). The prominent aquatic beetles in the suborder Adephaga (often referred to as Hydradephaga)and their relationship to terrestrial groups in the suborder has proved interesting given the mosaic of characteristics found in the relic family Trachypachidae. Features of these beetles have been interpreted as precursors to an aquatic adaptation (Roughley 1981; Bell 1982)or a vestige resulting from a shift away from an aquatic life history (Kavanaugh 1986). The discov ery and description of a new family of aquatic Ade phaga,the Meruidae(Spangler and Steiner 2005),led to an analysis of Adephaga that supports a placement of Trachypachidae as sister to the terrestrial family

Carabidae (Beutel et al. 2005) and a recent analysis focused on mtDNA places Hydradephaga sister to Trachypachidae -f- Carabidae (Lopez-Lopez and Vogler 2017). However, the number of times and

direction of habitat shifts still remain ambiguous because an aquatic life history can be interpreted as primitive for Adephaga with terrestriality reacquired in the Trachypachidae + Carabidae lineage, or that the freshwater habitat was invaded by two or three separate hydradephagan lineages (Beutel 1995). Most aquatic beetles are in the suborder Polyphaga and within this massively diverse group most aquatic beetles are in the Hydrophiloidea or Byrrhoidea. Within the various aquatic families of Coleoptera that are

members of the byrrhoid clade (e.g., Elmidae, Dryopidae, Lutrochidae, Psephenidae, etc.) there are closely related species that occur in regions as widely separated as Australia and South America. However, our present understanding of plate tectonics and continental drift is consistent with such patterns and draws attention to the antiquity of these insects and their success in aquatic habitats (Brown 1987). Archangelsky (2004) presents an explicit phylogeny for higher-level hydrophilid taxa and discusses the evolutionary trends in types of egg cases, larval morphology, and respiratory adaptations.

Chapter 9 Adaptations and Phylogeny of Aquatic Insects

183

Enicocephalamorpha Dipsocoromorpha Mesoveliidae

C

Hebridae

Paraphrynoveliidae Macroveliidae

Hydrometridae Hermatobatidae Veliidae Gerridae

Cimicomorpha Pentatomomorpha Leptopoddidae Omanlidae

Aepophilidae Saldidae

Nepidae Belostomatidae Corixidae

Aphelocheiridae Potamocoridae Naucoridae

Ochteridae Gelastocoridae

Notonectidae Pleidae

Helotrephidae Figure 9.7 Phylogeny of aquatic Hemiptera.

In the Staphyliniformia the Hydraenidae have evolved specialized exocrine secretion systems,including cuticular modifications and grooming behaviors that help maintain the respiratory bubble (Perkins 1997). Amphiesmenoptera is the supraordinal group including Lepidoptera and Trichoptera. This relation ship and the monophyly ofeach ofthe ordinal members are arguably the best supported groups among the higher-level classification of Holometabola. Characters supporting these taxa come from DNA, morphology,

The order Trichoptera appears to have arisen from an ancestor with a larval stage, much like that of the present-day caddisfly family Philopotamidae (Ross 1956), which were more or less free-living rather

and the fossil record. Based on the fossil record, the

and classification have included adult and larval

amphiesmenopteran stem group was probably among

characters, and DNA sequences (Kjer et al. 2001, 2002, 2016). Presently, two subordinal monphyletic groups are recognized, Annulipalpia and Integripalpia (Fig. 9.8). Other works also recognized a third group, Spicipalpia (Wiggins 2004 and Chapter 19). Spicipalpia may also be considered to be a set offam ilies arranged as a grade leading to the Integripalpia. Wiggins and Mackay (1979) suggested that eco logical diversification in Trichoptera can be attributed

taxa included in Necrotauliidae, which includes a few

partially terrestrial and marine species. Except for a few truly terrestrial caddisflies (Flint 1958; Anderson 1967; Wallace 1991), Trichoptera is wholly composed of aquatic species, whereas the vast majority of Lepidop tera are terrestrial. In Lepidoptera,the aquatic and semiaquatic habit is restricted to various groups in Dytrisia with the exception ofthe earlier branching Nepticulidae.

than case-makers. The evolution of the more than

fifty families of Trichoptera is more apparent in the structure and habits of the immature stages, whereas adult structures are valuable in elucidating the evolu tion of groups treated as genera within these families (Ross 1956). A series of refinements of the phylogeny

184

Chapter 9 Adaptations and Phyiogeny of Aquatic Insects

• Hydropsychidae

'

■

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■T'%r.*0,.

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'Ort,

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Hydrobiosidae Rhyacophilidae Plectrotarsidae

Phryganopsychidae Oeconesidae Kokiriidae Pfsuliidae

Brachycentridae Lepidostomatidae Phryganeidae Rossianidae

"Apataniidae" "Goeridae"

Limnephilidae "Thremmatidae"

rD Uenoidae

Atriplectididae Calamoceratidae

Leptoceridae Limnocentropodidae Molannidae Odontoceridae Philorheithridae Tasimiidae

Ceylanopsychidae Anomaiopsychidae Barbarochthonidae Chathamiidae

Helicophidae

Helicopsychidae Petrothrincidae

"Sericostomatidae"

Antipodoeciidae Beraeidae Parasericostomatidae

Conoesucidae Calocidae

Heloccabucidae

Hydrosalpingidae

Figure 9.8

Phyiogeny of Trichoptera.

'

zi.

u'

5L

•g

oj'

Chapter 9 Adaptations and Phylogeny of Aquatic Insects

to the many ways in which species of this order use silk for food gathering, and Wiggins and Wichard (1989)emphasized the importance of how silk is used in preparing for the pupal stage. Other notable studies on phylogeny and ecology of Trichoptera include works by Mackay and Wiggins (1979), Morse (1997), Weaver and Morse (1986), and Wiggins (2004). Hymenoptera includes a few taxa that are para sitic on aquatic invertebrates and so are associated with aquatic habitats. These are all taxa in the more derived Apocrita. Interestingly, although some

early branching groups of wasps are xylophagous (wood-feeding) or phytophagous (plant-feeding), none of these include species that have moved into aquatic systems to take advantage of these resources as have some Diptera and Lepidoptera. Diptera is an order of great diversity ofform and habits. Tipuliodea, Psychodomorpha, Culicomorpha, and Blephariceromorpha include major aquatic or semi-aquatic groups. All of these groups are arrayed as a grade of early branching lineages in the dipteran phylogeny (Fig. 9.9). Many species of more derived Deuterophlebiidae Nymphomyiomorpha Tipuliodea

Ptychopteromorpha I

Blepharicerldae T"

i i~ Psychodidae Psychodidae

1^

^Tanyderldae Tanyderldae

^

j~ Ceratopogonidae %

Chironomidae Simuliidae

41

Thaumaleidae

Dixidae Culicidae Chaoboridae Corethrellidae

Axymyiidae Rhagionidae Pelecorhynchidae Oreoieptidae Anthericidae

Tabanidae

Stratiomyidae I— Dolichopodidae

'""Empididae Phoridae

— Syrphidae

Aulacigastridae Canacidae

Chioropidae Sciomyzidae

Drosophilidae Ephydridae Fanniidae

Muscidae

Scathophagidae Sarcophagidae Calliphoridae

Figure 9.9 Phylogeny of aquatic Diptera.

185

n

c_

n' O

3 o

186

Chapter 9 Adaptations and Phylogeny of Aquatic Insects

flies, scattered across the order, have various levels of

association with the aquatic habitat, though the bulk of the species are terrestrial. Among the more derived families, Tabanidae, Sciomyzidae, and Ephydridae are rich in aquatic species. Chironomidae, a family of Culicomorpha,is particularly remarkable for its diver

sity and its species occupy a very wide range of envi ronmental conditions (Finder 1986). Systematically, chironomids were fundamental as model taxa for

developing an understanding of zoogeography (e.g., Brundin 1966). Of the major orders ofinsects, Diptera has arguably been the most intensively studied by biologists. No doubt this is in part because ofthe obvi ous impact of blood-feeding and disease pathogen vectoring by adults of the aquatic Diptera (Grimaldi and Engel 2005). The current phylogeny for the order by Wiegmann etal.(2011)is well established and there are reviews covering a variety of aspects of the evolu tionary biology for Diptera (Yeates and Wiegmann 2005)and various families,e.g., Chironomidae(Finder 1986) and Simuliidae(Adler et al. 2004).

LIFE CYCLE ADAPTATIONS

Of the major groups offreshwater insects, almost all are regularly represented in the tremendous variety of lentic and lotic habitats occurring throughout the world (see Chapter 3). However, several groups only occur in running-water habitats and many others reach their maximum diversity there. Hynes (1970a) suggests that this may be a consequence ofthe perma nence of streams and rivers when compared with the longevity of most lake and pond habitats. Many river systems have been in continuous existence for long periods of geologic time, whereas lakes persist for relatively short periods and have had little opportu nity to develop a purely lacustrine fauna. Lakes are certainly capable of producing evolutionary change, as shown in the endemic species that occupy ancient lakes such as Lake Baikal in Russia. In time, however, all lakes fill in and disappear; their faunas perish. River faunas have a much greater chance for continu ity and development because although rivers and river systems may change, they rarely disappear entirely. They are not "evolutionary traps" that lentic habitats are and, because of this, many species that retain many primitive features are freshwater organisms pri marily found in lotic environments (Hynes 1970a; Marten et al. 2006; Ribera and Vogler 2000). In addition, most ofthese less derived species (i.e., those that have retained more of the ancestral charac

ters) of aquatic insects are found in running-water habitats because many groups probably first became aquatic by using lotic habitats where high oxygen con

centrations provided more advantageous respiration conditions for larvae and along with resistance to des iccation for adults (see Chapter 4). These factors seem to apply well for the Flecoptera, Ephemeroptera, Odonata, Trichoptera, and Chironomidae. Thus, the absence of primitive members of these groups from lentic habitats could be explained by the hypothesis that they never lived there. Wiggins et al. (1980) give support to this speculation in that insects that occur in temporary pools are primarily derivative groups because the specialized features required to sustain dry periods arose from permanent lentic-dwelling species and these, in turn, arose from lotic species. In addition to differences in their occurrence in

stream and lake habitats, aquatic insects can also be viewed as wanderers between two worlds, the aquatic and the terrestrial. Depending on the species or even the order, the aquatic stage may consist of various combinations of egg, larva, pupa, and adult stages. Just as there is variation in the aquatic portion of the life cycle, there is variation in the terrestrial portion as well. However,the presence of a completely terrestrial larval stage, with the other stages being aquatic, is thus far only known in long-toed water beetles (Dryopidae)(Ulrich 1986). Ferhaps the larva, as the key food gathering stage, is of primary importance for being an aquatic insect. Very few aquatic insects have adapted to a completely submerged life cycle (but see Jewett 1963, for a possible exception among the Fle coptera). At one time or another, nearly all species spend a period in the terrestrial habitat. A major problem in being submerged for even part ofthe life cycle is respiration because,in order to respire while submerged, an insect must receive oxygen from the surrounding aquatic environment (see Chapter 4). Many species have evolved respiratory systems that function in well-aerated water but have not developed survival mechanisms for low oxygen concentrations.In regard to the latter, there is a major difference between running-and standing-water environments. Normally, streams have higher oxygen concentrations(because of turbulence) than do either ponds or lakes. This is certainly a factor in the distribution of Flecoptera and most species ofEphemeroptera and Trichoptera,which are groups that have their maximum diversity in run ning water. Oxygen saturation and temperature are integrally related, and the cooler temperatures that often prevail in running water can contain higher con centrations of oxygen and aid in survival. In high-alti tude (or latitude) lakes where water is cold and highly oxygenated at all times, the distinction between standing- and running-water faunas becomes less clear. Oxygen limitation can result in habitat isolation and consequent speciation. For example, there is a

Chapter 9 Adaptations and Phylogeny of Aquatic Insects

great deal of local, species-level endemicity among Plecoptera that occur in Asian streams and especially among island faunas. This reflects the limitation of tropical Plecoptera to swift streams in mountainous areas. The lack of suitable habitat in lowland areas

restricts their spread within river systems, resulting in typical stonefly habitats being widely separate and isolated (Dudgeon 1999). In addition to the adaptations in the immature and adult stages of aquatic insects, the life cycles often exhibit unique phenological patterns. Aquatic insect populations may produce single or multiple genera tions during a year or, in some species, greater than a year (see Chapter 5). Life-cycle completion time may vary greatly throughout the range ofa species, between populations of the same species (e.g., from more than one generation in a year in the warmer areas to more than one year for each generation in the colder areas), or even between the upper and lower reaches of the same stream (e.g., Resh and Rosenberg 1989). The presence of a diapause in the egg stage, or the formation of a quiescent prepupa, is an important modification that enables the insect to conserve

energy or survive unfavorable conditions. For exam ple, in limnephilid Trichoptera prepupae, the diges tive tract atrophies, but the legs remain functional, which allows the larvae to follow receding water levels (Cummins 1964). Certain life history patterns in aquatic insects may have a definite selective advan tage, particularly in maximizing the efficient use of food sources that have only seasonal availability (see Chapter 5). An example of this can be seen in the leptocerid caddisflies of the genus Ceraclea that feed on freshwater sponge. Because the sponge is available only certain times of the year (only the nonedible gemmules are present during the winter months), life cycles have been modified and alternative food sources are used (Resh 1976a). The significance of life cycle flexibility in the presence of coexisting, system

atically related species has yet to be fully understood, but the ability of these species to share available resources presents interesting implications for habitat partitioning and community evolution. This and many other fascinating aspects of aquatic insect evo lution provide fertile areas for future research.

PHYSIOLOGICAL, MORPHOLOGICAL, AND BEHAVIORAL ADAPTATIONS

The distribution of aquatic insects in the wide variety of habitats present in freshwater environments has led to the evolution of many types of adaptations. Because there is no singular evolutionary line of aquatic insects (i.e., they "got their feet wet" many

187

different times in the course of their evolution), they solved the problems that were inherent from living in freshwater in many different ways. It should be remem bered that when we speak of adaptations, most bio logical structures have an evolutionary plasticity that makes alternative functions possible, and a structure that evolved in one context may later be used for another function. In fact, it is not even necessary that a particular structure had any function or conferred any advantage in its incipient stages. Two processes present fundamental problems for insects to overcome in order to adopt an aquatic exis tence: respiration and osmoregulation. Ofcourse,these are often linked in that the latter can be a consequence of the former process. For example, when oxygen dif fuses through an insect's body surface, water also pen etrates into their bodies. Because the osmolarity of insect body fluids is much higher than that of the sur rounding water, water must be expelled(Wichard et al. 2002, and Chapter 4). This is opposite of the situation in marine and hypersaline environments. Respiration presents one of the most interesting views into the evolution of aquatic-insect adapta tions. Insects have solved the problems of respiration in many different ways. These include the use of airtubes to obtain atmospheric oxygen, cutaneous and gill respiration, the extraction of air from plants, hemoglobin pigments, air bubbles, and plastrons. Air-tubes, which tend to restrict activity to the water surface, have evolved independently in the Hemiptera (Nepidae)and the Diptera {Aedes, Culex, and Eristalis). Furthermore, cutaneous respiration and gill respira tion are widespread in the immature stages of most of the aquatic insect orders. These mechanisms enable the submerged insects to occupy habitats entirely below the water surface and within the substrate.

Most of the species that rely on this type of respira tion require well-oxygenated water, although certain species of the Chironominae that are found in the profundal regions of eutrophic lakes and that nor mally rely on cutaneous respiratory mechanisms may survive periods of oxygen depletion through the use of hemoglobin pigments that aid in oxygen transfer. Respiration by adult aquatic insects such as the bee tles and true bugs often is facilitated by the use of an air bubble, although certain species have evolved a more advanced respiratory mechanism, the plastron (which is a system of microhairs or papillae that hold an air film. Fig. 4.5). A plastron enables the adult to stay submerged for far longer periods than would be possible ifan air bubble mechanism were used(Thorpe 1950). (For a detailed description of aquatic insect respiration and osmoregulation see Chapter 4 and the text of Wichard et al. 2002.) An extraordinary example

188

Chapter 9 Adaptations and Phylogeny of Aquatic Insects

of how various constraints (respiratory needs, fixed number of instars, Dyar's rule) have driven the evolu tion of paternal care is found in the giant water bugs, the Belostomatidae (Smith 1977). BIOLOGICAL TRAITS AND ADAPTATIONS

The use of biological or species traits (such as number of generations per year) initially started with how aquatic insects evolved mechanisms for food acquisition (e.g., Merritt et al. 2002; Cummins et al. 2005). They also have been used in evaluating aquatic insect response to pollution effects (Usseglio-Polatera 1994; Doledec et al. 2000; Usseglio-Polatera et al. 2000), to describe the characteristics of aquatic insect assemblages (Statzner et al. 2001), and to evaluate susceptibility to climate change(Lawrence etal. 2010; Stoks et al. 2014). Buchwalter et al. (2008) demon strated that ecophysiological traits can be related to phylogenetically based differences in aquatic insects. Species traits represent, of course, different types of adaptations to the freshwater environment but they also are the result of evolutionary tradeoffs and com promises. This is important to remember when con sidering adaptations of aquatic insects. Adaptations have occurred in all stages of the insect life cycle, and are characterized by their great flexibility in terms of the evolution of several different mechanisms in response to a specific selective pressure. For example, many adaptations in the egg stage are found in species that occupy temporary pool habitats. The adult females of these species may deposit egg masses close to the ground on the underside of pieces of wood or bark, thereby gaining the advantage of increased humidity and protection from the sun and wind. Certain caddisfly species that are primarily adapted for life in temporary pools have evolved mech anisms that delay or suspend development of the immature stages until soil moisture or the surface water in the basin is sufficient to sustain the newly hatched larvae(Wiggins 1973a). In addition, modifica tions ofthe gelatinous egg-mass matrix can protect the eggs and larvae from desiccation and freezing for peri ods up to seven months(Wiggins 1973a). Other mech anisms used by species inhabiting temporary pools include delaying oviposition until the pools contain water, followed by the prompt hatching and develop ment of the eggs (Corbet 1964; Wiggins et al. 1980). The time spent in the egg stage varies consider ably from species to species, but the time may also vary within a single species population. Whereas all eggs of the damselfly Lestes sponsa will diapause through the late summer,autumn,and winter(Macan 1973), both diapausing and nondiapausing eggs will

be found in the same batch of the stonefly Diura bicaudata eggs(Hynes 1970a). These two egg types of this latter species imply that not all larvae hatch at the

same time. Similarly,eggs ofthe mayfly Baetis rhodani hatch at different intervals (Macan 1973) and other examples of egg diapause bet-hedging have been reported for stoneflies (Frutiger 1996; Zwick 1996). A diapause during the egg stage may enable insects to survive unfavorable periods (see Chapter 5). Like wise, a staggered hatching pattern may also prevent overcrowding of newly hatched larvae in an area that may have limited food resources. The exploding egg masses in the stonefly family Nemouridae, which disperse eggs over a wide area, and the swimming ability of newly hatched caddisfly larvae in genera such as Phryganea, Agrypnia, and Triaenodes may also aid in enhancing dispersal and preventing over crowding. In contrast,the "stickiness" ofmany aquatic insect egg masses prevents displacement. Morphological adaptations of larvae of freshwa ter insects (Table 9A, based on the discussion of Hynes 1970a,b; also see Hora 1930; Nielsen 1951b) are closely followed by behavioral adaptations. In running-water environments, many of the adapta tions can be directly related to hydraulic stress and the continuous struggle ofthe organisms to remain on the substrate (Statzner et al. 1988). There are, however, species that actively exhibit what Waters (1972) has termed behavioral drift, in which individuals enter the

water column and move downstream from their orig inal points of attachment during certain well-defined periods of their daily cycle (for a review on drift, see Brittain and Eikeland [1988]). In standing water, current does not force animals into the open water. However, insects such as Chaoborus (Diptera) and many Coleoptera and Hemiptera actively move through the water column in these environments. (For a description of aquatic insect behavior, see Chapters 5 and 6.) The wide range of biological traits that we see in the larval stages of aquatic insects is evident from the various methods of food acquisition (e.g., as grazers, shredders, gatherers, filterers, predators, and even parasitoids; see Chapters 5,6, and 22). Consequently, these organisms can have an important influence on nutrient cycles, primary productivity, secondary pro duction of fishes, decomposition, and other ecosys tem processes(Wallace and Webster 1996; Huryn and Wallace 2000; Gra9a 2001). Morphological modifica tions of larval stages of aquatic insects often relate to feeding mechanisms. For example, in the dragonflies we see nymphs of Cordulegastridae and Libellulidae with spoon-shaped appendages that, when held together, form a mask that may function as a basket

Chapter 9 Adaptations and Phylogeny of Aquatic Insects

to quickly "scoop up" small prey. In contrast, acshnid dragonflics have pinccr masks that can be extended rapidly to grasp their prey. Damselfly nymphs have masks that are intermediary between these two types and in some cases (e.g., the Calopterygidae) these could be an adaptation to running water because the deeply modified base (the ligula) of the mouthparts may guide water flow and reduce water pressure in front of the mask (Wichard et al. 2002). Aquatic ecologists long believed that the mor phological feature of dorso-ventral flattening of the body (Table 9A), such as modification of the head into the shape of a shield that slopes toward the front (e.g., as in the heptageneid mayflies), enables these nymphal forms to be close to the substrate where the

189

Psephenidae, and Noterus of the Noteridae. Of all aquatic beetles, the pupae of species in the Asian genus Psephenoides are the only ones surrounded with water and that could be described as truly aquatic; the other taxa mentioned above pupate in air-filled cocoons(Leech and Chandler 1956). The adult stages of aquatic insects are generally non-feeding or take only small quantities of liquid food (e.g., Ephemeroptera, Trichoptera, Megalop tera, but with notable exceptions in the Odonata, Diptera, and Coleoptera)and the adults serve entirely as agents of dispersal and reproduction. The mating

droughts and floods, of course, affect whether an

systems of aquatic insects show a wide range of types, including chemical pheromones (e.g., in caddisflies), visual recognition (e.g., note the large eyes in some of the mayfly species that swarm),acoustical recognition (e.g., drumming in stoneflies), and tactile responses (which appears to occur in several groups). Although each of these behaviors is fascinating, the drumming behavior in stoneflies, in which potential mates find each other through signals produced by sub strate vibrations is worth noting phylogenetically (see Chapter 16). For example, this complicated signaling system occurs in the stonefly suborder Arctoperlaria,

insect can become established in a certain aquatic habitat as well. Most stream ecosystems are not in

which is confined to the Northern Hemisphere. In contrast,the southern-hemisphere suborder, Antarcto-

current is reduced. However, we now know that even at the substrate level the current velocities around the

body and the flow forces acting on them are much more complicated (e.g., Statzner and Holm 1982). Benthic insects in fact have body shapes that are com promises between having to live under different cur rent conditions(and hydraulic stresses) as they grow. Water flow, and the extreme conditions of

long term equilibrium, but, unlike lakes, are continu

perlaria,lacks this behavior. It is especially interesting

ally being altered by floods of varying recurrence intervals. There are exceptions under extreme condi tions of dry and wet seasons (Gasith and Resh 1999; Bonada et al. 2006). This is in contrast to geological time in which rivers persist much longer than lakes. These two very different temporal regimes have major consequences in the evolution of aquatic insect com munities. For example, when fish addition or removal experiments are done in lakes, the response of the benthic macroinvertebrate fauna is generally predict able because lake communities are at equilibrium conditions and fish predation reduces numbers of

that the distribution ofthe two suborders coincides with that of the two former subcontinents Laurasia and

prey. In contrast, the results of these types of manip ulative experiments in streams are highly variable because these communities are not at equilibrium. Ofthe holometabolous insects, aquatic pupae are

found in nearly all species ofTrichoptera(cf. Anderson 1967) and aquatic Diptera. Some of the aquatic Lepidoptera (e.g., Petrophila confusalis) that pupate underwater do so in air-filled cocoons. Terrestrial

pupae are found in all Megaloptera and aquatic Neuroptera. Although the aquatic Coleoptera have spe cies of both types, a terrestrial pupa is by far the most common condition among the aquatic beetles. Aquatic pupae are found only in Donacia, Neohaemonia, and Macroplea ofthe Chrysomelidae, Llssorhoplrus ofthe Curculionidae, Psephenoides and Psephenus of the

Gondwanaland, and might reflect the splitting of the

original continent Pangaea and continental drift(Zwick 2000, 2004). The different climatic zones in which aquatic

insects exist expose them to a variety of abiotic fac tors, the most pronounced being temperature, which varies considerably on a yearly basis(although gener ally less than terrestrial temperatures) within the dif ferent geographical regions ofthe world. Temperature fluctuations greatly affect the poikilothermic (cold blooded)insects(Sweeney 1984). Possibly because ofthe relatively constant temperature, more types of emer gence patterns are seen in the tropical regions of the world than in either the arctic or temperate zones. In

permanent water bodies of the tropics, many insects have continuous emergence throughout the year (Corbet 1964; McElravy et al. 1982). This occurs primarily in areas that are located near the equator and undergo small fluctuations in temperature. How ever, continuous emergence has been demonstrated in constant low-temperature mountain brooks where two species of caddisflies have acyclic development and show continuous emergence (Malicky 1980). Distinct seasonal emergence is typical of insects living at high latitudes where regular changes in

190

Chapter 9 Adaptations and Phylogeny of Aquatic Insects

Table 9.A Examples of morphological adaptations of aquatic insects to running-water environments. Representative Groups

Adaptation

Significance

and Structures

Flattening of body

Allows species to live on top of

surface

flattened stones and allows them

Psephenidae (Coleoptera); Epeorus, Many stream inhabitants that do Rhithrogena (Ephemeroptera); not live on exposed surfaces are Gomphidae, Libellulidae (Odonata); also flattened

to crawl through closely compacted substrate

Exceptions or Comments

Molanna, Glossosoma, Ceradea

ancylus(Trichoptera) Streamlining

A fusiform body offers least resistance to fluids

Baetis, Centroptilum (Ephemeroptera); Simulium (Diptera); Dytiscidae (Coleoptera)

Except in the Coleoptera, this body shape is relatively rare Atherix (Diptera), Corydalidae (Megaloptera), and Gyrinidae have large lateral projections; gills of Baetis may reflect respiratory physiology

Reduction of

Projecting structures increase

Gills of Baetis and loss of central

projecting structures

water resistance

cerd in many mayflies (Ephemeroptera)

Provide attachment to smooth

Blephariceridae (Diptera); some Dytiscidae (Coleoptera)

Suckers

surfaces

Friction pads and marginal contact with substrate

Flooks and grapples

Close contact with substrate increases frictional resistance

and reduces chances of being dislodged by current Attachment to rough areas of substrate

Small size

Small sizes permit them to crawl through closely compacted substrates.

Rare in stream animals; if surfaces

are irregular because of moss, algae, etc., suckers are inefficient

Psephenus(Coleoptera); Dicercomyzon, Drunella doddsi, Rhithrogena (Ephemeroptera)

Marginal contact devices are not confined to insects living in the torrential parts of streams

Elmidae, Dryopidae (Coleoptera); Rhyacophilidae (Trichoptera); Corydalidae (Megaloptera)

claws, clawlike legs, and posterior prolegs

Water mites (Flydracarina) in streams are never as large as stillwater Flydracarina and lack the swimming hairs that are found in these latter water mites; large hydrophilids and dytiscids (Coleoptera) inhabit still water; nearly all beetles in fast water

Modified structures include tarsal

Stream animals in other groups are not noticeably smaller than stillwater relatives

are small

Silk and sticky secretions

Ballast

Allows attachment to stones in swift current

Incorporating large stones in

Rheotanytarsus, Orthocladiinae, Simulium (Diptera); Psychomyiidae, Flydropsychidae, Leptoceridae (Trichoptera); Paragyractis (Lepidoptera); Plecoptera and Ephemeroptera eggs

This is at least partially caused by these sized particles being the only ones available for case building

Calopterygidae (Odonata); some Taeniopteryx (Plecoptera); Ephemerellidae (Ephemeroptera)

Most animals that live in vegetation show no particular adaptations that distinguish them from still-water species in vegetation

and less easily swept away Attachment claws

Stout claws aid in attachment

and fixation to plants

Reduction in powers of flight

Fiairy bodies

A loss of individuals from

a restricted habitat may result in smaller populations in following generations Keeps sand and soil particles away while burrowing in substrate

species when they molt and pupate

Goera, Sfenopby/ax (Trichoptera)

cases makes the insects heavier

and dorsal processes

Even free-living caddisflies use silk attachments as do the cased

Reduced wings of stoneflies Reduced flight powers may be a Ailocapnia (Plecoptera); loss of disadvantage because it reduces hind wings in Elmidae (Coleoptera); dispersal ability wingless females of Dolophiloides (Trichoptera) Hexagenia, Potamanthus

(Ephemeroptera)

Permits open spaces for water to flow over body

Chapter 9 Adaptations and Phylogeny of Aquatic Insects

temperature occur. However, wet and dry season cli matic patterns may also lead to seasonal emergence patterns, as has been demonstrated in the tropics (Corbet 1964). As one moves from the tropics to higher latitudes, emergence periods become increas ingly shorter, with the extreme condition occurring in the arctic regions where many species have adapted to cycles in which emergence is limited to a few days out of the entire year. The most widespread rhythm exhibited by aquatic insects is the diel pattern (Remmert 1962), which may influence initial hatching from the egg, feeding behavior of the immature stages, emergence, flight activity, oviposition, and other life history fea tures(Resh and Rosenberg 1989). The significance of diel emergence patterns is that in a short-lived adult insect, such as a mayfly or caddisfly, simultaneous emergence and subsequent swarming by males help to ensure the continuity of the population through the next generation. Periodicity in mate attraction by caddisflies that use sex pheromones also may be an especially important adaptation in this regard (Jackson and Resh 1991). Furthermore, there are differences in diel patterns in the tropics and the temper ate zone. For example, many mayflies form daytime swarms in the former, whereas they form evening swarms in the latter, and perhaps this is to reduce predation. Several species of aquatic insects show rhythmic patterns, such as in the timing of emergence from the pupal stage. In addition, lunar emergence rhythms have been recognized in several aquatic insects that live in tropical climates (Tjonneland 1960; Corbet 1964). The peak in the emergence pat tern for these species coincides with different phases of the moon. For example, the chironomid, Tanytarsus balteatus, has an emergence pattern coinci dent with the new moon (Corbet 1964), whereas other species may have two emergence peaks where the minimum activity occurs during the new and full moon phases (Tjonneland 1960). In temperate regions, the lunar periodicity in adult emergence patterns is exemplified by the chironomid, Clunio marinus(Neumann 1976). This midge larva lives in the intertidal zone of sandy seashores and emer gence is restricted to a few days at the time of the new and full moon (Caspers 1951). A sporadic pat tern, in which emergence appears to occur at irregu lar intervals and seemingly without any environmental cues, seems to be present in only a few aquatic insects (Corbet 1964). Emergence synchrony can also be enhanced by adaptation to day length which, unlike temperature and other environmental cues, is the only one that has no variance.

191

FUTURE PROSPECTS AND QUESTIONS

A satisfactory explanation of why a particular insect is aquatic as a juvenile, has a brief adult sta dium, is cryptic, conspicuous, or even moves and behaves in a certain way requires an understanding of its evolutionary history. Testable evolutionary sce narios are the product of systematics, and names are the purview of taxonomy; both are vital for under standing the natural world and for providing units of information transmission. Therefore, systematics and integrative taxonomy are the underpinning of biology in general. The needs and areas of emphasis in the field of biology at any given time then in turn direct systematic research. This is especially true in consid ering evolutionary studies of aquatic insects. Many fundamental questions in aquatic insect biology remain unanswered and improved techniques in molecular, genomic, and developmental fields will undoubtedly be used to address these inquiries. The basic task of describing diversity for all insect groups remains a major and important focus that is coupled to and integrated with building the tree of life. Until these tasks are closer to completion only moderate gains in improving classifications can be made. Tech

nology is also being applied to streamline the identifi cation process. However, until the entities we wish to identify are characterized, these new techniques can offer only modest gain overall. Specific studies, in contrast, may benefit greatly from the application of new techniques.

Because the aquatic habitat is well defined and a relatively small number of effective sampling methods can collect most aquatic groups, thousands of species ofaquatic insects have been described. Unfortunately, in all groups, and particularly in the holometabolous orders, the immature stages are poorly known. Because it is the immature stages that are usually col lected by aquatic entomologists, the lack of associa tion of the immature and the taxonomically named

adult stage has limited the precision of many studies in aquatic ecology. Associations between immature and adult stages are needed, and they can be made by a variety of rearing techniques (see Chapter 3, espe cially the discussion of rearing methods). Association of the egg and pupal stages with the adult also has been important in elucidating phylogenetic relation ships (Wiggins 1966; Koss 1970). The association of immature and adult stages is an area of research in which all students of aquatic entomology can make valuable contributions because rearing techniques do not require elaborate equipment. These associations eventually can be used in constructing taxonomic keys. We also will likely see an increase in the use of

192

Chapter 9 Adaptations and Phylogeny of Aquatic Insects

DNA sequence data to identify and associate imma ture forms and their adult stages. Efforts using a single small portion of mitochondrial DNA to identify species have achieved some popularity(Hebert et al. 2003; Brownlee 2004; Janzen 2004). Beyond the limited use of sequence similarity or distance-based clustering for groups that have been taxonomically revised, this can only provide clusters of haplotypes that may or may not be equivalent to recognized species (Sperling 2003; Moritz and Cicero 2004; Will and Rubinoff 2004; Will et al. 2005)due to a variety ofissues such as lateral transfer,pseudogenes, and insufficient variation between closely related spe cies(Marcus 2018). For groups that are already taxo nomically well revised, especially in a clearly delimited aquatic system, a sample of DNA may provide the distinguishing characteristics for rapid identification (Curry et al. 2018). However, sequence data are not necessarily always the most efficient character system

for identification; many techniques exist for separa tion of morphologically similar taxa, including dis crimination of early instars (Zloty et al. 1993) and cryptic species (Jackson and Resh 1992) of aquatic insects. Work also needs to be done to link DNA

analysis to morphological/behavioral taxonomy using the gene sequences that actually control the traits used in taxonomic keys. Studies of systematically related, coexisting spe cies often reveal both obvious and subtle mechanisms

of resource partitioning and ecological segregation (e.g.. Cummins 1964; Grant and Mackay 1969; Resh 1976; Mackay 1972; Butler 1984). Similarly, studies of water quality tolerances of congeneric species have been useful in developing the important concept of biological indicators of environmental quality (e.g., Resh and Unzicker 1975 and Chapter 7). Oftentimes congeneric species co-occur, providing ample oppor tunity for such comparative studies. In almost all cases, evolutionary relationships of aquatic insects have been based on studies of morpho logical structures, and this has proven to be a useful way of analyzing systematically related groups. How ever, species are often assigned to different higher taxa because of conflicting opinions on the validity of the various morphological characteristics. In these cases, ecological and behavioral studies may be valu able (Dijkstra et al. 2014). For instance, behavior patterns of larvae of two European species of the mayfly genus Leptophlebia (Solem 1973) agree with that reported for a North American Leptophlebia (Hayden and Clifford 1974). If the diversity of such rhythms is comparable at the generic level, as has been proposed for trophic status by Wiggins and Mackay (1979) (but see Hawkins and MacMahon

[1989] for other considerations in this approach), behavioral studies could provide valuable informa tion on the higher classification of aquatic insects. By combining behavior and other alternative approaches to descriptive systematics, classification and taxon omy may soon be used extensively in predicting eco logical features of systematically related species. There are many generalities that we know about aquatic insects but specific explanations for them are often lacking. These present both interesting research questions and excellent material for "brainstorming sessions." For example,there is an old adage popular ized by Noel Hynes that large benthic macroinverte brates occur in small streams and small ones occur in

large rivers. Is this a function of substrate sizes, refugia availability, etc.? Fikewise, aquatic insects are

rarely obligatory herbivores on certain plants and, yet, in the group closely related to the Trichoptera— the Fepidoptera—host specificity is extremely com mon. Is this related to the timing of the diversification of Trichoptera in freshwater and the evolutionary "flowering" of the angiosperms that occurred later? Why are there so few parasites that attack underwater in aquatic ecosystems? And of course, why aren't there more marine insects? These questions should remind us that this is an exciting time to be studying the links between evolutionary biology and aquatic entomology. As noted recently by Dijkstra et al. (2014), the combination of phylogenetics with exten sive ecological data provides a promising avenue for future research, making aquatic insects highly suit able models for the study ofecological diversification. Finally, several chapters in this book have talked about the use of aquatic insects in biomonitoring, and likely this topic will be a major use for this text. It is interesting to consider whether anthropogenic activi ties have affected aquatic insect evolution? It should be remembered that although benthic macroinvertebrate biomonitoring is based on the different responses of these organisms to pollution, their responses are not evolutionary responses to human pollution per se even though pollution-tolerance scoring systems might imply that this is the case. However, human pollution has not really been around a long enough time to be a selective force. For example,the rat-tailed maggot of the Diptera family Syrphidae is a classic species associated with low oxygen conditions, espe cially given the presence ofa long air tube that enables it to be an atmospheric breather. However,this "indi cator" and many other organic pollution-tolerant dipterans likely evolved to live in rotting, putrefying vegetation or other organic matter. It was through this means that it likely became adapted to thrive in anthropogenically polluted habitats.

.A

"s-all. ,

wfeas^.,.»fe^SSfef AQUATIC INSECTS OF NORTH AMERICA: A PHOTOGRAPHIC OVERVIEW G. W. Courtney Iowa State University, Ames

S. A. Marshall

University ofGuelph, Guelph, Ontario

Macrophotography has increasingly emerged as an important activity for insect taxonomists and amateur entomologists alike, and photographs of living terrestrial insects are now routinely used to document behaviour, distribution and identification

of a wide variety oftaxa. Aquatic insect photography, however, presents some obstacles to the would-be photographer. Most aquatic insects are hidden from view and don't lend themselves to observation in situ,

and even the relatively few taxa easily visible through the water surface are difficult to photograph because ofirregularities and reflection at the water surface. To some extent these obstacles can be overcome by photographing insects in the still water of pools (Figs. 10.270 and 10.271), use of underwater cameras, or by the use of modifications of the "glass bottom bucket" approach. For example, we sometimes use a triangular aquarium with plexiglass sides and a plate

glass bottom (essentially a triangular glass-bottomed bucket) for photography in flowing water (Figs. 10.268 and 10.269). The plexiglass frame is clamped to a heavy swing arm tripod and the aquarium lowered into the water; photographs are taken through the glass bottom. Some photographers use a lighter version of the "glass-bottomed bucket" by inserting their macro lens into a waterproof tube with a glass bottom, whereas others go the whole distance by using a full waterproof housing for their 35 mm

cameras. Such techniques are rewarding because they allow

observation

and

documentation

of the

undisturbed insect and its surroundings, but they are challenging and time-consuming. It is generally more practical to capture aquatic insects prior to photography.

Photography of captured aquatic insects is easily done, either through the still surface of water in a con tainer or through the glass sides of an aquarium (Figs. 10.272 and 10.273). Specimens must be transferred into clean water, and generally require supplemental light (flash) for photography. Especially if shooting directly through the water surface, it is critical to posi tion the flash to avoid reflection into the lens. We find

that best results are obtained using a small aquarium made from microscope slides, with a well-braced cam era and a remote flash unit firing through the side of the aquarium. Some aquatic insects are easily trans ferred to the laboratory for aquarium photography, but some, such as delicate mayflies and stoneflies, are best photographed immediately in the water in which they were found, so we carry small aquaria into the field with us for that purpose. Field photography using aquaria also allows better duplication of the habitat, for example using the same sand or gravel, or frag ments of the same rotting wood on which the insects were collected(Figs. 10.274-10.276). If the substrate is unknown or cannot be duplicated, then it is best to shoot against a neutral grey background. Wherever possible, the photographed aquatic immatures should be retained alive and photographed again (as pupae and adults) as they develop.

The images in this chapter reflect our ongoing efforts to produce a photographic gallery of North American aquatic insects. Some of the images are older slide photographs of taxa we have not yet had the opportunity to re-shoot in recent years, but even a substandard photograph of a living insect often gives a better idea of what the taxon really looks like than a drawing or photo of a preserved specimen.

193

194

Chapter 10 Aquatic Insects of North America

^

C?

m Collembola. Figures 10.1-8. Figure 10.1 Podura (Poduridae), Iowa, Figure 10.2 Podura (Poduridae), Iowa. Figure 10.3 Lepidocyrtus (Entomobryldae), Ontario. Figure 10.4 Tomocerus (Tomocerldae), Virginia. Figure 10.5 Isotomidae, Ontario. Figure 10.6 Immature Sminthuridae & Podura (Poduridae), Ontario.

Figure 10.7 Sminthurinus (Sminthuridae), Virginia. Figure 10.8 Ceratophysella (Hypogastruridae), Ontario.

Figures 1-2 © G.W. Courtney. Figures 3-8 © S.A. Marshall.

Chapter 10 Aquatic Insects of North America

Ephemeroptera nymphs. Figures 10.9-16. Figure 10.9 Analetris (Acanthametropodldae), Montana.

Figure 10.10 Ameletus (Ameletidae), Alberta. Figure 10.11 Ametropus (Ametropodidae), Oregon. Figure 10.12 Arthroplea (Arthropleidae), Wisconsin.

195

Figure 10.13 Heterocloeon (Baetidae), New Mexico. Figure 10.14 Baetisca (Baetiscidae), North Carolina. Figure 10.15 Dolania (Behningiidae), Florida. Figure 10.16 Gaenis (Caenidae), Montana. All figures © G.W. Courtney.

196

Chapter 10 Aquatic Insects of North America

*1 V *

NT

Ephemeroptera nymphs. Figures 10.17-24. Figure 10.17 Drunella (Ephemerellidae), Montana, Figure 10.18 Ephemerella (Ephemerellidae), Michigan. Figure 10.19 Hexagenia (Ephemerldae), Alberta. Figure 10.20 Euthyplocia (Euthyploclldae), French

Colorado.

Guiana.

Figure 20 © C.R. Nelson.

Figure 10.21 Montana.

Raptoheptagenia (Heptagenlldae),

Figure 10.22 Isonychia (Isonychlldae), Michigan. Figure 10.23 Tricorythodes (Leptohyphidae),

Figure 10.24 Traverella (Leptophleblldae), Utah.

Figures 17-19, 21-24 ©G.W. Courtney.

Chapter 10 Aquatic Insects of North America

Ephemeroptera nymphs. Figures 10.25-32. Figure 10.25 Siphloplecton (Metretopodidae), Michigan. Figure 10.26 Neoephemera (Neoephemeridae), South Carolina.

Figure 10.27 Homoeoneuria (Ollgoneurldae), Iowa. Figure 10.28 Homoeoneuria (Ollgoneurldae), male (above) and female (below), Nebraska.

197

Figure 10.29 Ephoron (Polymltarcyldae), Iowa. Figure 10.30 Anthopotamus (Potamanthldae), Michigan. Figure 10.31 Pseudiron (Pseudlronldae), Alberta. Figure 10.32 Siphlonurus (SIphlonurldae), Ontario. Figures 25-31 © G.W. Courtney. Figure 32 © S.A. Marshall.

198

Chapter 10 Aquatic Insects of North America

v;;

Ephemeroptera adults. Figures 10.33-40. Figure 10.33 Analetris (Acanthametropodidae),

Figure 10.38 Hexagenia (Eptiemerldae), Ontario. Figure 10.39 Stenacron (Heptageniidae), Soutti

Montana.

Garolina.

Figure 10.34 Figure 10.35 Figure 10.36 Figure 10.37

Ametropus (Ametropodidae), Oregon. Callibaetis (Baetidae), Ontario. Baetisca (Baetiscidae), Ontario. Gaenis (Gaenidae), Ontario.

Figure 10.40 Isonychia (Isonychiidae), Nebraska. Figures 33, 35-38 © S.A. Marshall. Figures 34, 39, 40 © G.W. Gourtney.

Chapter 10 Aquatic Insects of North America

199

:S; irP.'hS

%

1

4

Odonata nymphs. Figures 10.41-46. Figure 10.41 Anax (Aeshnidae), Wyoming. Figure 10.42 Ophiogomphus (Gomphidae), South Carolina.

Figure 10.43 Cordulegaster (Cordulegastridae) mouthparts, Ontario. Figure 10.44 Cordulegaster (Cordulegastridae), Nevada.

Figure 10.45

Tramea (Libellulidae) mouthparts,

Iowa.

Figure 10.46 Somatochlora (Corduliidae), Illinois. Figures 41, 42, 44, 45 © G.W. Courtney. Figure 43 © S.A. Marshall. Figure 46 © M. Garrison.

200

Chapter 10 Aquatic Insects of North America

V >#■

.*4^.s. %*■ -*[ j ■''

•

$ '

i-m-A

\

Odonata nymphs. Figures 10.47-53. Figure 10.47 Tanypteryx (Petalurldae), Oregon. Figure 10.48 Macromia (Macromiidae), Louisiana. Figure 10.49 Enallagma (Coenagrionidae), Montana. Figure 10.50 Argia (Coenagrionidae), Arizona.

Figure 10.51 Calopteryx (Calopterygidae), New York. Figure 10.52 Lestes (Lestidae), South Carolina. Figure 10.53 Palaemnema (Platystictidae), Brazil. Figures 47-52 © G.W. Courtney. Figure 53 © U.G. Neiss.

Chapter 10 Aquatic Insects of North America

t

Odonata adults. Figures 10.54-61. Figure 10.54 Calopteryx (Calopterygidae), South Carolina.

Figure 10.55 Lestes (Lestidae), Arizona. Figure 10.56 Cordulegaster (Gordulegastridae), Ontario.

Figure 10.57 Gomphurus (Gomphidae), Iowa,

Figure 10.58 Epitheca (Gorduiiidae), Wisconisn. Figure 10.59 Ladona (Libeilulidae), Wisconsin. Figure 10.60 Macromia (Macromiidae), Ontario. Figure 10.61 Tanypteryx (Petaiuridae), Oregon. Figures 54, 57-59, 61 © G.W. Gourtney. Figure 55 © K. Tennessen. Figures 56, 60 © S.A. Marshall.

201

202

Chapter 10 Aquatic Insects of North America

Plecoptera nymphs. Figures 10.62-67. Figure 10.62 Allocapnia (Capniidae), Iowa. Figure 10.63 Leuctra (Leuctridae), Ontario.

Figure 10.64 Zapada (Nemouridae), Oregon. Figure 10.65 Taeniopteryx (Taenlopterygldae), Iowa.

Figure 10.66 Tallaperia (Peltoperlidae), North Carolina. Figure 10.67 Pteronarcys (Pteronarcyidae), Oregon. Figures 62, 64-67 © G.W. Courtney. Figure 63 © S.A. Marshall.

Chapter 10 Aquatic Insects of North America

203

to A.BCfc

Plecoptera nymphs. Figures 10.68-73. Figure 10.68 Paraperia (Chloroperlidae), Oregon. Figure 10.69 Sweltsa (Chloroperlidae), Ontario. Figure 10.70 Acroneuria (Perlldae), Iowa. Figure 10.71 Paragnetina (Perlldae), North Carolina.

Figure 10.72 Clioperia (Perlodldae), North Carolina. Figure 10.73 Skwala (Perlodldae), Oregon. Figures 68, 70-73 © G.W. Courtney. Figure 69 © S.A. Marshall.

204

Chapter 10 Aquatic Insects of North America

i

M

Plecoptera adults. Figures 10.74-81. Figure 10.74 Allocapnia (Capniidae), Iowa. Figure 10.75 Paraleuctra (Leuctridae), Oregon. Figure 10.76 Zapada (Nemouridae), Oregon. Figure 10.77 Taeniopteryx (Taeniopterygidae), Iowa. Figure 10.78 Pteronarcys (Pteronarcyidae), Washington.

Figure 10.79 Callineuria (Perlldae), Oregon. Figure 10.80 Isoperia (Perlodldae), North Carolina. Figure 10.81 Sweltsa (Chloroperlidae), Oregon. All figures © G.W. Courtney.

Chapter 10 Aquatic Insects of North America

Orthoptera. Figures 10.82-86. Figure 10.82 Tetrix (Tetrigidae), Iowa. Figure 10.83 Tettigidea (Tetrigidae), South Carolina. Figure 10.84 Conocephalus (Tettigoniidae), Ontario. Figure 10.85 Neotridactylus (Tridactylidae), Ontario.

Figure 10.86 Gryllotalpus (Gryllotalpidae), South Carolina.

Figure 82 © G.W. Courtney. Figures 83-86 © S.A. Marshall.

205

206

Chapter 10 Aquatic Insects of North America

Heteroptera-Nepomorpha. Figures 10.87-95.

Figure 10.87 Belostoma (Belostomatidae), Iowa. Figure 10.88 Abedus (Belostomatidae), New Mexico. Figure 10.89 Ambrysus (Naucoridae), Idaho. Figure 10.90 Ranatra (Nepidae), Ontario. Figure 10.91 Neoplea (Pleldae), Iowa.

Figure 10.92 A/epa (Nepidae), Iowa. Figure 10.93 Notonecta (Notonectldae), Ontario. Figure 10.94 Gelastocorus (Gelastocorldae), Arizona. Figure 10.95 Sigara (Gorlxidae), Ontario. Figures 87-89, 91, 92 © G.W. Courtney. Figures 90, 93-95 © S.A. Marshall.

Chapter 10 Aquatic Insects of North America

207

%

103

Heteroptera-Gerromorpha and Leptopodomorpha. Figures 10.96-103. Figure 10.96 Metrobates (Gerridae), Ontario. Figure 10.97 Limnoporus (Gerridae), North Carolina. Figure 10.98 Rhagovelia (Veliidae), South Carolina. Figure 10.99 Microvelia (Veliidae), Ontario. Figure 10.100 Mesovelia (Mesoveliidae), Iowa.

Figure 10.101 Hebrus (Hebridae), Ontario. Figure 10.102 Hydrometra (Hydrometridae), South Carolina.

Figure 10.103 Pentacora (Saldidae), Ontario. Figures 96, 98, 99, 101, 103 © S.A. Marshall. Figures 97, 100, 102©G.W. Courtney.

208

Chapter 10 Aquatic Insects of North America

y*■ >

•a

Neuropteroids. Figure 10.104 Figure 10.105 Figure 10.106 Figure 10.107 Figure 10.108

Figures 10.104-111. Nigronia (Corydalidae) larva, Ontario. Corydalus (Corydalidae) larva, Iowa. Sialls (Sialidae) larva, Oregon. Climacia (Sisyridae) larva, Oregon. Nigronia (Corydalidae) adult, Ontario.

Figure 10.109

Corydaius (Corydalidae) adult, Costa

Rica.

Figure 10.110 Siaiis (Sialidae) adult, Ontario. Figure 10.111 Ciimacia (Sisyridae) adult, Ontario. Figures 104, 108-111 ©S.A. Marstiall. Figures 105-107 © G.W. Courtney.

Chapter 10 Aquatic Insects of North America

-

209

■

^ 5^;^ *Vl16'

Coleoptera-Myxophaga and Adephaga. Figures 10.112-119. Figure 10.112 Hydroscapha (Hydroscaphidae) larva, Arizona.

Figure 10.113 Hydroscapha (Hydroscaphidae) adult, Arizona.

Figure 10.114 Amphizoa (Amphizoidae) larva, Montana.

Figure 10.115 Amphizoa (Amphizoidae) adult, Oregon. Figure 10.116 Chlaenius (Carabidae) larva, Iowa. Figure 10.117 Omophron (Carabidae) adult, Texas. Figure 10.118 Dytiscus (Dytiscidae) larva, Iowa. Figure 10.119 Celina (Dytiscidae) adult, Iowa. All figures © G.W. Courtney.

210

Chapter 10 Aquatic Insects of North America

Coleoptera-Adephaga. Figures 10.120-126. Figure 10.120 DIneutus (Gyrinidae) larva, Ontario. Figure 10.121 Dineutus (Gyrinidae) adult, Iowa. Figure 10.122 Peltodytes (Haliplidae) larva, Ontario. Figure 10.123 Haliplus (Haliplidae) larva, Iowa. Figure 10.124 Peltodytes (Haliplidae) adult, Iowa.

Figure 10.125 Hydracanthus (Noteridae) larva, Ontario.

Figure 10.126 Hydracanthus (Noteridae) adult, Iowa. Figures 120, 122, 125©S.A. Marshall. Figures 121, 123, 124, 126©G.W. Courtney.

Chapter 10 Aquaticlnsects of North America

Coleoptera-Polyphaga. Figures 10.127-133. Figure 10.127 Donacia (Chrysomelidae) larva, Ontario.

Figure 10.128 Donacia (Chrysomelidae) adult, Ontario.

Figure 10.129 Bagous (Gurculionidae) adult, Ontario.

211

Figure 10.130 Helichus (Dryopidae) adult, Iowa. Figure 10.131 Stenelmis (Elmidae) adult, Ontario. Figure 10.132 Lara (Elmidae) larva, Montana. Figure 10.133 Stenelmis (Elmidae) larva, Ontario. Figures 127-129, 131, 133 ©S.A. Marshall. Figures 130, 132©G.W. Courtney.

212

Chapter 10 Aquatic Insects of North America

A v.

m

.JftliPTJAliiMkll

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is

:W #^r, orflOT«s Vein A, of forewings not forked near margin, attached to hind margin by three or more small veins(Fig. 13.168); abdomen of most species with distinctive dark contrasting patterns on terga and sterna 93 Cubital intercalary veins offorewings consist of a series of small veins, often forking or sinuate that attach vein CuA to hind margin of wing (Figs. 13.169, 13.175) 9 Cubital intercalary veins of forewings variable, but not as above (Figs. 13.176, 13.181, 13.183, 13.186-13.187); sometimes absent (Fig. 13.184) 10 Remnants of gill tufts (often purplish colored) present at sides of vestigial maxillae and bases of forecoxae; forelegs largely or entirely dark, but middle and hind legs pale; vein MP of hind wings forked near margin (Fig. 13.175); terminal filament vestigial ISONYCHIIDAE.../son>'c/H'fl

9'

Remnants of gill tufts not present on vestigial maxillae and forecoxae; legs not colored as above; vein MP of hind wings forked near base or near middle, but

10(8')

All three caudal filaments well-developed

10'

Only two caudal filaments (cerci) well-developed and apparently present, terminal filament rudimentary or absent Hind wings relatively large with one or more veins forked; costal projection shorter than wing width (Figs. 13.176, 13.181, 13.183) Hind wings small with only two or three simple veins or hind wings absent (Figs. 13.189, 13.190); if hind wings present, costal projection long (1.5 to 3.0 times width of wing); costal projection straight or recurved (Fig. 13.188)

not near wing margin (Figs. 13.169, 13.177); terminal filament variable

11(10) 1r

297

19 11 14 12

18

HIND LEG

Figure 13.159

terminal filament

) ) ) ) ) ) ) ) ) ) ) )))) ) ) ) ) ) ) ))) ) )

Figure 13.158 Dorsal view of genitalia of Ephemerella sp. male imago (Ephemerellidae).

Figure 13.159 Dorsal view of apex of abdomen and caudal filaments of

MIDDLE LEG

3

ABDOMEN

HIND WING

CAUDAL FILAMENTS

Ephemerella sp. female imago (Epfiemerellldae).

trochanter

femur

eye

Figure 13.158

SUBGENITAL PLATE

Figure 13.157 Lateral view of Ephemerella sp. male imago (Ephemerellidae),

frons

oce ti

PROTHORAX

veins

intercalary

unattached

outer margin

ANp = anteronotal protuberance.

Figure 13.157

antenna

1 (basal)

MESOTHORAX

coastal projection

builae

FOREWING

stigmatic area •

Chapter 13 Ephemeroptera

Sc

299

Ri

Figure 13.161

Figure 13.160

MESOTHORAX

hind margin

Figure 13.163

METATHORAX

Figure 13.162

Figure 13.165 Figure 13.164

Figure 13.160 Forewing (a) and hind wing (b) of Lachlania sp.(Ollgoneurlidae), arrows indicate primary veins and reduced crossveins.

Figure 13.161 Ventral view of male genitalia of Euthyplocia sp. (Euthyplociidae). Figure 13.162 Lateral view of thorax and wing of Ephemerella sp.(Ephemerellidae). Figure 13.163 Forewing (a) and hind wing (b) of Euthyplocia sp. (Euthyplociidae).

Figure 13.164 Forewing (a) and hind wing (b) of Dolania americana (Behningiidae), arrow indicates first intercalary vein behind vein CuA. Figure 13.165 Forewing (a), arrows indicate curved base of veins MPg and CuA and weak costal crossveins and hind wing (b), arrow indicates acute costal angle in Neoephemera sp.(Neophemeridae).

Figure 13.167

Figure 13.166

forceps

penes

forceps

Figure 13.168 Figure 13.172

Figure 13.170

,(

'

./

Figure 13.173

Figure 13.169

Figure 13.171

Figure 13.166 Forewing (a) and hind wing (b) of Ephoron sp.(Poiymitarcyidae), arrows indicate curved

Figure 13.174

Figure 13.170 Ventral view of male genitalia of Dolania americana (Behningiidae).

bases of veins MPg and CuA and anastomosed

Figure 13.171

crossveins near wing margin. Figure 13.167 Forewing (a) and hind wing (b) of Anthopotamus sp.(Potamanthidae), arrows indicate curved bases of veins MP2 and CuA and forked vein A^. Figure 13.168 Forewing (a) and hind wing (b) of Ephemera sp.(Ephemeridae). Figure 13.169 Forewing (a) and hind wing (b) of Siphlonurus sp.(Siphlonuridae), arrows indicate cubital intercalary veins.

americana female imago (Behningiidae), arrow indicates prominent anteroiateral projections. Figure 13.172 Ventral view of male genitalia of Pentagenia vittigera (Palingeniidae). Figure 13.173 Dorsal view of pronotum of Ephemera sp. male imago (Ephemeridae). Figure 13.174 Dorsal view of pronotum of Pentagenia vittigera male imago (Ephemeridae).

300

Dorsal view of head of Dolania

Figure 13.176

Figure 13.175

Figure 13.179

Figure 13.180 Figure 13.178 Figure 13.177

Figure 13.182

Figure 13.181

Figure 13.183

Figure 13.184

Figure 13.185

Figure 13.175 Forewing (a) and hind wing (b) of Isonychia sp. (isonychlldae), arrows indicate cubital intercalary veins. Figure 13.176 Forewing (a) and hind wing (b) of Ametropus sp.(Ametropodidae), arrows indicate cubital intercalary veins. Figure 13.177 Forewing (a) and hind wing (b) of Acanthametropus sp.(from Russia) (Acanthametropodidae). Figure 13.178 Hind wing of Acanthametropus sp. (from Russia)(Acanthametropodidae). Figure 13.179 Hind wing of Analetris eximia (Acanthametropodidae). Fig.13.180 Ventral view of male genitalia of Analetris eximia (Acanthametropodidae).

Figure 13.181

Forewing (a) and hind wing (b) of

Ephemereiia sp. (Ephemerellidae), arrows indicate detached marginal intercalary veins. Figure 13.182 Ventral view of male genitalia of Acanthametropus sp.(Acanthametropodidae)[from Russia], Figure 13.183 Forewing (a) and hind wing (b) of Paraleptophlebia debiiis (Leptophlebiidae), arrows indicate attached marginal intercalary veins.

Figure 13.184 Forewing (a), arrow indicates vein A, and hind wing (b), arrow indicates marginal intercalary veins of Baetisca rogersi (Baetiscidae). Figure 13.185 Forewing (a) and hind wing (b) of Acentreiia sp.(Baetidae), arrow indicates paired detached marginal intercalary veins. 301

302

12(11)

12' 13(12')

Chapter 13 Ephemeroptera

Vein Aj of forewings attached to hind margin by series of small veins (Fig. 13.176); forewings with two pairs of cubital intercalary veins, anterior pair long, posterior pair very short AMETROPODIDAE..../i»/ef»'(?pKs Vein of forewings not attached to hind margin as above (Figs. 13.181, 13.183); cubital intercalary veins not as above 13 Short, basally detached marginal intercalary veins present between primary wing veins along entire outer margins of fore- and hind wings(Fig. 13.181); male forceps with one short terminal segment (Figs. 13.240-13.246)

13'

14(10')

EPHEMERELLIDAE....74

Short basally detached marginal intercalary veins usually absent along outer margins of wings (occasionally a small single unattached marginal vein may occur irregularly along wing margins); most marginal intercalary veins attached (Fig. 13.183); male forceps with two or three short terminal segments (Figs. 13.229-13.230, 13.281-13.283) LEPTOPHLEBIIDAE Hind wings with numerous, long, free marginal intercalary veins(Fig. 13.184); cubital intercalary veins absent in forewings with vein A|terminating in outer margin of wings(Fig. 13.184)

BAETISCIDAE...Raet/sca

14'

Hind wings not as above or absent; if hind wings present then cubital intercalary veins present in forewings with vein Aj terminating in hind margin of wings

15(14')

Short, basally detached, single or double marginal intercalary veins present in each interspace of forewings and veins MAjand MPjdetached basally from their respective stem veins(Fig. 13.185); hind wings small or absent; penes of male

(Figs. 13.185-13.187

15

membranous; upper portion of compound eyes of male turbinate (i.e., raised on a stalk-like lower portion; Fig. 13.195 BAETIDAE*

15'

64

25

Marginal intercalary veins attached basally to other veins; veins M Aj and MP2 attached basally (Figs. 13.187-13.188); hind wings relatively large; penes of male well-developed; compound eyes of male not turbinate

16

16(15')

Hind tarsi apparently four segmented with basal segment fused or partially fused to

16'

tibiae (Figs. 13.192-13.193); hind tarsi longer than hind tibiae; one or two dissimilar pairs of cubital intercalary veins present (Figs. 13.186-13.187) 17 Hind tarsi distinctly five segmented (as in Fig. 13.194); hind tarsi shorter than hind tibiae; two pairs of cubital intercalary veins present similar to those in Fig. 13.186 HEPTAGENIIDAE.(in part)/ARTHROPLEIDAE....48

17(16)

Compound eyes of male contiguous (i.e., touching) or nearly contiguous dorsally (similar to Fig. 13.219); foretarsi three times the length of foretibiae; abdomen offemale not noticeably long and slender, posterior margin of sternite IX evenly convex

17'

Compound eyes of male separated dorsally by twice the width of median ocellus;

without median notch

METRETOPODIDAE

63

foretarsi two times the length of foretibiae; abdomen of female distinctly long and slender, posterior margin of sternite IX with median notch (Fig. 13.191); rare, sand dominated rivers

18(11')

PSEUDIRONIDAE.../'sra 3 3 ) J 3 3 )> 3 D D

)

■1^ o

)

Family Genus

Macrodiplax

Syntetrum)

Libellula (19) {=Belonia, Eolibellula, Eurothemis, Holotania, Leptetrum, Neotretrum,

Leucorrhinia (7)

Ladona (3)

Idiataphe (=Ephidatia)

Erythrodiplax (7)

Continued

balteata

(brackish water)

Lentic—littoral

(sediments)

depositional

lotic—

hydrophytes):

vascular

(silt and detritus.

Lentic—littoral

hydrophytes (Including bogs)

Lentic—vascular

(sediments in ponds)

Lentic—littoral

water)

zone—brackish

Lentic—vascular

hydrophytes (emergent

(=Ephidatia)

hydrophytes

Lentic—^vascular

Habitat

cubensis

Species

)

)

)

)

)

)

)

r)

)

)

Habit

)

Sprawlers

Sprawlers

Climbers

Sprawlers

Sprawlers

Climbers

'SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic '*Emphasis on trophic relationships

Order

parentheses)

(number of species in

Taxa

Table 14A

)

North

)

(engulfers)

Predators

(engulfers)

Predators

)

(engulfers. await prey; Diptera, Coleoptera, Trichoptera, Ephemeroptera)

Predators

(engulfers)

Predators

(engulfers)

Predators

(engulfers)

Predators

)

Southeast)

)

(particularly

South

Widespread

in North

Widespread

Widespread

Florida

South, East

Trophic American Relationships Distribution

)

9.8

SE

)

9

UM

)

M

Ecological

)

9

8

)

)

2164, 4283

420, 556, 4110, 4436, 4824, 5077, 5526, 6577, 6691, 6692, 6490, 1204, 573, 3334

5560, 6240, 2982, 2983, 2984, 3334

4811, 4824, 2529, 5526,

4283,4413,

438, 2164, 4283, 4699

)

420, 421,422,

2164, 4283

5526

1624, 4283,

NW MA* References**

Tolerance Values

)

J

o

Family Genus

Predators

Predators

Sprawlers

Lentic—littoral

Orthemis(3)

Pantala (2)

lineatipes

Paltothemis

Lotic—erosional

ponds)

(sediments and macroalgae in temporary

Lentic—littoral

(among rocks) Sprawlers (active foragers)

Sprawlers

ODONATA

Chironomidae)

(engulfers;

Predators

(engulfers)

Predators

(engulfers)

Predators

(detritus and silt); lotic— depositional (detritus)

Sprawlers

longipennis

Pachydiplax Lentic—littoral

(engulfers)

{=Neocysta)

Nannothemis

(engulfers)

Widespread

Southwest

North)

Widespread (except far

South

East

Arkansas

Climbers—

Texas,

Predators

Extreme

South

Predators

(engulfers)

(engulfers)

Arizona, Texas

Predators

(engulfers)

sprawlers

Sprawlers

Climbers

Sprawlers

Habit

hydrophytes (emergent zone in small puddles away from water's edge)

hydrophytes)

(vascular

Lentic—littoral

hyacinth roots)

(in water

Lentic—littoral

Lentic—littoral

Habitat

Lentic—vascular

bella

marcella

Species

North American Trophic Relationships Distribution

(=Aino)

Micrathyria (4)

(—Nothifixis)

Miathyria

Macrothemis(4)

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

(number of species in parentheses)

Taxa

Table 14A

9,6

SE

UM

M

NW

Tolerance Values

(.continued)

3334

5526,

367,818, 4283, 2187,

5526

1543, 4283,

5077, 6120, 6692,3334

1539, 4137, 4283, 5526,

4283, 5526

3334

2164,4283,

2164, 4283

2164, 4283

2164, 4283

MA* References*

Ecological

))))))

Family

Predators

Sprawlers

Lentic—littoral

Lentic—littoral

(silt and detritus,

Tholymis Tramea (7)

(=Trapezostigma)

hydrophytes, and macroalgae)

vascular

Sprawlers

Lentic—littoral

Tauriphila (3)

Sprawlers

(engulfers)

Predators

(engulfers)

Predators

(engulfers)

Predators

vascular

—Tarnetrum)

hydrophytes in ponds)

Predators

(engulfers)

(engulfers) Climbers—

Sprawlers

Predators

(engulfers)

sprawlers

depositional

Lotic—

Lentic—littoral

hydrophytes in ponds) Sprawlers

(engulfers)

sprawlers

(detritus and vascular

Predators

Climbers—

Lentic—littoral

atnna

Predators

(engulfers)

Lentic—littoral

Sprawlers (active)

Lotic—

Habit

depositional (margins)

Habitat

(detritus and

superbus

Species

North

Widespread

Florida, Texas

Florida, Texas

Widespread

Southwest

Widespread

Texas

North)

Widespread (except

Trophic American Relationships Distribution

Sympetrum (14) {=Diplax,

Pseudoleon

Plathemis(2)

Planiplax {=Platyplax)

Perithemis(3)

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

parentheses)

(number of species in

Taxa

Table 14A

7.3

' 10

10

8

8.2

NW

Tolerance Va ues

Ecological

4

4

3334

490, 1378, 3456, 4283, 5526, 6121, 6122, 6691, 6692, 6694,

4550

4283

4002, 4699, 5526, 6764, 6692, 3334

4323,4811,

1122,4283,

5526

5526

4283, 5077,

6692

2164,4137, 4283, 5077,

MA* References**

) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ))) ) ) ) ) )) ) )

o

4^

o

depositional; Ientic—littoral

Damselflies

Climbers

Climbers

**Emphasis on trophic relationships

ODONATA

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic

hydrophytes

Ientic—vascular

hydrophytes);

vascular

(detritus and

depositional

Lotic—

Lotic—

detritus)

(engulfers)

Predators

(engulfers)

Predators

Predators

(engulfers)

Climbers—

clingers

Lotic—erosional

Lestidae (19)-

Archilestes(2)

Predators

(engulfers)

and depositional (margins and

Spread-Winged

Hetaerina (3)

detritus)

and depositional (margins and

Lotic—erosional

(detritus)

(margins and

Climbers

Predators

Generally climbers

Generally lotic— erosional

(engulfers)

Predators

(engulfers)

Generally climbers

Both ientic and

Habit

North

West

Widespread

Widespread

Trophic American Relationships Distribution

lotic habitats

Habitat

depositional

Species

Damselflies

Calopteryx(5) {=Agrion)

Genus

detritus) and

Family

Continued

6.2

8.3

SE

6

5

UM

2.8

3.7

M

6

NW

Tolerance Values

2164, 3179,

574

2851, 5526,

6238

2164, 3179, 4599, 5526,

2043

5526, 6238,

6238, 3334

856, 3753, 4812, 5526,

6238

4599, 5526,

{continued)

6

5

6466

35,603, 1223, 1344, 2164, 2995, 3179, 3449, 3450, 4558, 4599, 5063, 5526, 6212, 6238,

MA* References**

Ecological

j ) 3 ))))) j ))))) > :) ) )

Calopterygidae (8) (=Agrionidae, =Agriidae) Broad- Winged

Zygoptera (damselflies)

Order

parentheses)

(number of species in

Taxa

Table 14A

j j j j

Lotic—

Amphiagrion (2)

hydrophytes (emergent zone) including bogs

lentic—^vascular

hydrophytes);

(vascular

depositional

hydrophytes

Lentic—vascular

Acanthagrion (=Myagrion) quadratum

habitats

(=Agrionidae)Narrow-Winged

Damselflies

Wide range of

depositional

Lotic—

hydrophytes)

(vascular

depositional

lotic—

Climbers

Climbers

climbers

Generally

Climbers

Climbers— swimmers

hydrophytes;

Habit

Lentic—^vascular

Habitat

lentic and lotic

domina

Species

Coenagrionidae

Palaemnema

Lestes(17)

Genus

(109)

Damselflies

Shadow

Platystictidae (1)-

Family

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

species In parentheses)

(number of

Taxa

Table 14A

North

(engulfers)

Predators

(engulfers)

Predators

(engulfers)

Predators

(engulfers)

Predators

(engulfers)

Predators

North)

Widespread (particularly

Texas

Southwest

Widespread

Trophic American Relationships Distribution SE

UM

6.1

M

5

9

9

9

5526, 6238, 6470, 3334

2070

6238

2164,3179, 4599, 5526,

4550

871, 1122, 1830, 2045, 2851, 3603, 4268, 5291, 2469, 5526, 6238, 6437, 266, 3003, 4423, 3334

Ecological NW MA* References**

Tolerance Values

) ) ) ) )) )) ) ) ) )) ) ) ) ) ) ) ) ) ))) ))

o o^

Taxa

Family

Coenagrion (3) {=Agrion)

Chromagrion

Argia (32) i=Hyponeura)

Apanisagrion

Genus

conditum

lais Predators

depositional; and littoral

bogs)

(marshes and

hydrophytes

vascular

hydrophytes at margin); lentic—

vascular

and depositional (emergent

Lotic—erosional

streams)

hydrophytes In small spring

vascular

(detritus and

depositional

(detritus) and

Lotic—erosional

(sediments)

Climbers

Climbers

ODONATA

North, East, West

Predators

East

Widespread

Arizona

(engulfers)

(engulfers)

Predators

Predators

(engulfers)

Climbers—

(engulfers)

clingers— sprawlers

lentic—erosional

North American

Relationships Distribution

Lotic—erosional

Climbers

Habit

Trophic

(sediments and detritus) and

hydrophytes

vascular

depositional (spring fed regions)—

Lotic—

Habitat

*SE = Southeast, DM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atiantic **Emphasis on trophic relationships

Order

species in parentheses)

Continued

SE

UM

5.1

M

7

NW

Tolerance Values

265, 267,874, 4525, 5526, 266, 2982, 2983, 6238, 2984, 2986, 486, 2057, 2187, 1204

3334

4586, 6238,

2047, 2164,

599, 1347, 2161, 2162, 3107, 3271, 4815, 5526, 6238, 2043

(continued)

6

4550

MA* References**

Ecological

j ))) J 3 ))>)))) ) ) > ) ) )) ) )

Table 14A

J

(number of

j j J

1

■t^

1

o oe

Family Genus

lotic—

Anomalura, Bedfordia, Celaenura, Ceratura, Ischnosoma, Ischnuridia,

)

)

)

Predators

Trichocnemis)

)

)

)

)

)

)

)

)

)

(engulfers)

Predators

Climbers

Lentic—vascular

hydrophytes (including edges of bog mats)

(=Argiallagma,

Predators

)

Cladocera, Chironomidae)

(engulfers;

Nehalennia (5)

)

Predators

(engulfers)

(engulfers)

Climbers

9

9

)

Ecological

9

9

9

8

)

2047,2757,

NW MA* References**

)

)

East, North

Florida, Texas

Widespread

>

)

)

3334

)

2164, 6238,

2042, 2070,

)

5,2186, 4549

859,1476, 2050, 2055, 2164, 35, 2237, 2993, 2996, 3107, 4268, 4355, 4525, 5526, 2565, 5966, 5967,6238, 269, 271, 272, 2234, 2487, 3965, 4423, 5078, 3334

2164

3108, 3271, 4269, 4280, 4824, 5526, 5928, 6238, 1020, 1021, 1022, 1347, 183, 269, 555, 6126, 2997, 3963, 4700, 6527, 3334

9.4

9

M

Cladocera)

Southwest

Widespread

UM

2758, 2852,

Predators

SE

Tolerance Values

(engulfers;

Loticytentlc

seeps

hydrophytes). small spring

(vascular

Climbers

Climbers

Climbers

Habit

North

Trophic American Relationships Distribution

Leptobasis (3) (=Chrysobasis)

Micronympha)

hydrophytes; depositional

Lentic—^vascular

hydrophytes

Lentic—vascular

depositional (vascular hydrophytes)

lotic—

alkaline waters):

brackish and

hydrophytes (including

Lentic—vascular

Habitat

(=Anomalagrion,

heterodoxum

Species

Ischnura (14)

Hesperagrion

Enallagma (38) i=Chromatallagma, Teleallagma)

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Ennphasis on trophic relationships

Order

species in parentheses)

(number of

Taxa

Table 14A

)

so

o

Family Genus

Sparganium)

(vascular

hydrophytes Sparganium)

(bases of

Lotic—

depositional

Zoniagrion exdamationis

Climbers

ODONATA

(engulfers)

California

Southwest

in South,

Predators

Widespread Predators

Climbers

Lentic—vascular

hydrophytes

Telebasis(3)

(=Erythragrion, Helveciagrion)

Texas

Texas

Florida, Texas

(engulfers)

leaves)

hydrophytes)

(vascular

Predators

(engulfers)

Climbers—

clingers(on floating

Lotic—

Protoneura

depositional

Predators

(engulfers)

Climbers—

cara

Predators

(engulfers) clingers

Climbers

Habit

Lotic—erosional

hydrophytes

Lentic—vascular

Habitat

(on rocks)

cultellatum

Species

North

Trophic American Relationships Distribution

Neoneura (2) (=Caenoneura)

Neoerythromma

Continued

*SE = Southeast, DM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

species in parentheses)

(number of

Taxa

Table 14A

j j j j l )) ))))))))

SE

UM

M

Ecological

9

3108,5627

6432

5204, 5526,

4550

2164

2042

NW MA* References**

Tolerance Values

SEMIAQUATIC ORTHOPTERA Hojun Song Texas A&M University, College Station, Texas

INTRODUCTION

With more than 28,000 extant species, Orthoptera is the most diverse order among the polyneopteran insect lineages (Cigliano et al. 2018). The order includes familiar singing insects, such as crickets and katydids, as well as often-devastating pests, such as grasshoppers and locusts. Orthopteran insects have diversified into numerous lineages that occupy every conceivable terrestrial habitat outside the polar regions and play integral roles in their ecosystems. Aquatic habitats have also been colonized by several orthopteran lineages, and some species even have unique morphological or behavioral adaptations that allow them to swim and breathe underwater. However,

most of the species associated with aquatic habitats should be considered semiaquatic for they do not pos sess any traits that allow them to cope with water. Some semiaquatic species occur on or under wet sub strates(damp sand, muck,organic litter, moss close to the ground). Others reside on emergent aquatic plants growing near the shoreline or throughout a body of water (bogs, fens, swamps, fresh and salt water marshes, ponds, lakes, streams). Some can readily dive into water and swim below the surface to feed on

aquatic plants or algae. Additional species live on plants growing in wet soil (damp meadows) or near the edge of open water. Within Orthoptera, Acrididae, Tetrigidae, Tridactylidae,and Anostostomatidae collectively include some of the most unusual aquatic species. Gryllidae and Tettigoniidae include a number of semiaquatic species that prefer to inhabit near the edges of aquatic habitats, but do not directly interact with water. Most of these orthopterans are known from the tropical regions around the world, and there is a relatively small number of species known from North America. Amedegnato and Devriese (2008)conducted a global survey of orthopterans associated with aquatic habi tats and recognized that there are at least 188 species in 50 genera from Acrididae and Tetrigidae alone.

The natural history of many orthopteran species is simply unknown so it is possible that there are more unusual species that have evolved adaptations to aquatic habitats. Within Acrididae, the most unusual aquatic grasshoppers are found in Marelliinae (Marellia remipes (Uvarov, 1929)) and Pauliniinae {Paulinia acuminata (De Geer, 1773)), both of which are monotypic and endemic to South America. These species live on broad,floating leaves ofaquatic plants, feeding and ovipositing on them, and their entire life cycle takes place on these plants. Their hind femora are flat and dilated, which help them swim underwa ter (Carbonell 1957). Cornops aquaticum (Bruner, 1906) (Leptysminae) from the Neotropics and Gesonula punctifrons(Stal, 1861)(Oxyinae)from India and Southeast Asia have convergently evolved to feed on water hyacinth and oviposit endophytically (Amedegnato and Devriese 2008; Capello et al. 2012). Some members of Leptyminae, Copiocerinae, Oxy inae, Hemiacridinae, and Tropidopolinae prefer to feed on reed species in the riparian habitats. Many species within Tetrigidae are limno-terrestrial and capable of swimming, and often found at the margins of rivers and lakes. Within Tetrigidae, the subfamily Scelimeninae can be considered truly aquatic because

they can dive under water to hide and to feed on algae that grow on the underside of boulders(Amedegnato and Devriese 2008). These insects have sharp spines protruding from pronotum, which are presumed to be a defensive structure against predatory fish. The members of Tridactylidae can often be found in the same habitats as Tetrigidae. These insects are very small, and have modifications to the legs for swim ming and walking on the water surface. The most recently discovered aquatic orthopteran belongs to Anostostomatidae, which include king crickets and wetas(common name for these orthopteran species). In 1999, a new genus of cave cricket Hydrolutos (Issa and Jaffe 1999) was discovered in the cave systems of

tepuis (flat table-top mountains) in Venezuela, and

411

412

Chapter 15 Semiaquatic Orthoptera

subsequently a total of seven species have been described. These insects are characterized by having a plastron-like structure on the pleurosternal area of the thorax and abdomen covered with fine microtri-

chia, which presumably holds an air bubble and allows them to be submerged and move about for 20 min (Issa and Jaffe 1999; Derka and Fedor 2010). In North America, many orthopteran species are probably semiaquatic, but ecological studies on whether these species truly prefer to feed on aquatic plants or show clear habitat preferences are lacking. Certainly both tetrigids and tridactylids are associ ated with aquatic habitats, but other examples described in this chapter are based on unpublished observations. The most widely cited and the only empirical study of semiaquatic orthopterans in the United States is by Squitier and Capinera (2002) who examined host plant preference of six grasshopper species commonly encountered in aquatic habitats in Florida. They performed laboratory choice tests involving 19 semiaquatic plant species on two species in Leptyisminae, Stenacris vitreipennis (Marschall, 1836), Leptysma marginicollis (Serville, 1838), three species in Melanoplinae, Gymnoscirtetes pusillus Scudder, 1897, Paroxya clavuliger (Serville, 1838), Paroxya atlantica (Scudder 1877), and one species of Romaleidae, Romalea microptera (Palisot de Beauvois, 1817). They showed that both leptysmines showed a strong preference for aquatic grasses, while other species showed a mixed preference for both grasses and forbs associated with semiaquatic habitats.

GENERAL BIOLOGY

The order Orthoptera is characterized by the presence of a cryptopleuron, developed from the lat eral extension of the pronotum over the pleural sclerites, and jumping hind legs(Kevan 1982). As in other polyneopteran insects, the orthopteran insects are fully winged (although microptery and aptery have evolved multiple times), have chewing mouthparts, and incomplete metamorphosis. The order consists of two suborders, Caelifera and Ensifera. The Caelifera

includes grasshoppers, locusts, and their relatives, and can be characterized by antennae with less than 30 flagellomeres, asymmetrical mandibles each with a heavy molar, mostly exposed thoracic pleura, three or fewer tarsal segments, and abdominal tympana. The Ensifera includes crickets, katydids, wetas, and their relatives, and can be characterized by long and thread like antennae that are usually longer than the body, symmetrical mandibles, thoracic pleura concealed by lateral pronotal lobes, three or four tarsal segments, and tympana often present on the front tibia. Many

orthopterans are capable of producing sound and engage in acoustic communication between males and females. Katydids and crickets have a stridulatory apparatus at the base of the tegmina. To produce sound, the front wings are elevated and the inner edge at the base of one tegmen is rilbbed against a toothed ridge at the base ofthe other tegmen. Grasshoppers in the subfamily Gomphocerinae rub a longitudinal ridge or a series of pegs on the inside of the hind fem ora across a raised vein on the tegmina. Those in the subfamily Oedipodinae can produce snapping sounds during flight, known as crepitation. Eggs are deposited in loose soil, stems, clumps of vegetation, burrows, or on the surface of leaves and twigs, depending on the species. Nymphs develop into adults through the process of incomplete metamor phosis. Most species occurring in temperate regions have a 1-year life cycle, although some in cold cli mates may require up to 3 years to become adults; in contrast, two generations per year may occur in the southern United States. All caeliferans are virtually phytophagous in a broad sense, although specific preferences on different plant types have evolved numerously throughout different lineages within Caelifera. Unlike caeliferans, ensiferans demonstrate

incredible variety in their diet. Most crickets are omnivorous, feeding on detritus, dead insects, and plants. Many basal ensiferans, including the Anostostomatidae, Gryllacrididae, Rhaphidophoridae, and Stenopelmatidae, are scavengers or are predatory on small insects. While many katydids are herbivorous, some groups are predatory and others may feed on flowers, pollen, or nectar. Grasshoppers (family Acrididae and Romalei dae) are characterized by short antennae and three-segmented tarsi; females have short, stout ovi positors. Approximately 26 out of some 630 species in North America north of Mexico are semiaquatic. Only afew species(e.g.,Leptysma marginicollis(Serville, 1838)), Stenacris vitreipennis{y[dirx\\s.\\, 1836), Metaleptea brevicornis {ioh&nnson, 1763), Paroxya clavuligera (Serville, 1838)) occur almost exclusively in wet areas. Habitats for semiaquatic grasshoppers include edges of bogs, fens, swamps, fresh and saltwater marshes, ponds, lakes and streams, as well as muck, wet meadows, peatlands, and tundra. In North America, the members of Acridinae, Gomphoceri nae, and Oedipodinae prefer grasses, whereas other subfamilies feed on a wide variety of herbaceous plants. Most grasshoppers are polyphagous,although many species can be narrowly oligophagous (Chap man and Sword 1997).

Pygmy grasshoppers(family Tetrigidae)are small insects with a distinctive pronotum that extends

Chapter 15 Semiaquatic Orthoptera

backward to the tip of the abdomen or beyond. The tegmina are reduced to small pads but the hind wings are usually long and used for flying. Fourteen of the 27-30 species in North America north of Mexico are associated with wet habitats at ground level. They occur on damp sandy, mucky, muddy, and algaecovered edges of swamps, marshes, ponds, lakes, and streams, on mossy soils, and in damp meadows. Indi viduals may leap into water and swim short distances to submerged objects to evade capture. Tetrigids feed on mosses, algae, decaying organic matters, fungi, and low, succulent seedlings. Pygmy mole crickets (family Tridactylidae) are small (4-10 mm in length) orthopterans with small eyes, prognathous mouthparts, front tibiae modified for digging. Their hind tibiae usually have long, slen der, movable plates for swimming and walking on the water surface, and their hind femora are enlarged for jumping. Five ofthe seven tridactylid species in North America north of Mexico excavate and live in bur

rows along sandy banks of ditches, ponds, lakes, and streams. They feed on organic debris and algae. Katydids(family Tettigoniidae) are characterized by tegmina that are held roof-like over the abdomen,a male subgenital plate with a pair of styles, a sword-like ovipositor in females, and four-segmented tarsi. Stridulation is achieved by rubbing the left tegmen over the right. Generally, katydids are the most commonly seen and heard ofthe orthopterans in semiaquatic environ ments. Most of the 35 semiaquatic species of katydids listed out of approximately 265 species in North Amer ica north of Mexico are more common in the wet, humid eastern half ofthe United States. They occur on vegetation on the edges of bogs, fens, swamps, ponds, lakes, and streams and the edges and interior of fresh and saltwater marshes. All are plant feeders although a few are predaceous on occasion (e.g., some

413

Orchetimum species), and one (Sphagniana sphagnorum (Walker, 1869))is presumed to be primarily preda ceous on other insects. Conocephalus and Orchelimum species are particularly common in and around the edges of marshes. Conocephalus spartinae(Fox, 1912), O. concinnum Scudder, 1862, and 0.fidicinium Rehn and Hebard, 1907 are major consumers of leaves, flowers, and seeds of dominant salt marsh perennials such as Juncus(rushes) and Spartina (grasses). Crickets (family Gryllidae) are characterized by long antennae, a generally quadrate pronotum, teg mina positioned flat across the dorsum, long cerci, and a needle-like ovipositor. In many species, males produce melodic songs by rubbing scrapers on the left tegmen against stridulatory files on the right tegmen. Cricket wings have modified veins that form the mir ror and harp, which function as resonators when stridulation takes place. Virtually all crickets stridu-

late by passing the right tegmen over the left. Crickets are omnivorous scavengers and typically nocturnal. About eleven species out ofapproximately 120 species in North America north of Mexico are semiaquatic. Semiaquatic crickets are found on the edges of bogs, fens, swamps (also under tidal litter of mangrove swamps), on grasses and reeds of fresh and saltwater marshes, and on edges of ponds, lakes, and streams. Mole crickets (family Gryllotalpidae) are charac terized by a small and conical head, legs modified for digging and burrowing, hind legs not modified for jumping, tegmina of males lacking a mirror, and a highly reduced ovipositor in females. They comprise seven species in North America north of Mexico, of which two live primarily in semiaquatic habitats. The

two semiaquatic species among them frequent muck and wet sand on the edges of marshes, ponds, lakes, and streams. They feed on plant materials,insects, and other soil arthropods.

KEY TO FAMILIES OF ORTHOPTERA WITH SEMIAQUATIC SPECIES IN NORTH AMERICA

1. T.

2(1).

Antennae long and thread-like and usually longer than the body (Figs. 15.17-15.20, 15.26, 15.30); tympanum present on the front tibiae Antennae short and robust with less than 30 flagellomeres and usually shorter than the body (Figs. 15.1, 15.10, 15.16); tympanum not present on the front tibiae All tarsi 4-segmented; tegmina held roof-like over the abdomen; a male subgenital plate with a pair of styles; a sword-like ovipositor in females (Figs. 15.17-15.20, 15.23-15.24)(katydids)

2'.

3(2').

2 4

TETTIGONIIDAE

All tarsi 3-segmented; tegmina positioned flat across the dorsum (Figs. 15.26, 15.30, 15.34); a male with long cerci; a needle-like or reduced ovipositor in females (Figs. 15.31-15.33) 3 Head small and conical; front legs modified for digging and burrowing (Fig. 15.34); hind legs not modified for jumping; tegmina of males lacking a mirror, and a highly reduced ovipositor in females(mole crickets) GRYLLOTALPIDAE

414

Chapter 15 Semiaquatic Orthoptera

fastlBlum of vertex

abdomen

pronotum

femur

tegmlna

jvi.iaugMt

subgenltal plate

antenna

tibia! spur

Figure 15.1

ovipositor

tarsus

Figure 15.2

width

length

lobe

Figure 15.5

Figure 15.4 Figure 15.3 spine

foveolae

lateral carinae

Figure 15.6

Figure 15.7

Figure 15.1 Male grasshopper, Leptysma marginicollis (Acrididae). Figure 15.2 Ovipositor of a female grasshopper. Figure 15.3 Prosternal spine between front coxae (Acrididae)(after Capinera et al. 2004). Figure 15.4 Mesosternal lobes longer than wide {Schistocerca: Acrididae)(after Bland 2003). Rgure 15.5 Mesosternal lobes as wide as long (Acrididae)(after Bland 2003).

Figure 15.8

Figure 15.9

Figure 15.6 Lateral foveolae of vertex visible from above (Acrididae). Figure 15.7 Lateral foveolae of vertex not visible from above (Acrididae). Figure 15.8 Lateral carinae of pronotum straight (Dichromorpha: Acrididae)(after Capinera etal. 2004). Figure 15.9 Lateral carinae of pronotum incurved in middle and diverging posteriorly (Orphulella: Acrididae) (after Capinera et al. 2004).

Chapter 15 Semiaquatic Orthoptera

3'.

4(1').

Head not conical; front legs normal; hind legs modified for jumping; tegmina of males with a mirror; a needle-like ovipositor in females (Figs. 15.31-15.33)(crickets) Pronotum extended posteriorly to or beyond tip of abdomen (Fig. 15.10) (pygmy grasshoppers)

GRYLLIDAE TETRIGIDAE

4'.

Pronotum not extended posteriorly

5(4').

Size small (less than 1 cm); prognathous mouthparts; front legs well-modified for digging (Fig. 15.16)(pygmy mole crickets)

5'.

6(5').

5

TRIDACTYLIDAE

Size medium to large(more than 1 cm); hypognathous mouthparts; front legs normal(Fig. 15.1); tympanum present on the lateral sides of first abdominal segment; ovipositor offemale stout, consisting offour short, curved projections at tip of abdomen (Fig. 15.2)

6

External apical spur present on the hind tibiae; often large, sluggish, and colorful(lubber grasshoppers)

6'.

415

ROMALEIDAE

External apical spur present on the hind tibiae; size and color highly variable (grasshoppers)

ACRIDIDAE

KEYS TO GENERA WITH SEMIAQUATIC SPECIES

Tettigoniidae 1.

Front tibiae with three large dorsal spines; tegmina broad and usually short, covering half(males)to one-fourth (females) of abdomen; edges of sphagnum bogs and spruce swamps; southern half Canada from eastern British Columbia to western Quebec; one species, sphagnorum (Walker, 1869)(Fig. 15.17). .. . Sphagniana Zeuner, 1941

r.

Front tibiae without three large dorsal spines, typically spineless dorsally; tegmina broad or narrow, if broad then longer than abdomen

2

2(1').

Prosternal spines between front coxae (cf. Fig. 15.3); tegmina narrow and sometimes do not extend beyond tip of abdomen; head conical on species longer than 24 mm

3

No prosternal spines between front coxae; tegmina broad,long, extend beyond tip of abdomen; head rounded, not conical

6

2'. 3(2).

Body length less than 17 mm excluding ovipositor; fresh and salt water marshes, swamps,edges of ponds, lakes, and streams; eastern half United States, TX,CA,southern Quebec and Ontario; nine species, aigialus Rehn and Hebard, 1915, attenuatus(Scudder, 1869)(Fig. 15.18), brevipennis(Scuddsr, 1862), hygrophilus Rehn and Hebard, 1915, nigropleuroides Fox, 1912, nigropleurum (Bruner, 1891), spartinae(Fox, 1912), spinosus(Morse, 1901), stictomerus Rehn and Hebard, 1915 Conocephalus Thunberg, 1815

3'.

Body length usually 17 mm or longer excluding ovipositor

4(3').

Body length 17-27 mm (rarely less than 17 mm)excluding ovipositor; head without conical projection; fresh and salt water marshes, swamps, edges of ponds, lakes, and streams; United States, southern fifth Canada; 12 species, agile(De Geer, 1773), bradleyi Rehn and Hebard, 1915, bullatum Rehn and Hebard, 1915, campestre Blatchley, 1893, concinnum Scudder, 1862(Fig. 15.19), delicatum Bruner, 1^92,fidicinium Rehn and Hebard, 1907, gladiator Bruner, 1891, militare Rehn and Hebard, 1907, nigripes Scudder, IS15,pulchellum Davis, 1909,

4'.

Body length 27 mm or more excluding ovipositor; head with conical projection (fastigium)(Figs. 15.20-15.22)

vo/unrum McNeill, 1891

4

Orchelimum Serville, 182S

5

416

Chapter 15 Semiaquatic Orthoptera

pronotum

Figure 15.10

fastigium

fastigium

fastlgium -

Figure 15.11

Figure 15.12

Figure 15.13

lateral carlnae of

fastigium

Figure 15.14

Figure 15.15

pronotum

Figure 15.16

Figure 15.10 Female pygmy grasshopper, Tetrix subulata (Tetrlgidae)(after Rehn and Grant 1961). Figure 15.11 Fastigium of vertex slightly extended in front of eyes {Paratettix cucullatus: Tetrlgidae)(after Rehn and Grant 1961). Figure 15.12 Fastigium of vertex greatly extended in front of eyes and broadly arched in profile (Neotettix femoratus: Tetrlgidae)(after Rehn and Grant 1961). Figure 15.13 Fastigium of vertex greatly extended in front of eyes and angular in profile (Tetrix subulata: Tetrlgidae)(after Rehn and Grant 1961),

tarsus

Figure 15.14 Lateral carlnae of frontal costa strongly divergent ventrally (Neotettix femoratus: Tetrlgidae) (after Rehn and Grant 1961). Figure 15.15 Lateral carlnae of frontal costa slightly divergent ventrally (Tetrix subulata: Tetrlgidae)(after Rehn and Grant 1961). Figure 15.16 Male pygmy mole cricket, Eiiipes minutus (Tridactylidae) (after Hebard 1934).

Chapter 15 Semiaquatic Orthoptera

5(4').

417

Fastigium with a broad tooth beneath, large gap between lower face of fastigium and median facial ridge (Fig. 15.21); edges of bogs, fens, fresh and saltwater marshes; eastern two-thirds United States, southern Ontario; six species, caudellianus(Davis, 1905), exiliscanorus(Davis, 1887), lyristes(Rehn and

5'.

Hebard, 1905)(Fig. 15.20), melanorhinus(Rehn and Hebard, 1907), palustris (Blatchley, 1893), retusus(Scudder, 1878) Neoconocephalus Kamy, 1907 Fastigium without a broad tooth beneath, narrow or no gap between lower face of fastigium and median facial ridge (Fig. 15.22); fresh and salt water marshes; Atlantic and Gulf coasts, AR;one species, malivolans

6(2').

(Scudder, 1878) Tegmina distinctly broader in middle; fastigium about twice as wide as

Bucrates Burmeister, 1838

first antennal segment; edges of marshes, swamps, ponds, and lakes; eastern two-thirds United States, southern Quebec; one species, oblongifolia(De Geer, 1773)

6'. 7(6').

(Fig. 15.23) Amblycorypha Stk\, 1873 Tegmina not distinctly broader in middle, tegmina narrow and elongated; fastigium about same width as first antennal segment 7 Tegmina green, strikingly marked with black and brown, sometimes as stripes; cypress swamps; southeastern United States, Gulf Coast states to LA,north to IL; three species, strigata (Scudder, 1898), taxodii Caudell, 1921, walkeri Hebard, 1925

7'. 8(7').

Inscudderia Csiuddl, 1921

Tegmina without striking black and brown markings Male subgenital plate long, upwardly curved (Fig. 15.24); male supra-anal plate (Fig. 15.24) elongate and notched at apex (Fig. 15.25); front and middle femora not spined below; edges of marshes, swamps, and lakes; United States except

8

northwestern and southwestern regions; one species, texensis Saussure and Pictet, 1897

8'.

(Fig. 15.24) ScudderiaSikX, 1873 Male subgenital plate short, broad, not curved upward; male supra-anal plate triangular, without apical notch; front and middle femora strongly spined below; swamps, on water hyacinth; southern United States north to IN and MD;

one species, modesta (Brunner von Wattenwyl, 1878)

Montezumina Hebard, 1925

Gryllotalpidae 1. Front tibiae with two dactyls (blade-like claws or finger-like projections); wet sand or muck on edges of ponds and streams; southern United States; one species, borellii(Giglio-Tos, 1894) Neoscapteriscus Cadena-Castaneda, 2015 1'. Front tibiae with four dactyls(Fig. 15.34); wet sand or muck on edges of marshes, ponds, lakes, and streams; eastern two-thirds United States, southern Ontario; one species, hexadactyla (Perty, 1832)(Fig. 15.34) Neocurtilla Kirby, 1906 Gryllidae* 1. Ventral side of second tarsal segment of hind tarsi with a brush-like pad (Fig. 15.27); edges of fresh and saltwater marshes, mangrove and other swamps,lakes, and streams; eastern two-thirds United States, southern Ontario; four species, delicatula (Scudder, 1878), exigua (Say, 1825)(Fig. 15.26), litarena Fulton, 1956, scia Hebard, 1915 Anaxipha Saussure, 1874 *Receiitly, the subfamily Trigonidiinae, which includes all of the genera covered in this key, was elevated to the family Trigonidiidae. However, this change is not yet widely accepted, and thus we follow the traditional family concept, Gryllidae, here.

418

r. 2(1'). 2'.

3(2').

3'.

Chapter 15 Semiaquatic Orthoptera

Ventral side of second tarsal segment of hind tarsi without a brush-like pad 2 Hind tibiae with three spines on each upper margin; mangrove swamps; south FL coast; one species, alleni(Morse, 1905) Hygronemobius Hebard, 1913 Hind tibiae with four spines on each upper margin 3 Paired disto-ventral spurs of hind tibiae about equal in length (Fig. 15.28); end of ovipositor with coarse teeth dorsally and very fine teeth ventrally (Fig. 15.31); edges of bogs, fens, marshes, mangrove and other swamps, lakes, and streams; United States, southeastern Canada; two species, carolinus (Scudder, 1877), melodius(Thomas and Alexander, 1957) Eunemobius Hebard, 1913 Paired disto-ventral spurs of hind tibiae distinctly unequal in length (Fig. 15.29); end of ovipositor with fine teeth dorsally and no teeth ventrally (Figs. 15.32-15.33)

4(3').

4'.

4

Body length of males less than 9 mm; ovipositor gently curved upward (Fig. 15.32) and not more than two-thirds length of hind femora; edges of bogs (especially sphagnum bogs), fens, and mangrove and other swamps; eastern half of United States, southern Canada; two species, cubensis(Saussure, 1874), palustris (Blatchley, 1900) Neonemobius Hebard, 1913 Body length of males usually greater than 9 mm; ovipositor nearly straight (Fig. 15.33), at least three-fourths length of hind femora; edges of bogs, fresh and saltwater marshes, mangrove and other swamps, ponds, lakes, and streams; northern half United States, Midwest south to TX,

southern third Canada; 2 species,fasciatus(De Geer, 1773)(Fig. 15.30), spawa/.sM.s(Fulton, 1930)

.

Allonemobius Hehard, 1913

Tetrigidae

1.

Less than 15 antennal segments; front femora with distinct dorso-longitudinal ridge

2

r.

More than 15 antennal segments; front femora with a shallow, broad, dorso-longitudinal groove

4

2(1).

2'. 3(2').

3'.

4(1').

4'.

Fastigium of vertex slightly or not extended in front of eyes in profile (Fig. 15.11); edges of coastal marshes, swamps, ponds, lakes, and streams, on mats of algae; United States, southern Ontario; four species, aztecus(Saussure, 1861), cucullatus (Burmeister, 1838), (Saussure, 1861), rwgoiMi(Scudder, 1862) . .Paratettix Bolivar, 1887 Fastigium of vertex greatly extended in front of eyes in profile (Figs. 15.12-15.13) 3 Frontal costa with lateral carinae strongly divergent ventrally (Fig. 15.14); fastigio-facial angle distinctly and broadly arched in profile (Fig. 15.12); edges of salt marshes and swamps; NY to southeastern United States, west to IN and TX; one species,femoratus(Scudder, 1869) Neotettix Hancock, 1898 Frontal costa with lateral carinae only slightly divergent ventrally (Fig. 15.15); fastigio-facial angle angular or weakly rounded in profile (Fig. 15.13); edges of bogs, marshes, ponds, and streams; Alaska, Canada, United States; three species, arenosa Burmeister, 1838, ornata (Say, 1824), subulata(Limaaeus, 1758)(Fig. 15.10) TernA Latreille, 1802

Body distinctly swollen; front half of pronotum moderately arched in profile; wet meadows and woods, edges of ponds; southeastern United States west to LA; one species, obesa (Scudder, 1877) Paxilla Bolivar, 1887 Body weakly swollen, relatively slender; front half of pronotum not arched in profile; edges of bogs, fresh and salt water marshes, swamps, wet woods, and ponds; eastern two-thirds United States, southeastern Canada; four species, acuta Morse, 1895, armata Morse, 1895, lateralis(Say, 1824), prorsa Scudder, 1877 Tettigidea Scudder, 1862

^

Chapter 15 Semiaquatic Orthoptera

419

Figure 15.17

Figure 15.18

ovipositor

Figure 15.19

Figure 15.17 Male katydid, Sphagniana sphagnorum (Tettigoniidae)(after Heifer 1987). Figure 15.18 Male katydid, Conocephalus attenuatus (Tettigoniidae).

Figure 15.19 Male katydid, Orchelimum concinnum, and ovipositor of a female (Tettigoniidae).

420

Chapter 15 Semiaquatic Orthoptera

fastigium

tooth

fastigium

Figure 15.21 Figure 15.20 ovipositor

Figure 15.22

Figure 15.23

supra-anai piate

Figure 15.25 notch

subgenital

Figure 15.24

ovipositor

Figure 15.20 Male katydid, Neoconocephalus lyristes, and ovipositor of a female (Tettlgonlldae) (modified after VIckery and Kevan 1986). Figure 15.21 Fastigium with a broad tooth and wide gap between lower face and median facial ridge (Neoconocephalus: Tettlgonlldae)(after Caplnera et al. 2004). Figure 15.22 Fastigium without a broad tooth and with a narrow gap or no gap between lower face and median facial ridge (Bucrates: Tettlgonlldae)(modified after Caplnera etal. 2004).

Figure 15.23 Male katydid, Amblycorypha oblongifolia (Tettlgonlldae)(after VIckery and Kevan 1986). Figure 15.24 Male katydid, Scudderia texensis, and ovipositor of a female (Tettlgonlldae)(modified after Bland 2003). Figure 15.25 Male supra-anal plate (dorsal view) with broad apical notch (Scudderia texensis: Tettlgonlldae) (after Bland 2003).

Chapter 15 Semiaquatic Orthoptera

421

Tridactylidae 1. Body length usually less than 5.5 mm; prosternum without a conical protuberance; tarsus of hind leg absent; wet, sandy banks of ponds, lakes, and streams; eastern two-thirds United States, southwestern United States, CA,southern Quebec, Ontario, and Manitoba; four species, gurneyi Giinther, 1977, minimus Bruner, 1916, minuta (Scudder, 1862)(Fig. 15.16), monticolus Giinther, 1977 Ellipes Scudder, 1902

r.

Body length usually more than 5.5 mm; prosternum with a conical protuberance; tarsus of hind leg present; habitats same as Ellipes; eastern half United States, southwestern United States, southern Quebec, Ontario, and Manitoba; 1 species,

apicialis(Sny, 1825)

Neotridactylus Giinther, 1972

Romateidae

1.

Hind tibiae with immovable apical spine on outside surface; short hind wings are pinkish red; large, stout grasshopper; color varying from orange yellow to black, edges of marshes and ponds, on water hyacinth; southeastern United States;

one species, microptera (Palisot de Beauvois, 1817)

Romalea Serville, 1831

Acrididae

1.

Prosternum with a prominent cylindrical spine (prosternal process) between

front coxae (Fig. 15.3)

2

r.

Prosternum without a prominent cylindrical spine between front coxae

2(1).

Male cerci vertically hooked; lower external lobe of the hind knee angular; second tarsal segment of the hind legs very short; face strongly angled backwards in profile (Fig. 15.1)

3

2'.

Male cerci triangular, quadrate, or variable shape, but not vertically hooked; lower external lobe of the hind knee round; second tarsal segment of the hind legs not short; face nearly vertical or slightly angled backwards in profile

4

3(2).

10

Head as long as or longer than pronotum; fastigium of vertex (Fig. 15.1) with a deep median groove; edges of bogs, marshes, ponds, lakes, and streams; southern half United States southwest to CA;one species, marginicollis

(Serville, 1838)(Fig. 15.1)

Leptysma StM, 1873

3'.

Head shorter than pronotum; fastigium of vertex without a median groove; edges of bogs, marshes, ponds, lakes, and streams; southeast and Gulf Coast states; one species, vitreipennis (Marschall, 1836) Stenacris Walker, 1870

4(2').

Mesosternal lobes longer than wide with rectangular inner angle (Fig. 15.4);

edges of bogs, fresh and saltwater marshes, ponds, and lakes; eastern half United States; one species, alutacea (Harris, 1841)

Schistocerca Stal, 1873

4'.

Mesosternal lobes as wide as or wider than their length with round inner angle (Fig. 15.5)

5

5(4'). 5'.

Tegmina reduced to small pads or completely absent Tegmina and hind wings fully developed

6 9

6(5). 6'. 7(6).

Tegmina reduced to slender pads or round pads 7 Tegmina and hind wings completely absent 8 Tegmina modified as slender pads; green, with two white stripes running laterally from head to thorax; edges of fresh and salt water marshes; southern U.nited States; one species, sphenarioides Scudder, 1878 Aptenopedes Scudder, 1878 Tegmina modified as round or elongate-oval pads; pronotum tectiform; edges of marshes; eastern half United States; two species, palustris Morse, 1904, s/gnalMi Scudder, 1897 Eotettix Scudder, 1897

7'.

422

Chapter 15 Semiaquatic Orthoptera

dactyls

Figure 15.26

Figure 15.30

2nd tarsal segment

Figure 15.34

Figure 15.27

coarse teeth

tibia

Figure 15.31 fine teeth

Figure 15.28

distoventrai spurs equal

Figure 15.32 fine teeth

Figure 15.29 distoventrai

spurs unequal

Figure 15.33

Figure 15.26 Male cricket, Anaxipha exigua (Gryliidae)(after Froeschner 1954). Figure 15.27 Brush-like pad on ventral side of second tarsal segment(Anaxipha: Gryliidae)(after Bland 2003). Figure 15.28 Paired distoventrai spurs of hind tibiae nearly equal in length (Eunemobius: Gryliidae)(after Bland 2003). Figure 15.29 Paired distoventrai spurs of hind tibiae distinctly unequal in length (Neonemoblus, Allonemobius: Gryliidae)(after Bland 2003). Figure 15.30 Male cricket, Allonemobius fasclatus (Gryliidae).

Figure 15.31 Short, slightly curved cricket ovipositor with coarse dorsai teeth apically (Eunemobius: Gryliidae)(modified after Vickery and Kevan 1986). Figure 15.32 Short, slightly curved cricket ovipositor with fine dorsal teeth apically (Neonemoblus: Gryliidae) (after Vickery and Kevan 1986). Figure 15.33 Long, nearly straight cricket ovipositor with fine dorsal teeth apically (Allonemobius: Gryliidae) (after Vickery and Kevan 1986). Figure 15.34 Male mole cricket, Neocurtllla hexadactyla (Gryllotalpidae).

Chapter 15 Semiaquatic Orthoptera

423

8(6').

Brownish green body, dark ivory patch on side of pronotum; bogs, fens; North Central states, southern Ontario; two species, glacialis(Scudder, 1862), variegata (Scudder, 1897) Boomacris Rehn and Randell, 1962

8'.

Greenish or brownish yellow or gray body,longitudinal dull black stripe on side of body; edges of bogs and marshes; southern U.S.; two species, wor.se/Hebard, 1918,pms///ms Scudder, 1897 Gymnoscirtetes Scudder, 1897

9(5').

Dorsal surface of pronotum twice as long as average width; antennae much longer than head and pronotum combined; edges of bogs, fresh and saltwater marshes, ponds, and lakes; eastern third United States, southern Ontario; three species, atlantica Scudder, 1877, clavuligera (Serville, 1838), hoosieri(Blatchley, 1892) Paroxya Scudder, 1877

9'.

Dorsal surface of pronotum less than twice as long as average width; antennae as long as or shorter than head and pronotum combined; tundra, peatlands, edges of bogs and streams; Alaska, Canada, northern third United States, southwestern United States; two species, borealis(Fieber, 1853), herbaceus Bruner, 1893 Melanoplus Stdl, 1873

10(1'). 10',

Lateral foveolae or foveolar area of vertex visible from above (Fig. 15.6) Lateral foveolae or foveolar area of vertex not visible from above (Fig. 15.7)

11(10).

Dorsum of pronotum without dark lateral markings posteriorly; tegmina reach much beyond tip of abdomen; underside of hind femora usually reddish, sometimes yellowish; edges of bogs, marshes, swamps, wet meadows,lakes, and streams; Alaska, Canada, northern half United States except Pacific Coast states,

11 12

CO,OK,southeastern U.nited States; three species, celatum Otte, 1979, gracile (Scudder, 1862),

//nea/MW (Scudder, 1862)

StethophymaP{?,chex, 1853

11'.

Dorsum of pronotum with dark lateral markings posteriorly; tegmina often short, extending about two-thirds of distance to tip of abdomen but sometimes beyond abdomen; underside of femora pale brown or yellowish; tundra, wet meadows, edges of bogs,fens, marshes, and lakes; N. America except southernmost areas; one species, curtipennis(Harris, 1835) Pseudochorthippus Defaut, 2012

12(10').

Antennae flattened, sword-shaped (ensiform), basal third slightly to greatly widened

13

12'.

Antennae threadlike (filiform), basal third may be slightly flattened but not wider than distal segments

15

13(12).

Head longer than pronotum; front of face strongly slanted backward and concave in profile; tegmina shorter than abdomen; body very slender; wet meadows, edges of fresh and salt water marshes and ponds; southeastern United States to FL; one species, carinalum (Walker, 1870) Achurum Saussure, 1861

13'.

Head equal to or shorter than pronotum; face moderately slanted, not concave in profile; tegmina longer than abdomen; body not highly slender

14

14(13'). Tegmina rounded apically; in males, a stridulatory file consisting of a row of pegs along the inside of each hind femur present, dark dorso-longitudinal stripe; wings rounded apically; edge of marshes; coastal region from NJ to FL; one species, intertexta Scudder, 1899 Mermiria StM, 1873 14'. Tegmina nearly square (truncated) apically; in males, a stridulatory file consisting of a row of pegs along the inside of each hind femur absent; wet meadows,edges of fresh and salt water marshes, swamps, ponds, lakes, and streams; eastern half United States, southern Ontario; one species,

15(12').

brev/com/.s(Johannson, 1763) MetalepteaBrmmv won Wattenwyl, 1893 Lateral carinae of pronotum straight and nearly parallel in dorsal view (Fig.15.8); edges of marshes, swamps, ponds and lakes; eastern half U.S. southwest to NM;two species, elegans(Morse, 1896), viridis (Scudder, 1862) .... Dichromorpha Morse, 1896

424

15'.

Chapter 15 Semiaquatic Orthoptera

Lateral carinae of pronotum distinctly incurved in middle and diverge posteriorly in dorsal view (Fig. 15.9); wet meadows, muck,fresh and salt water marshes; United States, southern Canada; one species, (Burmeister, 1838) ... Orphulella Giglio-Tos, 1894

General Blatchley (1920); Capinera et al. (2004); Rehn and Grant(1961); Triplehorn and Johnson (2005).

Gryllotalpidae: Bland (2003); Blatchley (1920); Capinera et al. (2004); Otte et al. (2001); Vickery and Kevan (1986); Walker and Moore (2005). Tetrigidae: Bland (2003); Blatchley (1920); Capinera et al.(2004); Dakin and Hays (1970); Heifer (1987); Otte et al.(2001);

Taxonomic treatments at the family and generic levels

Tettigoniidae: Bland (2003); Blatchley (1920); Capinera et al. (2004); Dakin and Hays (1970); Hebard (1925); Heifer (1987); McCafferty and Sein (1976); Otte et al.(2001);

ADDITIONAL TAXONOMIC REFERENCES

Rehn and Grant (1961); Strohecker et al.(1968).

Acrididae; Bland (2003); Blatchley (1920); Capinera et al.(2004); Dakin and Hays (1970); Heifer (1987); Otte (1981); Otte et al.(2001); Rehn and Eades(1961); Strohecker et al. (1968); Vickery and Kevan (1986). Gryllidae: Bland (2003); Blatchley (1920); Capinera et al.(2004); Dakin and Hays(1970); Fulton (1956); Otte et al.(2001); Vickery and Johnstone (1970); Vickery and Kevan (1986); Walker and Moore (2005).

Rehn and Hebard (1915a,b,c); Thomas and Alexander (1962); Vickery and Kevan (1986); Walker(1971); Walker and Moore (2005). Tridactylidae: Bland (2003); Blatchley (1920); Capinera et al. (2004); Dakin and Hays(1970); Gunther (1975); Heifer (1987); Otte et al. (2001); Vickery and Kevan (1986).

Ki

C/l

Family Genus

Ecological

ORTHOPTERA

"Emphasis on trophic relationships

Melanoplinae

Leptysminae

Aptenopedes(1)

Stenacris(1)

Southern U.S.

states

Southeast and Gulf Coast

to California

U.S., southern Canada Southern half U.S. southwest

Orphulella (1)

Florida

Coastal region New Jersey to

Leptysma (1)

Mermiria (1)

Eastern half U.S. southwest to New Mexico

(continued)

4897, 5028

4897

4478

4478

4478

Chorthippus(^) Dichromorpha (2)

4478

N. America except southernmost areas

4478

Southeastern U.S. to Florida

Achurum (1)

4478

4478

535, 525, 532, 885, 1317, 3734, 5797, 6177, 6050, 2515, 1993, 4478, 4479, 2491

References**

Gomphocerinae

states

half U.S. except Pacific Coast

Alaska, Canada, northern

Ontario

Eastern half U.S., southern

Distribution

North American

Eastern half U.S.

Stethophyma (3)

Metaleptea (1)

(chewers)

Shredders—herbivores

Trophic Relationships

Cyrtacanthacridinae Schistocerca (1)

Acridinae

climbers

hydrophytes(emergent zone, margins)

Grasshoppers

Skaters; "swimmers"

Lentic—vascular

Habit

Acrididae (26)-

Habitat

Short-Florned

Orthoptera - Grasshoppers, Locusts and Crickets

Order

of species In parentheses)

(number

Taxa

Table 15A Summary of ecological and distributional data for semiaquatic Orthoptera (grasshoppers, crickets, etc.)(For definition of terms see Tables 6A-6C; table prepared by R. G. Bland, K. W. Cummins, R. W. Merritt, and M. B. Berg.)

Romalea (1)

Romaleinae

Paroxya (3)

Eastern two-thirds U.S.,

Texas

U.S., west to Indiana and

New York to southeastern

southeastern Canada

Ecological

322

2491

5797, 6177, 6050, 2515, 1993, 4479,

535, 4898, 525, 532, 885, 1317,

123

123

References*

1 ) ) > ))) ) 1 )

Louisiana

U.S., southern Ontario

Generally shredders— herbivores; collectors— gatherers

Southeastern U.S. west to

Sprawlers

Paxilla (1)

Neotettix

Tetriginae

Lentic—vascular hydrophytes (emergent zone, margins)

Paratettix(4)

Tettigidea (4)

Batrachidelnae

Pygmy Grasshoppers

Southeastern U.S.

southern Ontario

North Central states,

Ontario

Eastern third U.S., southern

southwestern U.S.

northern third and

Alaska, Canada,

Melanoplus(2)

Distribution

North American

Eastern half U.S.

Booneacris(2)

Tetrigidae (14) -

Trophic

Relationships Southern U.S.

Habit

Gymnoscirtetes(2)

Habitat

Eotettix(2)

Genus

Podisminae

Family

Continued

"Emphasis on trophic relationships

Order

of species in parentheses)

(number

Taxa

Table 15A

1 ) } } ^ ) ) ) ) J )) 1

OS

20; ventral comb of lacinia with 50 fine intercalary surface hairs, femur with a few long dorsal fringe hairs (Fig. 16.83); abdominal terga with long, thich marginal hairs medially (Fig. 16.81)

41'.

40

42

Femur and tibia of foreleg with >20 fine intercalary surface hairs, femur usually laching long dorsal fringe hairs (Fig. 16.84); abdominal terga laching long, thich marginal hairs medially (Fig. 16.82); mostly Western North America Capnia Pictet, sensu lato, Arsapnia Banhs, Siermcapnia Bottorff and Baumann

* Broome et at. 2019. Illiesia 15: 1-26.

Figure 16.65

Figure 16.67

16.68

Figure 16.71

Figure 16 Figure 16.70

I \ Figure 16.72

Figure 16.73 Figure 16.74

Figure 16.75

Figure 16.76

Figure 16.65 Nanonemoura wahkeena (Nemouridae)

Figure 16.71 Paranemoura perfeota (Nemouridae)

nymphal right foreleg.

Figure 16.66 Ostrocerca sp.(Nemouridae) nymphal

nymphal right foreleg. Figure 16.72 Lednia tumana (Nemouridae) nymphal

head and pronotum, dorsal.

female terminalia, ventral.

Figure 16.67 Podmosta sp.(Nemouridae) nymphal

Figure 16.73 Isocapnia Integra (Capniidae) nymphal right cercus, lateral. Figure 16.74 Nemocapnia Carolina (Capniidae) nymphal right cercus, lateral. Figure 16.75 Bolshecapnia spencerl(Capniidae)

head and pronotum. Figure 16.68 Podmosta sp.(Nemouridae) nymphal right foreleg. Figure 16.69 Prostoia sp.(Nemouridae) nymphal right foreleg.

Figure 16.70 Shipsa rotunda (Nemouridae) nymphal right foreleg.

nymphal right cereal segments.

Figure 16.76 Paracapnia angulata (Capniidae) nymphal terminalia, lateral. 447

448

42(41').

42'.

Chapter 16 Plecoptera

Ventral comb of lacinia with about 10 long and 3 short stout teeth (Fig. 16.85); abdominal sterna clothed with stout hairs interspersed with fine hairs(Fig. 16.86); mostly Western North America Mesocapnia Rauser Ventral comb of lacinia with 13-16 teeth of gradually diminishing length toward base (Fig. 16.80); abdominal sterna clothed only with fine surface hairs (Fig. 16.87); Western North America, one

43(6').

43'.

species, Northeastern North America Utacapnia Gaufin Body robust, its length 8X width (Fig. 16.12); abdomen nearly naked (Fig. 16.90) or with variable coverage of short(Fig. 16.91) or long curved setae (Fig. 16.92); abdominal terga with or without a posterior setal fringe; male paraprocts and female 8th abdominal sternum unmodified

44

44(43'). 44'.

Abdominal terga with a posterior fringe of short or long setae (Figs. 16.91 and 16.92) Abdominal terga without a posterior fringe of setae (Fig. 16.90)

45(44).

Abdominal terga with a posterior fringe of short setae, and its last few segments with 2-4 long

45'.

setae (Fig. 16.91); abdominal segments 1-4 divided ventrolaterally by a membranous pleural fold; Eastern North America Lewcfra Stephens Entire body clothed with long, curved hairs(Eigs. 16.92 and 16.93); abdominal segments 1-6 divided ventrolaterally by a membranous pleural fold; Western North America

46(44').

45 46

Moselia Ricker

Tuft of long setae on corners of pronotum (Eig. 16.94); mesosternal Y-stem widely double and with median longitudinal suture (Fig. 16.95); abdominal segments 1-6 divided ventrolaterally by a membranous pleural fold; widely distributed

Pamleuctm Hanson

46'.

Pronotum without long marginal or corner setae; mesosternal Y-stem single (or narrowly double in the rare Western Pomoleuctra) and without a median longitudinal suture (Fig. 16.96); abdominal

47(46').

Paraprocts bare and appearing fused or touching medially for entire length (Fig. 16.97); apical circlet hairs of cereal segments less than half length of segments(Fig. 16.98); abdominal

segments 1-5, 1-6, or 1-7 divided ventrolaterally by membranous pleural fold

47

segments 1-7 divided ventrolaterally by membranous pleural fold; Western

North America

47'.

Perlomyia Banks

Paraprocts with short or long apical bristles or setae; apical circlet hairs of cereal segments longer than half length of segments(Fig. 16.99); abdominal segments 1-5, 1-6 or 1-7 divided ventrolaterally by membranous pleural fold

48(47').

48

Paraprocts fused and clothed with short setae (Fig. 16.100); pronotum with scattered short surface setae; abdominal segments 1-6 or 1-7 divided by ventrolateral membranous pleural fold; Eastern North America

48'.

Zealeuctm Ricker

Paraprocts unfused, with 2 or more long apical bristles; pronotum glabrous without surface or marginal setae (Fig. 16.101) or with only sparse anterior marginal hairs; abdominal segments 1-5 or 1-7 divided by ventrolateral membranous pleural fold

49(48').

49

Apical circlet hairs of middle cereal segments about half length of segments and directed caudally (Fig. 16.102); abdominal segments 1-7 divided by ventrolateral membranous pleural fold; CA, rare

49'. 50(49').

50'.

laterally (Fig. 16.103); abdominal segments 1-5 divided by pleural fold 50 Pronotum with sparse long setae on anterior margin (Fig. 16.101); mesosternal Y-stem undivided (Fig. 16.96); terminal 4 cereal segments with single, short apical hair (Fig. 16.103); Western North America Despaxia Ricker (one species, D. augusta (Banks)) Pronotum without long setae; Y-stem narrowly divided (Fig. 16.104); Western North America; uncommon

51(7).

Calileuctra Shepard and Baumann

Apical circlet hairs of middle cereal segments long, half or more length of segments and directed

Pomoleuctra Stark and Kyzar

Occiput with transverse row of regularly spaced spinules(Fig. 16.105) or distinctly elevated ridge

52

Figure 16.78

Figure 16.77

Figure 16.79

/ ,\

{ IM

I I I ! 1/ ,1 . ,1

,, ,!!'', h,!i, c'I" i'

rmrrrtrfx^

fiUl'lN

Figure 16.80

Figure 16.83

Figure 16.81

!

\y

Figure 16.82

Figure 16.84

7,4

'"Vxii'V

'/Ivff'r, , \

I' »

yi^ kmm i'

Hium Figure 16.85

Figure 16.87 Figure 16.86

Figure 16.77 Bolshecapnia spenceri(Capniidae) nymphal mesosternum. Figure 16.78 Capnura venosa (Capniidae) nymphal

Figure 16.83 Mesocapnia frisom (Capniidae) nymphal right foreleg. Figure 16.84 Capnia vernalis (Capniidae) nymphal

mesosternum.

right foreleg.

Figure 16.79 Eucapnopsis brevicauda (Capniidae)

Figure 16.85 Mesocapnia frisoni(Capniidae) nymphal right lacinia, ventral. Figure 16.86 Mesocapnia frisoni (Capniidae) nymphal terminalia, ventral. Figure 16.87 Utacapnia lemoniana (Capniidae) nymphal terminalia, ventral.

nymphal right lacinia, ventral.

Figure 16.80 Utacapnia lemoniana (Capniidae) nymphal right lacinia, ventral. Figure 16.81 Mesocapnia frisoni(Capniidae) nymphal terminalia, dorsal. Figure 16.82 Capnia vernalis (Capniidae) nymphal terminalia, dorsal.

449

Jf \

Figure 16.88

Figure 16.89

Figure 16.91

Figure 16.90

■

i

Figure 16.95

^

Figure 16.93

Figure 16.94

Figure 16.92

Figure 16.96

Figure 16.100 Figure 16.97 Figure 16.98

Figure 16.88 Megaleuctra kincaidi(Leuctridae)

Figure 16.99

Figure 16.95 Paraieuctra occidentaiis (Leuctridae)

nymphal habitus.

nymphal mesosternum.

Figure 16.89 Megaleuctra kincaidi(Leuctridae)

Figure 16.96 Despaxia augusta (Leuctridae) nymphal

nymphal termlnalla, ventral.

mesosternum.

Figure 16.90 Paraieuctra occidentaiis (Leuctridae)

Figure 16.97 Periomyia utahensis (Leuctridae)

nymphal termlnalla, dorsal.

nymphal termlnalla, ventral.

Figure 16.91

Figure 16.98 Periomyia utahensis (Leuctridae) nymphal right cercus, lateral. Figure 16.99 Zeaieutra ciaassenia (Leuctridae) nymphal right cercus, lateral. Figure 16.100 Zeaieuctra ciaasseni(Leuctridae) nymphal termlnalla, ventral.

Leuctra sp.(Leuctridae) nymphal

termlnalla, dorsal.

Figure 16.92 Moseiia infuscata (Leuctridae) nymphal termlnalla, dorsal.

Figure 16.93 Moseiia infuscata (Leuctridae) nymphal head and pronotum, dorsal. Figure 16.94 Paraieuctra occidentaiis (Leuctridae) nymphal head and pronotum, dorsal. 450

Chapter 16 Plecoptera

51'.

451

52(51).

Occiput without spinules, except possibly laterally near the eyes (Figs. 16.106 and 16.107), or with a sinuate, irregularly spaced spinule row (Fig. 16.108) 55 Two ocelli (Fig. 16.109) Neoperla Needham

52'.

Three ocelli

53(52').

Abdominal terga with more than 5 intercalary bristles (Fig. 16.110); Western North America Clmssenia Wu(one species, C. sabulosa (Banks)) Abdominal terga with no more than 4 intercalary bristles; Eastern North America 54 Posterior spinule fringe of abdominal sternum 7 complete (Fig. 16.111); cerci without a long setal fringe Agnetina Klapalek Posterior spinule fringe of abdominal sternum 7 incomplete (Fig. 16.112); cerci with at least a few long silky setae Paragnetina Klapalek Occipital spinules in a sinuate, irregularly spaced row, more or less complete behind posterior ocelli (Figs. 16.108 and 16.113) 56 No distinct occipital spinule row (Fig. 16.107) of a few scattered spinules may be present near the postocular setal fringe (Fig. 16.106) 60

53'. 54(53'). 54'. 55(51). 55'. 56(55). 56'. 57(56').

57'.

58(57'). 58'. 59(58').

Ab terga with 5 intercalary bristles 57 Pronotum laterally fringed with a complete, close-set row of long setae (Fig. 16.114); posterior fringe of Ab terga with numerous long setae whose length is three-fourths or more the length of abdominal segments; Eastern North America, uncommon Attaneuria Ricker (one species, A. ruralis(Hagen)) Pronotum fringed laterally with short setae, not so closely set (Fig. 16.113); posterior fringe of abdominal terga mostly of short setae whose length is about one-fourth the length of Ab segments 58 Cerci without a dorsal fringe of long silky setae; abdomen of most species speckled (dark pigment at bases of intercalary setae); primarily Eastern North America, common Perlesta Banks Cerci with prominent dorsal fringe of long silky setae (Fig. 16.115); abdominal terga not speckled 59 Dorsum of thorax and abdomen with a mesal, longitudinal row long, fine, silky setae (Fig. 14.116) (best seen in lateral view); abdominal sternum 7 usually with incomplete posterior fringe; Western North America

59'.

60(55'). 60'. 61(60).

53

Doroneuria Needham and Claassen

No mesal longitudinal row of silky hairs on thorax and abdominal dorsum; abdominal sternum 7 usually with a complete posterior fringe; Western North America Calineuria Ricker(one species, C. californica (Banks)) Postocular fringe reduced to 1-3 long setae (Fig. 16.107); eyes set forward on head; pronotal fringe of 2-3 setae at corners 61 Postocular fringe with a close-set row of several thick spinules(Fig. 16.106); pronotal fringe well developed, consisting of a close-set row of spinules or setae, occasionally incomplete laterally 62 Femora and tibia with dorsal(outer) and ventral(inner)fringes of long silky setae; Eastern North America

Perlinella Banks

61'.

Femora and tibia with only dorsal(outer)fringe of long, silky setae: Eastern North America, rare Hansonoperla Nelson

62(60'). 62'. 63(62').

Two ocelli (Fig. 16.117); lateral pronotal fringe complete; AZ,TX Amcroneuria Klapalek Three ocelli; lateral pronotal fringe incomplete 63 Head with large area of yellow in front of median ocellus (Fig. 16.106); Eastern North America, Appalachian Mts. and foothills Eccoptum Klapalek (one species, E. xanthenes(Newman))

63'.

Head mostly brown (Fig. 16.118), often with yellow M-shaped mark in front of median ocellus

64

Figure 16.102

Figure 16.104

Figure 16.103

Figure 16.101

transverse

of spinufes

Figure 16.107 Figure 16.106

Figure 16.105

Figure 16.109

Figure 16.113

Figure 16.108

' T

T_f

1 p

^

a

..♦ ♦♦ ♦ J

3

Figure 16.110

Figure 18.112

Figure 16.101 Despaxia augusta (Leuctridae) nymphal head and pronotum, dorsal. Figure 16.102 Calileuctra ephemera (Leuctridae) nymphal right cercus, basal and distal segments, lateral. Modified from Stewart etal.(2013). Figure 16.103 Despaxia augusta (Leuctridae) nymphal right cercus, basal, medial, distal segments, lateral.

Figure 16.104 Pomoieuctra andersoni(Leuctridae) nymphal mesosternum. Figure 16.105 Claassenia sabulosa (Perlldae) nymphal head and pronotum, dorsal. Figure 16.106 Eccoptura xanthenes (Perlldae) nymphal head and pronotum, dorsal. 452

Figure 16.111

Figure 16.107 Perlinella drymo (Perlldae) nymphal head and pronotum, dorsal. Figure 16.108 Hesperoperia pacifica (Perlldae) nymphal head and pronotum, dorsal. Figure 16.109 Neoperia ciymene (Perlldae) nymphal head and pronotum, dorsal. Figure 16.110 Claassenia sabuiosa (Perlldae) nymphal mid-abdomlnal tergum, dorsal. Figure 16.111 Agnetina capitata (Perlldae) nymphal termlnalla, ventral.

Figure 16.112 Paragnetina fumosa (Perlldae) nymphal termlnalla, ventral. Figure 16.113 Periesta sp.(Perlldae) nymphal head and pronotum, dorsal.

Chapter 16 Plecoptera

64(63').

64'. 65(8).

Cerci with fringe of long, silky setae, sometimes reduced, but at least on basal segments (Fig. 16.119); pronotal flange wider at posterior angles than along lateral margins; widespread Acroneuria Pictet Cerci without basal fringe of silky setae; pronotal flange narrow throughout; Eastern North America, Southern Appalachians or Piedmont Belonemia Needham and Claassen Eyes set far forward and posterolateral corners of head form nearly a right angle (Fig. 16.120, several Eastern North American Alloperla species approach similar compound eye location but are much smaller); body length of mature nymph 18-25 mm (except Utaperla which is smaller)

65'.

66(65).

66

Eyes set midlaterally and posterolateral corners of head convex (Fig. 16.121); body length of mature nymph 12 mm or less (except some Sweltsa which are larger); apical hairs of cereal segments directed at posterior angles (Fig. 16.122) Epicranial suture truncate (Fig. 16.123); body length of mature nymph 6-8 mm; apical hairs of cereal segments perpendicular to cercus(Fig. 16.124) and marginal pronotal hairs long (Fig. 16.123); mostly Western North America, but U. gaspesiana Harper and Roy, Eastern North America, rare

66'. 67(66'). 67'. 68(65'). 68'. 69(68'). 69'.

70(69).

Epicranial suture Y-shaped (Fig. 16.120); body length of mature nymph 16-25 mm 67 Body length of mature nymph 20-25 mm; head longer than wide (Fig. 16.120); lacinia semiquadrate (Fig. 16.125); Western North America KathroperlaBmks Body length of mature nymph 18-20 mm;head about as long as wide; lacinia subtriangular (Fig. 16.126); Western North America Paraperla Banks Cerci usually with a vertical fringe of intrasegmental hairs (Fig. 16.122); pronotal setae largely restricted to corners; widely distributed Alloperla Banks Cerci without vertical fringe of intrasegmental hairs (Fig. 16.131); pronotum with variable marginal setation, but always with setae on anterior and/or posterior margin (Fig. 16.139).... 69 Thick,depressed dark clothing hairs present on thoracic sterna,inside coxae(Fig. 16.127) 70 Thick, depressed dark hairs absent inside coxae, hairs if present at this location fine, erect and usually light colored (Fig. 16.128) 71 Mesosternal Y-arms well developed, mesosternum usually without erect bristles lateral to clothing hair patch (Fig. 16.127); silky setae well developed on tibial fringes(Fig. 16.129); widely

71'. 72(71'). 72'. 73(72). 73'.

Sasqiiaperla Stark and Baumann (one species,

S. hoopa Stark and Baumann) Longest apical hairs of distal cereal segments shorter than their following segment

(Fig. 16.131) Suwallia Ricker and Neaviperla Ricker Longest apical hairs of distal cereal segments as long or longer than their following segment (Fig. 16.133, 16.134) 72 Distal cereal segments with only a single dorsal and ventral long apical hair in addition to short circlet hairs (Fig. 16.133) 73 Distal cereal segments with more than two long apical circlet hairs in addition to short and medium circlet hairs (Fig. 16.134) 74 Fringe of long marginal hairs complete around pronotum (Fig. 16.135); abdomen pigmented with dark striped or checkered pattern in mature nymphs; Western North America Triznaka Ricker Fringe of long marginal hairs of pronotum with a lateral gap (Fig. 16.136); abdomen concolorous; Western North America

74(72').

Sweltsa Ricker

Mesosternal Y-arms poorly developed, mesosternum with a few erect bristles lateral to clothing hair patch (Fig. 16.130); silky setae sparse on tibial fringes

(Fig. 16.132); CA 71(69').

68

Utaperla

distributed

70'.

453

Plumiperla Surdick

Twelve or fewer long marginal pronotal setae; mature nymph with ocelli enclosing a dark pigment patch (Fig. 16.137); lacinia with a close-set comb of setae just below apical tooth (Fig. 16.138); Baja California Norte, CA,OR,uncommon Bisancora Surdick

fine hair

fringe

Figure 16.115

Figure 16.119

Figure 16.122

Figure 16.114

Figure 16.124

Figure 16.117

Figure 16.116

Figure 16.118

Figure 16.121

Figure 16.120

Figure 16.114 Attaneuria ruralis (Perlidae) nymphal head and pronotum, dorsal.

Figure 16.115 Doroneuria baumanni(Perlidae) nymphal right cercus, basal, medial, and distal segments, lateral. Figure 16.116 Doroneuria baumanni(Perlidae) nymphal terminalia, dorsal. Figure 16.117 Anacroneuria sp. (Perlidae) nymphal head and pronotum, dorsal.

Figure 16.118 Beloneuria georgiana (Perlidae) nymphal head and pronotum, dorsal. Figure 16.119 Acroneuria arenosa (Perlidae) nymphal right cercus, basal, medial, and distal segments, lateral. 454

Figure 16.123

Figure 16.120 Kathroperia perdita (Chloroperlidae) nymphal head and pronotum, dorsal. Figure 16.121 Alloperia imbecilla (Chloroperlidae) nymphal head and pronotum, dorsal. Figure 16.122 Alloperia Imbecilla (Chloroperlidae) nymphal right cercus, lateral. Figure 16.123 Utaperia sopladora (Chloroperlidae) nymphal head and pronotum, dorsal. Figure 16.124 Utaperia sopladora (Chloroperlidae) nymphal right cercus, lateral.

Chapter 16 Plecoptera

74'. 75(74').

75'.

455

More than 20 long marginal pronotal setae (Figs. 16.139 and 16.140); head with variable pattern; lacinia without close-set comb of setae just below apical tooth (Fig. 16.141) 75 Abdomen patterned with dark spots forming 4 incomplete longitudinal stripes in mature nymphs; lacinial basal width about 1/3 length of outer margin (Fig. 16.141); Eastern North America, rare Rasvena Richer Abdomen concolorous; lacinial basal width about 1/2 length of outer margin (Fig. 16.142)

76

76(75').

Pronotal fringe hairs long,0.3-0.4 pronotal width, sparse on anterior margin (Fig. 16.143); lacinia broadly triangular (Fig. 16.144); widely distributed, common Haploperla Navas

76'.

Pronotal fringe hairs shorter, about 0.25 pronotal width, numerous on anterior margin (Fig. 16.140); lacinia narrowly triangular (Fig. 16.142); AK, Northwest Territories, Yukon Alaskaperla Stewart and DeWalt(one species, A. ovibovis(Richer))

77(8'). IT.

Gills present on 1 or more thoracic segments (Fig. 16.145) 78 Gills absent from thoracic segments 82 Prothoracic gills present(Fig. 16.145) 79 Prothoracic gills absent; Western North America Setvena lilies Lateral abdominal gills present(Fig. 16.146); CA,NY Oroperla Needham (one species, 0. barbara Needham) Lateral abdominal gills absent 80 Cervical gills present(Fig. 16.145); Western North America Perlinodes Needham and Claassen (one species, P. aureus(Smith)) Cervical gills absent 81 Prothoracic gills reduced to nipple-like stubs, meso- and metathoracic gills forked (Fig. 16.35); CA, OR Salmoperla Baumann and Lauck (one species, S. sylvanica Baumann and Lauck) Prothoracic gills well developed; meso- and metathoracic gills simple, thumb-like (Fig. 16.147); Western North America Megarcys Klapalek Lacinia unidentate (Fig. 16.148)(early instar Kogotus bidentate) 83 Lacinia bidentate (Fig. 16.149) 86 Abdomen with dark longitudinal pigment bands(Fig. 16.15); widely

78(77). 78'. 79(78). 79'. 80(79'). 80'. 81(80'). 81'. 82(7T).

82'. 83(82).

distributed

Isoperla Banks(in part)

83'. 84(83').

Abdomen without dark longitudinal pigment bands Lacinia broad basally, abruptly narrowed into a long terminal spine(Fig. 16.150); Eastern

84'.

Lacinia gradually narrowed from base to terminal spine (Fig. 16.148)

85(84').

Mesosternum without transverse anterior furrow; Western North America,

North America

common

85'.

84

Remenus Richer

85 Kogotus Richer

Mesostemal furcal pits often connected by a transverse anterior furrow, sometimes difficult to discern in young nymphs (Fig. 16.151) CA,NY,OR,WA,rare Rickem Jewett(one species, R. sorpta (Needham and Claassen))[note: often nymphs of Kogotus and Rickera cannot be satisfactorily separated]

86(82').

Abdomen with longitudinal pigment bands (Fig. 16.15)

87

86'.

Abdomen without longitudinal pigment bands

95

87(86).

Abdomen with pale, median, longitudinal pigment band

88

87'. 88(87).

Abdomen with dark, median, longitudinal pigment band 90 Apical lacinial tooth about as long as rest of lacinia (Fig. 16.152); mesostemal Y-arms with secondary furrows extending to anterior corners of furcal pits (Fig. 16.153); BC,CA,OR, WA Osobenus Richer one species, O. yakimae(Hoppe))

Figure 16.125

Figure 16.126

Figure

Figure 16.129

Figure 16.130

Figure 16.135 Figure 16.131

Figure 16.132

Figure 16.134 Figure 16.133

Figure 16.138 Figure 16.136

Figure 16.137

Figure 16.125 Kathroperia perdita (Chloroperlidae) nymphal right lacinia, ventral. Figure 16.126 Paraperia frontalis (Chloroperlidae) nymphal right laclnia, ventral. Figure 16.127 Sweltsa oregonensis (Chloroperlidae) nymphal mesonotum. Figure 16.128 Suwallia pallidula (Chloroperlidae) nymphal mesonotum. Figure 16.129 Sweltsa oregonensis (Chloroperlidae) nymphal right foreleg. Figure 16.130 Sasquaperia hoopa (Chloroperlidae) nymphal mesosternum. Figure 16.131 Suwallia pallidula (Chloroperlidae) nymphal right cercus, lateral. Figure 16.132 Sasquaperia hoopa (Chloroperlidae) nymphal right foreleg. 456

Figure 16.139

Figure 16.133 Plumiperia diversa (Chloroperlidae) nymphal right cercus, lateral. Figure 16.134 Rasvena terna (Chloroperlidae) nymphal right cercus, lateral. Figure 16.135 Triznaka pintada (Chloroperlidae) nymphal head and pronotum, dorsal. Figure 16.136 Plumiperia diversa (Chloroperlidae) nymphal head and pronotum, dorsal.

Figure 16.137 BIsancora rutrlformis (Chloroperlidae) nymphal head and pronotum, dorsal. Figure 16.138 BIsancora rutrlformis (Chloroperlidae) nymphal right laclnia, ventral. Figure 16.139 Rasvena terna (Chloroperlidae) nymphal head and pronotum, dorsal.

Chapter 16 Plecoptera

88'. 89(88'). 89'.

90(87'). 90'. 91(87). 91'. 92(91). 92'.

Apical lacinial tooth much shorter than rest of lacinia (Fig. 16.154); mesosternal Y-arms lacking secondary furrows extending to anterior corners of furcal pits 89 Inner lacinial margin with row of at least 4-5 long seta; no prominent knob bearing pegs below subapical lacinial tooth (Fig. 16.155); widely distributed Isoperla Banks (in part) Inner lacinial margin lacking a row of long setae; prominent knob below subapical lacinial tooth bearing 3-4 stout peg-like setae(may appear as a 3rd tooth under low-power magnification; Fig. 16.154); CA Susulus Bottorff and Stewart(one species, S. venustus (Jewett)) Lacinia quadrate (Fig. 16.156) 91 Lacinia triangular or subquadrate (Fig. 16.152) 94 Lacinia with a dense brush of stout setae (Fig. 16.156); inner lacinial margin glabrous 92 Lacinial with one or more rows of stout setae(Figs. 16.155 and 16.157); inner lacinial margin setose. ... 93 Apical cereal segments fringed with fine hairs dorsally and ventrally (Fig. 16.158); Western North America Cascadoperla Szczytko and Stewart(one species, C. trictura(Hoppe)) Apical cereal segments fringed along ventral margins or not at all; widely distributed

93(91'). 93'. 94(90'). 94'. 95(86'). 95'.

96(95).

97(96').

Isoperla Banks (in part)

Cerci without marginal fringe; mesosternal Y-arms enclosing an area of abundant clothing hairs (Fig. 16.159); CA,OR, WA Calliperla Banks(one species, C. luctuosa (Banks)) Cerci with ventral fringe on apical segments(Fig. 16.160); mesosternal Y-arms enclosing an area with few or no clothing hairs; widely distributed Isoperla Banks(in part) Mesosternal Y-arms sinuate (Fig. 16.161); cerci lacking fine setal fringe; CA Cosumnoperla Szczytko and Bottorff Mesosternal Y-arms straight (Fig. 16.162); cerci with fine setal fringe (Fig. 16.160); widely distributed Isoperla Banks (in part) Mesosternal Y-arms meet or approach anterior corners of furcal pits (Fig. 16.163); mandibles deeply cleft, separating teeth into two major cusps 96 Mesosternal Y-arms meet or approach posterior corners of furcal pits (as in Fig. 16.162): mandibles not deeply cleft 98 Apical lacinial tooth short, much less than one-third the outer lacinial length, ventral submarginal lacinial setae extending well onto apical tooth as a closely set row (Fig. 16.164); BC, CA, NV,OR, WA

96'.

457

Frisonia Ricker(one species, F. picticeps (Hanson))

Apical lacinial tooth long, greater than one-third the outer lacinial length; ventral submarginal lacinial setae end at inner base of apical tooth 97 Outermost cusp of both mandibles serrate (Fig. 16.165); mesal tufts of silky setae on occiput; abdominal segments 1-2, divided by pleural fold; Western North America

97'.

98(95').

98'. 99(98').

99'. 100(99'). 100'.

Skwala Ricker

Outermost cusp of both mandibles unserrated, or with indistinct serrations on left mandible only; occiput without mesal tuft of silky setae; abdominal segments 1-3 divided by pleural fold; Western North America and higher latitudes of Eastern North America Arcynopteryx Klapalek (one species, A. dichroa (McLachlan)) Femora and tibiae without long setal fringe; posterolateral margins of pronotum notched; Southern Appalachians, rare Oconoperla Stark and Stewart (one species, O. innubila (Needham and Claassen)) Femora or tibiae or both with long setal fringe; posterolateral margins of pronotum smoothly rounded; widespread; some genera common 99 Mesosternum with median longitudinal suture connecting fork of Y-arms with transverse suture (Fig. 16.166); widely distributed Isogenoides Klapalek Mesosternum without median longitudinal suture 100 Submental gills conspicuous, projecting portion usually 2 times or more as long as basal diameter (Fig. 16.37) 101 Submental gills absent or barely projecting beyond submentum 104

Figure 16.141

Figure 16.142

Figure 16.140

Figure 16.143

y.

Figure 16.151

Figure 16.147

Figure 16.145 Figure 16.146

Figure 16.144

Figure 16.148

Figure 16.149

Figure 16.140 Alaskaperia ovibovis (Chloroperlidae) nymphal head and pronotum, dorsal. Modified from Stewart etal. (1991). Figure 16.141 Rasvena terna (Chloroperlidae) nymphal right laclnia, ventral. Figure 16.142 Alaskaperia ovibovis (Chloroperlidae) nymphal right laclnia, ventral. Figure 16.143 Haploperia brevis (Chloroperlidae) nymphal head and pronotum, dorsal. Figure 16.144 Haploperia brevis (Chloroperlidae) nymphal right laclnia, ventral. Figure 16.145 Perlinodes aureus (Perlodidae) nymphal head and thorax, ventral. Figure 16.146 Oroperia barbara (Perlodidae) nymphal body, ventral. 458

Figure 16.150

Figure 16.152

Figure 16.147 Megarcys signata (Perlodidae) nymphal head and thorax, ventral. Figure 16.148 Kogotus nonus (Perlodidae) nymphal left laclnia, ventral.

Figure 16.149 Malirekus hastatus (Perlodidae) nymphal left laclnia, ventral. Figure 16.150 Remenus bilobatus (Perlodidae) nymphal left laclnia, ventral. Figure 16.151 Rickera sorpta (Perlodidae) nymphal mesothorax.

Figure 16.152 Osobenus yakimae (Perlodidae) nymphal right laclnia, ventral.

Chapter 16 Plecoptera

101(100).

Apical lacinial tooth about half the total outer lacinial length; Western North America, uncommon

lOr.

459

Pictetiella lilies

Apical lacinial tooth about one-third or less the total outer lacinial length (Fig. 16.149); Eastern North America, usually common

102

102(101'). Ventral lacinial surface with basal patch of about fifty dark clothing hairs (Fig. 16.149) Malirekus Richer (in part) 102'. Ventral lacinial surface without dark clothing hairs, a patch of about 10 setae may be present (Fig. 16.167) 103 103(102'). Transverse dark pigment band of frons lateral to median ocellus interrupted by circular yellow areas(Fig. 16.168); ventral lacinial surface with outer patch of about 10 setae (Fig. 16.169); right mandible with 4 teeth; Eastern North America

103'.

Hydroperla Frison

Transverse dark pigment band of frons uninterrupted by enclosed yellow areas(Fig. 16.170); ventral lacinial surface without outer patch of setae; right mandible with 5 teeth; Eastern North America Helopicus Kicker

104(100'). Inner lacinial margin with a low knob below subapical tooth (Fig. 16.149) 104'. Inner lacinial margin without low knob below subapical tooth (Fig. 16.155)

105 107

105(104'). Outer ventral lacinial surface with basal patch of about 50 dark clothing hairs (Fig. 16.149); Eastern North America Malirekus Kicker (in part) 105'. Outer ventrallacinial surface with fewer than 50 basal clothing hairs(Fig. 16.167) 106 106(105'). Marginal lacinial setal row extending from near subapical tooth to near base (Fig. 16.167); labrum with yellow longitudinal mesal band; submental gills very short, if present; Eastern North America Yugus Kicker

106'.

Marginal lacinial seta! row restricted to apical half (Fig. 16.171); labrum without longitudinal band; submental gills absent; Western North America and higher latitudes of Eastern North America

Dima Billberg (in part)

107(104'). Occiput or anterolateral prothoracic margins or both with a row of short stout setae 107'.

108

Occiput and anterolateral prothoracic margins without rows of short, stout setae, a few long setae may be present

112 108(107). Ocellar region transversed by dark band connecting lateral ocelli and eyes but not covering anterior ocellus (Fig. 16.38), a dark form lacking the light medial area of head, with light M-line; abdominal terga with pale medial, paired spots; Eastern North America Clioperla Needham and Claassen (one species, C. clio(Newman)) 108'. Ocellar triangle more or less completely covered by dark pigment; abdominal terga with 3 pairs pale or dark spot, or lacking such spots 109 109(108'). Outer mandibular cusp serrate; abdominal terga with anterior narrow dark basal bands and conspicuous transverse row of dark spots; CA Baumannella Stark and Stewart (one species, B. alameda(Needham and Claassen)) 109'. Major mandibular cusp without serrations; abdominal terga with variable color pattern 110 110(109'). Mesosternal Y with indistinct anterior extensions which approach furcal pits (Fig. 16.172); CA,OK 110'.

Chernokrilus Kicker (one species, C. misnomus(Claassen))

Mesosternal Y without anterior extensions; widely distributed

Ill

111(110'). Occipital area with a pair of pale oval markings adjacent to compound eyes; posterior margins of ovals marked by dense irregular rows of short, thick setae (Fig. 16.173); Western North America and higher latitudes of Eastern North America Dima Billberg (in part) 111'. Occipital area without pale oval markings margined by short, thick setae; widely distributed Isoperla Banks (in part) 112(107'). Mesosternal Y with basal stem and fork (Fig. 16.174); widely distributed Cultus Kicker 112'.

Mesosternal Y without stem and fork (Fig. 16.175); Eastern North America

Drp/oper/a Needham and Claassen

?k

Figure 16.153

Figure 16.161

Figure 16.159

T Figure 16.162

Figure 16.163

Figure 16.154

Figure 16.166

Figure 16.155

Figure 16.156

serrations

Figure 16.158

Figure 16.160

Figure 16.157

Figure 16.153 Osobenus yakimae (Perlodldae) nymphal mesothorax. Figure 16.154 Susulus venustus (Perlodidae) nymphal right iacinia and inset of knob, ventral. Figure 16.155 Isoperia bilineata (Perlodidae) nymphal right Iacinia, ventral. Figure 16.156 Cascadoperia trictura (Perlodidae) nymphal right Iacinia, ventral. Figure 16.157 Calliperia luctuosa (Perlodidae) nymphal right Iacinia, ventral. Figure 16.158 Cascadoperia trictura (Perlodidae) nymphal right cercus, lateral. Figure 16.159 Calliperia luctuosa (Perlodidae) nymphal mesosternum. 460

Figure 16.164

Figure 16.165

Figure 16.160 Isoperia bilineata (Perlodidae) nymphal right cercus, basai, medial and distal segments, lateral. Figure 16.161 Cosumnoperia hypocrena (Perlodidae) nymphal mesosternum. Figure 16.162 Isoperia bilineata (Perlodidae) nymphai mesosternum. Figure 16.163 Skwaia amerlcana (Perlodidae) nymphal mesosternum. Figure 16.164 Frisonia picticeps (Perlodidae) nymphal left Iacinia, ventral. Figure 16.165 Skwaia amerlcana (Perlodidae) nymphal left mandible, ventral. Note serrations on outer cusp. Figure 16.166 Isogenoides zionensis (Perlodidae) nymphal mesosternum.

Chapter 16 Plecoptera

461

Figure 16.170 Figure 16.168

i

Figure 16.172

Figure 16.173

Figure 16.167 t

Figure 16.174

Figure 16.169

Figure 16.171

t

Figure 16.175

Figure 16.167 Yugus bulbosus (Perlodldae) nymphal left laclnia, ventral.

Figure 16.168 Hydroperia crosbyi(Perlodldae) nymphal head and pronotum. Figure 16.169 Hydroperia crosbyi(Perlodldae) nymphal left laclnia, ventral. Figure 16.170 Heiopicus nalatus (Perlodldae) nymphal head and pronotum. Figure 16.171 Diura knowitoni (Perlodldae) nymphal left laclnia, ventral.

Figure 16.172 Chernokrilus misnomus (Perlodldae) nymphal mesosternum. Figure 16.173 Diura knowitoni(Perlodldae) nymphal head and pronotum. Figure 16.174 Cuitus aestivaiis (Perlodldae) nymphal mesosternum.

Figure 16.175 Diploperia dupiicata (Perlodldae) nymphal mesosternum.

462

Chapter 16 Plecoptera

KEY TO THE FAMILIES AND GENERA OF NORTH AMERICAN PLECOPTERA ADULTS

1.

Basal(1st) tarsal segment about as long, or slightly longer than apical(3rd)segment(Fig. 16.176);

r.

Basal tarsal segment much shorter than apical segment(Fig. 16.177); mid and basal tarsal segments with well-developed ventral membranous pads

tarsi completely sclerotized on venter

2(1).

5

Mid (2nd) tarsal segment about as long as basal (1st) segment(Fig. 16.178); gill scar present(Fig. 16.179) or absent on inner coxal surface

2'.

2

TAENIOPTERYGIDAE

9

Mid tarsal segment much shorter than basal segment(Fig. 16.176); gill scar absent from inner coxal surface

3(2'). 3'.

3

Cerci multisegmented (Fig. 16.180); 2nd anal vein of forewing simple and unforked (Fig. 16.181); usually 1 or 2 intercubital crossveins CAPNIIDAE 16 Cerci 1-segmented (Fig. 16.182); 2nd anal vein of forewing forked (Fig. 16.183); usually 5 or more intercubital crossveins

4(3').

4'.

gills absent

5(1').

LEUCTRIDAE

PELTOPERLIDAE .... 86

Two slightly enlarged ventroapical spurs occur on each tibia, each surrounded by subapical membranous area (Fig. 16.190); sternacostal sutures extend laterad of anterior corners of metathoracic furcal pits, often to margins of basisternum (Fig. 16.191), or sutures incomplete near furcal pits (Fig. 16.192); posterolateral angles of metasternum not projecting (Fig. 16.192); two or three ocelli

6

6(5').

Gill remnants conspicuous on thoracic sterna between coxae and on first 2 or 3 abdominal sterna (Fig. 16.193); forewing anal region with two or more rows of crossveins (Fig. 16.194) PTERONARCYIDAE

6'.

Thoracic gill remnants, if present, restricted to area behind coxae (Fig. 16.195) and absent from basal abdominal sterna; forewing anal region with, at most, one row of crossveins Lateral margins of pronotum not bent downward; second anal vein of forewing often forked between anal cell and wing margin (Fig. 16.196); hind wing anal region usually with less than 5 longitudinal veins; body color variable but often green or yellow in life; apical maxillary palpal segment often much reduced in size relative to penultimate segment;

7(6').

gill remnants absent

7'.

8(7').

71

Ventroapical tibial spurs small and arranged in one or two rows(Fig. 16.188); metathoracic sternacostal sutures, if present, arise from posterior corners of furcal pits, posterolateral angles of metasterna usually project behind coxae (Fig. 16.189); two ocelli

5'.

4

Apical segment of labial palpus circular and larger than preceding segment(Fig. 16.184); wings lying flat over abdomen at rest; forewing often with an X-pattern of crossveins at cord (Fig. 16.183); cervical gills sometimes present (Fig 16.185) NEMOURIDAE 51 Apical segment of labial palpus similar to preceding segment(Fig. 16.186); wings rolled around abdomen at rest giving body a slender, needle-like appearance; forewings without an X-pattern of crossveins at cord (Fig. 16.187);

CHLOROPERLIDAE

91

7

92

Lateral margins of pronotum usually bent sharply downward, partially covering sides of prothorax (Fig. 16.197); second anal vein of forewing forked or unforked between anal cell and wing margin; hind wing anal region with 5 or more longitudinal veins; body color yellow, brown or black; apical maxillary palpal segment not greatly reduced in size; gill remnants present or absent 8 Metathoracic sternacostal sutures along posterior margin of basisternum not reaching furcal pits (Fig. 16.195), obscure arched grooves extend anterolaterad from furcal pits; cubitoanal crossvein of forewing usually touching, or very near anal cell(Fig. 16.199); thorax(and sometimes paraprocts) often with branched gill remnants or ragged gill stubs (Fig. 16.195) PERLIDAE 105

Figure 16.179

Figure 16. En^sobasjsternufti

tarsal pad

Figure 16.177

gill scar

cercus

Figure 16.178

paraproct

Figure 16.180

apical costal space

basal curve

ve icle

Intercubltal crossvein

paraproct

Figure 16.181

lobes

Figure 16.182

apical palpal segment

x-pattern

Figure 16.183

intercubltai crossveins Figure 16.184

Figure 16.176 Megaleuctra complicata (Leuctridae) adult foreleg tarsal segments, lateral. Figure 16.177 Acroneuria arenosa (Perlidae) adult foreleg tarsal segments, lateral. Figure 16.178 Taeniopteryx maura (Taenloptetygidae) adult foreleg tarsal segments, lateral. Figure 16.179 Taeniopteryx maura (Taeniopterygidae) adult mesosternum.

Figure 16.180 Allocapnia virginiana (Capniidae) adult male terminalia, ventral.

Figure 16.181 Mesocapnia frisoni(Capniidae)forewing. Figure 16.182 Leuctra grandis (Leuctridae) adult male terminalia, ventral.

Figure 16.183 Amphlnemura wui(Nemouiidae)forewing. Figure 16.184 Amphlnemura wui(Nemouridae) adult fiead, ventral. 463

464

8'.

Chapter 16 Plecoptera

Metathoracic sternacostal sutures extend laterad from anterior corners of furcal pits to margins of basisternum (Fig. 16.198); cubitoanal crossvein of forewing often absent, or removed from anal cell by at least its own length (Fig. 16.200); thorax and paraprocts without branched gill remnants but finger-like (or forked) gills sometimes occur on submentum, thorax or sides of abdomen (Fig. 16.198)

9(2). 9'. 10(9').

PERLODIDAE

Gill scar present on inner surface of coxa (Fig. 16.179); abdominal sternum 9 not strongly produced over sternum 10 (Fig. 16.201); widespread Taeniopteryx Pictet Gill scar absent from inner coxal surface; abdominal sternum 9 strongly produced over sternum 10 (Fig. 16.202) 10 Forewing with humeral crossvein and sometimes an apical crossvein present in costal spaces, but without other costal crossveins (Fig. 16.203); forewings of some males shortened or malformed

10'. 11(10).

11'. 12(11').

12'.

138

11

Forewing with at least 1 costal crossvein in addition to humeral and apical crossveins (Fig. 16.204); forewings of males not shortened or malformed 13 Forewing without crossvein in apical costal space (Fig. 16.205); males of some species with shortened or malformed forewings and long hind wings; male abdominal sternum 9 with or without vesicle; widespread Oemopteryx Klapalek (in part) Forewing with an apical costal crossvein (Fig. 16.203); forewings not shortened or malformed; males with vesicle on abdominal sternum 9(Fig. 16.206) 12 Upturned tip of male abdominal sternum 9 slightly asymmetrical, notched and bearing a dorsal asymmetrical bulbous process (Fig. 16.206); female abdominal sternum 9 gradually narrowed, triangular in outline and sclerotized band of female abdominal sternum 8 small(Fig. 16.207); Eastern North America Bolotoperla Ricker and Ross (only one species, B. rossi(Frison)) Upturned tip of male abdominal sternum 9 symmetrical, unnotched and without dorsal bulbous process; female abdominal sternum 9 broadly rounded, parabolic in outline and sclerotized band of female abdominal sternum 8 extends across most of segment length (Fig. 16.208); widespread Oemopteryx Klapalek (in part)

13.

Eastern North America

14

13'.

Western North America

15

14(13).

Apex of male abdominal sternum 9 process usually sharply upturned in lateral aspect (Fig. 16.209), and narrowed in ventral aspect; free portion of female abdominal sternum 9 about as long as basal width (Fig. 16.210); Eastern North America Strophopteryx Frison Apex of abdominal sternum 9 not upturned (Fig. 16.211), and broad in ventral aspect; free portion of female abdominal sternum 9 shorter than basal width (Fig. 16.212); widespread Taenionema Banks(in part) Rs vein of forewing with three branches and Cu vein with four or five branches (Fig. 16.213); upturned tip of male abdominal sternum 9 strongly narrowed in dorsal aspect; Western North America Doddsia Needham and Claassen (one species, D. occidental^ (Banks)) Rs vein of forewing with two branches and Cu vein with two or three branches(Fig. 16.204); tip of male abdominal sternum 9,if upturned, not strongly narrowed; widespread.... Taenionema Banks (in part) Epiproct present on abdominal tergum 10(Fig. 16.214); abdominal sternum 9 covers much of sternum 10(Fig. 16.180); abdominal sternum 8 unmodified

14'.

15(13').

15'. 16(3).

Males

16'.

Epiproct absent, tergum 10 unmodified; abdominal sternum 9 not extending over abdominal sternum 10; abdominal sternum 8 at least slightly modified as subgenital plate (Fig. 16.215) Females

17(16).

17

35

Vesicle present on abdominal sternum 9(Fig. 16.216)

18

17'.

Vesicle absent from abdominal sternum 9(Fig. 16.180)

20

18(17).

R vein of forewing curved slightly cephalad distal to origin of Rs (Fig. 16.181); Western North America; uncommon Bolshecapnia Ricker, Eurekapnia Stark and Broome, Sasquaeapnia Baumann and Broome*

* Broome et al. 2019. Illiesia 15: 1-26.

tibiai spur row cervical gill remnant

Figure 16.185

Figure 16.188

Figure 16.186

arculus

tibiai spur

Figure 16.187 sternacostal suture

Figure 16.190

Figure 16.189 projecting metasternal angle

sternacostal suture

Y-ridge

*■

I

st^nacostal \

I

/

futures

furcal pit

Figure 16.192 Figure 16.191

Figure 16.185 Zapada oregonensis (Nemourldae) head cervical gill remnants. Figure 16.186 Leuctra grandis (Leuctridae) adult head, ventral.

Figure 16.187 Calileuctra dobryi (Leuctridae) forewing. Figure 16.188 Tallaperia anna (Peltoperlidae) adult foreleg tibiai apex, ventral.

Figure 16.189

Tallaperia anna (Peltoperlidae) adult

mesosternum.

Figure 16.190 Acroneuria arenosa (Perlidae) adult foreleg tibiai apex. Figure 16.191 Yugus kondratleffi (Perlodidae) adult meso- and metasterna.

Figure 16.192 mesosternum.

Hesperoperia paclfica (Perlidae) adult 465

466

Chapter 16 Plecoptera

18'. 19(18').

R vein of forewing straight beyond Rs origin (Fig. 16.217) Mesosternum postfurcal plates separated from spinasternum and furcasternum by membrane (Fig. 16.218); cerci consisting of ■

Figure 16.211

[-i

iM'k

0ji W:/

Figure 16.210

Figure 16.212

Figure 16.215

Figure 16.213

Figure 16.214

straight R

epiproct

Figure 16.217

Figure 16.216

Figure 16.209 Strophopteryx cucullata (Taeniopterygidae) adult male terminalia, lateral. Figure 16.210 Strophopteryx cucullata (Taeniopterygidae) adult female terminalia, ventral. Figure 16.211 Taenionema atlantlcum (Taeniopterygidae) adult male terminalia, lateral. Figure 16.212 Taenionema atlantlcum (Taeniopterygidae) adult female terminalia, ventral. Figure 16.213 Doddsia occidentalls (Taeniopterygidae) forewing.

Figure 16.214

Capnia gracilaria (Capniidae) adult

male terminalia, dorsal.

Figure 16.215

Capnia gracilaria (Capniidae) adult

female terminalia, ventral.

Figure 16.216

Isocapnia grandls (Capniidae) adult

female terminalia, ventral.

Figure 16.217 forewing.

Isocapnia grandls (Capniidae)

469

470

Chapter 16 Plecoptera

31(30').

Dorsal aspect of epiproct housing a large membranous, eversible crest or apically frayed duct (Figs. 16.222 and 16.225), terminal spine absent; duct may be retracted and lie within trough formed by epiproct sclerite, or expanded into a balloon-like structure arising in the apical half or third of epiproct; widespread Pamcapnia Hanson (in part)

31'.

Dorsal aspect of epiproct variously modified but without retracted or expanded balloon-like eversible crest duct; postfurcasternum not fused, separate

32

32(31').

Epiproct with small basally projecting lower arm (Fig. 16.228); abdominal tergum 7, and in 1 species also tergum 8, with dorsal hump Capnura Banks (in part)

32'.

Epiproct without projecting lower arm; humps on abdominal terga present or absent, if present, humps may occur on one or more of abdominal segments 5-9(including 7, or 7-8) 33

33(32).

Epiproct with basal sclerite; dorsolateral horns prominent, arching 15-30% of epiproct length

33'.

(Fig. 16.229); CA, NV, OR Sierracapnia Bottorff and Baumann Epiproct with basal sclerite absent or vestigial; dorsolateral horns reduced 34

34(33). 34'.

Dorsal epiproct sclerite divided entire length (Fig. 16.230a); lower portion of epiproct sclerite absent(Fig. 16.230b); Western North America Arsapnia Banks Dorsal epiproct sclerite not fully divided (Fig. 16.23 la); lower portion of epiproct present or vestigal Capnia Pictet (see Muranyi et al. (2014)for additional characters to distinguish Capnia s.s. and Capnia s.l.) Wingless or with very short wings 36

35(16'). 35'. 36(35).

Postfurcal plates of mesosternum fused with spinasternum and furcasternum

36'.

(similar to Fig. 16.220) Pamcapnia Hanson (in part) Postfurcal plates of mesosternum completely separated by membrane from spinasternum and

37(36').

furcasternum (as in Fig. 16.218) Eastern North America

37'.

Western North America

38(37'). 38'.

Subgenital plate deeply emarginate mesally (Fig. 16.232); known from Colusa Co., CA Pamcapnia Hanson (in part){P. boris Stark and Baumann) Subgenital plate rounded or truncate mesally 39

39(38').

Known from Lake Tahoe, CA, NV

39'.

Known from AK

40(39).

Subgenital plate uniformly pigmented. Lake Tahoe, CA, NY Capnia Pictet (in part){Capnia lacustra Jewett, adults aquatic)

40'.

Subgenital plate with dark longitudinal median band (Fig. 16.233)

41(35'). 41'.

Macropterous or brachypterous

41

37 Allocapnia Claassen (in part) 38

40

Mesocapnia Rauser (in part)(M bergi(Ricker))

f/tacapn/a Gaufin (in part) Postfurcal plates of mesosternum fused with spinasternum and furcasternum (Fig. 16.220) 42 Postfurcal plates of mesosternum completely separated by membrane from spinasternum and

42(41).

furcasternum (Fig. 16.218) R vein of forewing curved slightly cephalad distal to origin of Rs (Fig. 16.181)

42'.

R vein of forewing straight beyond origin of Rs (Fig. 16.217)

43(42').

Cerci consisting of < 11 segments; abdominal sternum 8 posterior to subgenital plate with a transverse, hairy sclerite (Fig. 16.234) Nemocapnia Banks(one species, N. Carolina Banks) Cerci of most species consisting of >11 segments (only /. vedderensis(Ricker) with 6-9); abdominal sternum 8 without transverse, hairy sclerite posterior to subgenital plate 44 Eastern North America Pamcapnia Hanson (in part) Western North America Isocapnia Banks R vein of forewing straight beyond origin of Rs(as in Fig. 16.217) 46

43'. 44(43'). 44'. 45(41').

45

P«rac«/>n/a Hanson (in part) 43

Chapter 16 Plecoptera

471

epiproct

furcasternum

horns

furcasternum

"N postfurcal

spinasternum

plate ^postfurcal plate

spinasternum

Figure 16.218

Figure 16.220

Figure 16.219

dorsat humps

1 /

,: ■ . i \ r J

y I I

1 1 I 'i

I' l ■epiproct trough / ' < ' ,

9^^ 1 forked

Figure 16.223

Figure 16.222

Figure 16.221

^ "s /i

epiproct spin^Jj

ventral arm

dorsal arm

Figure 16.225

Figure 16.226

Figure 16.224

Figure 16.218

Eucapnopsis brevlcauda (Capnildae)

Figure 16.223 Allocapnia granulata (Gapniidae) adult

adult mesosternum.

male terminalia, dorsal.

Figure 16.219 Eucapnopsis brevlcauda male (Capnildae) adult terminalia, dorsal. Figure 16.220 Isocapnia grandls (Gapniidae) adult

aduit male terminalia, dorsal.

mesosternum.

male terminalia, dorsal.

Figure 16.221

Allocapnia granulata (Gapniidae) adult

male terminalia, lateral.

Figure 16.222

Paracapnia disala male (Gapniidae)

adult terminalia, lateral.

Figure 16.224

Figure 16.225 Figure 16.226

Utacapnia lemonlana (Gapniidae)

Paracapnia enslcala (Gapniidae) adult Mesocapnia frisoni (Gapniidae) adult

male terminalia, dorsal.

472

45'. 46(45). 46'.

47(45'). 47'. 48(47'). 48'. 49(48').

49'. 50(49').

50'.

Chapter 16 Plecoptera

R vein of forewing curved slightly cephalad distal to origin of Rs(Fig. 16.181) 47 Cerci with 15 segments; center of sternum 8 sclerotized; Eastern North America Allocapnia Claassen (in part) Subgenital plate usually with dark median band and small posterior projection and median notch (Fig. 16.233) [/tflca/w/a Gaufm (in part) Subgenital plate with or without dark median band, if present, posterior margin of plate is not a small, notched projection 48 Median field of subgenital plate membranous, often with an impressed Y-shaped sclerite (Fig. 16.235) C«/;MHra Banks Median field of subgenital plate variable, but not membranous with Y-shaped sclerite 49 Abdominal sternum 8 uniformly pigmented except for a dark pair of lateral spots(Fig. 16.236); abdominal sternum 9 with a basolateral pair of L-shaped pigment spots; subgenital plate scarcely projecting or narrowed to a small nipple-like projection Mesocapnia Rauser Abdominal sternum 8 usually more darkly pigmented along posterior margin or on subgenital plate; abdominal sternum 9 without L-shaped pigment spots; subgenital plate variable 50 Subgenital plate usually narrow and projecting over base of abdominal sternum 9; body length usually at least 9 mm Bolshecapnia Ricker, Eurekapnia Stark and Broome, Sasquacapnia Baumann and Broome* Subgenital plate usually broad and typically not projecting over base of abdominal sternum 9; body length usually less than 8 mm Arsapnia Banks, Capnia Pictet (in part), Sienacapnia Bottorff and Baumann

51(4).

Cervical or submental gill remnants present(Fig. 16.185)

52

51'.

Gill remnants absent

57

52(51).

Wings reduced to minute stubs; hind legs much longer than others; known from spring seeps near Wahkeena Falls, Columbia River Gorge, OR Nanonemoura

52'.

Macropterous or slightly brachypterous; hind legs only slightly longer than other legs; widely

Baumann and Fiala(one species, N. wahkeena (Jewett)) distributed

53(52'). 53'. 54(53').

54'. 55(54').

55'.

56(55').

56'.

53

Each gill remnant unbranched (Fig. 16.185); mountains east(uncommon)and west(common) Zapada Ricker (in part) Each gill remnant with multiple branches(Fig. 16.238) 54 Gill remnants arise from submentum; Western North America Visoka Ricker(one species, V. cataractae(Neave)) Gill remnants arise from cervical region (Fig. 16.238) 55

Male with dorsal membranous or sclerotized lobe on cereal base (Fig. 16.239); female subgenital plate notch narrow and extending forward from posterior margin of sternum 8 to near mid length of abdominal sternum 8(Fig. 16.240); Western North America Malenka Ricker Male without dorsal lobe on cerci (Fig. 16.241); female subgenital plate notch present or absent, if present, notch or plate is set forward of posterior margin of abdominal sternum 8 (Fig. 16.242) 56 Gill remnants with at least 6 branches; male paraprocts divided into three lobes, mid and outer lobes bearing spines(Fig. 16.241); female abdominal sternum 7 not usually produced over most of abdominal sternum 8; subgenital plate of abdominal sternum 8 forming a notched, sclerotized band set anterior to the hind margin of sternum 8(Fig. 16.242); widespread Amphinemura Ris Gill remnants with 4 or fewer branches(Fig. 16.185); male paraprocts quadrate, divided into two lobes, inner lobe often concealed and spines absent from paraprocts (Figs. 16.243 and 16.244); female abdominal sternum 7 usually produced over abdominal sternum 8 (Fig. 16.245); subgenital plate a dark mesal patch over gonopore; mountains east(uncommon)and west(common) Zapada Ricker (in part)

* Broome et al. 2019. Illiesia 15: 1-26.

straight R

ventral arm

Figure 16.227

m tev^8^f-is P^M dorsolateral horns

;:v.i»

x.fciBSSiiiija'

(vr—r...-i'^"*i^.>.-^jS^;

Figure 16.228

Figure 16.229

undivided divided

bwer sc erite

Figure 16.231

Figure 16.230

If''/' i ,>,

i\h l,K V

\

^' r 'i^

/

^'

oi-Vv"

Iran verse sclerite

Figure 16.232

Figure 16.233

Figure 16.227 Nemocapnia Carolina (Capniidae) forewing. Figure 16.228 Capnura elevata adult male terminalia,

Figure 16.234

Figure 16.235

Figure 16.232 Paracapnia borisi(Capniidae) adult female terminalia, ventral. From Stark and Baumann

lateral.

(2004). Figure 16.233 Utacapnia lemoniana (Capniidae)

Figure 16.229 Sierracapnia washoe (Capniidae)

adult female terminalia, ventral.

adult male terminalia, lateral. Modified from Bottorff and

Figure 16.234 Nemocapnia caroiina (Capniidae)

Baumann (2015). Figure 16.230 Arsapnia decepta (Capniidae) adult male epiproct.(a) dorsal,(b) lateral. From Nelson and Baumann (1989). Figure 16.231 Capnia californica (Capniidae) adult male epiproct.(a) dorsal,(b) lateral. From Nelson and Baumann (1989).

adult female terminalia, ventral.

Figure 16.235 Capnura wanica (Capniidae) adult female terminalia, ventral. From Nelson and

Baumann (1987).

473

474

Chapter 16 Plecoptera

57(51').

A1 and A2 veins united near forewing margin (Fig. 16.246); cerci small and unsclerotized; male epiproct bilaterally asymmetrical; widespread Soyedina Ricker

57'.

A1 and A2 veins not united in forewings (Fig. 16.247): cerci variable; male epiproct bilaterally symmetrical

58(57').

Epiproct enlarged into a probe-like structure on abdominal tergum 10(Figs. 16.248 and 16.249); vesicle usually present on abdominal sternum 9(Fig. 16.250) Males

58'.

59

Epiproct simple, abdominal tergum 10 without probe-like structure; vesicle absent from abdominal sternum 9 Females

59(58).

58

65

Vesicle absent from abdominal sternum 9

60

59'.

Vesicle present on abdominal sternum 9(Fig. 16.250)

61

60(59).

Forewing terminal costal crossvein joins Sc in apical costal space(Fig. 16.247); abdominal tergum 10 simple, without lobes; Eastern North America, rare

60'.

Paranenwura Needham and Claassen

Forewing terminal costal crossvein joins R in apical costal space (Fig. 16.251); abdominal tergum 10 with a pair of elevated spiny lobes (Fig. 16.252); Western North America; glacial or snowmelt-fed streams; rare

Lednia Ricker

61(57'). 61'.

Cerci elongate, sclerotized or with subapical spines(Fig. 16.249) Cerci unmodified and poorly sclerotized (Fig. 16.248)

62(59).

Cerci with subapical spines (Fig. 16.253); abdominal segments uniformly sclerotized and about equal in width; apex of abdominal sternum 9 only slightly produced (similar to Fig. 16.250);

62'.

Cerci elongate, curved and without subapical spines; anterior abdominal segments weakly sclerotized and narrower than abdominal segments 9 and 10; apex of abdominal segment 9 produced into a long probe which extends almost to, or beyond tips of cerci (Fig. 16.249); Eastern

Eastern and Western North America

62 63

Nemoura Latreille

North America

Ostrocerca Ricker

63(61').

Body of epiproct a long, simple, completely sclerotized, acute probe with median suture (Fig. 16.248), sometimes bearing sclerotized basolateral arms; widespread Prostoia Ricker

63'.

Body of epiproct a short to moderately long, complexly lobed, incompletely sclerotized, blunt probe (Fig. 16.254)

64

64(63').

Abdominal tergum 10 without long curved lobes; shorter bulbous lobes may partially cover cereal

64'.

Abdominal tergum 10 bearing a pair of long curved lobes which cover small, sub-ventrally located cerci completely in dorsal aspect (Figs. 16.254 and 16.255); Eastern North America Shipsa Ricker (one species, 5'. rotunda (Claassen)) Forewing terminal costal crossvein joins Sc in apical costal space (Fig. 16.247); Eastern North

bases

65(58').

America, rare

Podmosta Ricker

Paranemoum Needham and Claassen

65'.

Forewing terminal costal crossvein joins R in apical costal space (Fig. 16.251)

66(65').

Abdominal sternum 7 covers most or all of sternum 8(Fig. 16.256); cerci lightly sclerotized and apically truncate; widespread Nemoura Latreille

66'.

Abdominal sternum 7 covers less than half of sternum 8; cerci membranous and apically rounded

67(65').

Subgenital plate of abdominal sternum 8 covers most of abdominal sternum 9(Fig. 16.257); abdominal sternum 7 sometimes with median nipple-like projection; Eastern North America

67'.

66

67

Ostrocerca Ricker

68(67').

Subgenital plate of abdominal sternum 8 covers little if any of abdominal sternum 9; abdominal sternum 7 without median nipple-like projection 68 Abdominal sternum 7 slightly enlarged, covering base of abdominal sternum 8 69

68'.

Abdominal sternum 7 not extending over base of abdominal sternum 8

70

Figure 16.236

Figure 16.240

Figure 16.237

Figure 16.238

Figure 16.239

Figure 16.242

Figure 16.241

Figure 16.244

paraproct vesicle

Figure 16.245 Figure 16.243

Figure 16.236 Mesocapnia arizonensis (Capniidae)

Figure 16.242 Amphlnemura delosa (Nemourldae)

adult female termlnalla, ventral. Modified from

adult female termlnalla, ventral.

Baumann etal.(1977). Figure 16.237 Botshecapnia sasquatchi(Capniidae)

adult male termlnalla, lateral. Modified from Baumann

Figure 16.243 Zapada oregonensis (Nemourldae)

Figure 16.238 Maienka coloradensis (Nemourldae) adult prothoracic sternum and cervical gill remnants. Figure 16.239 Maienka coloradensis (Nemourldae)

(1975). Figure 16.244 Zapada oregonensis (Nemourldae) adult male paraproct, caudal. Modified from Baumann (1975). Figure 16.245 Zapada oregonensis (Nemourldae)

adult male termlnalla, lateral.

adult female termlnalla, ventral. Modified from

Figure 16.240

Baumann (1975).

adult female termlnalla, ventral. Modified from Baumann etal. (1977).

Maienka coloradensis (Nemourldae)

adult female termlnalla, ventral.

Figure 16.241

Amphlnemura delosa (Nemourldae)

adult male termlnalla, lateral.

475

476

Chapter 16 Plecoptera

69(68).

Abdominal sternum 8 with median membranous band and small subgenital plate which projects over base of abdominal sternum 9(Fig. 16.258) Shipsa Richer (one species, S. rotunda (Claassen))

69'.

Abdominal sternum 8 with median pigment band and without projecting plate (Fig. 16.259)

70(68').

Podmosta Kicktx

Abdominal sternum 8 slightly produced and bearing a median dark submarginal spot and two lateral pigment spots on plate margin (Fig. 16.260); Western North America glacial or snowmelt waters; rare

Lednia Richer

70'.

Abdominal sternum 8 not produced, bearing median dark spot or uniformly pigmented on posterior margin; Eastern and Western North America, common Prostoia Richer

71(4').

Flind wing with 6 anal veins; apical costal space of forewing usually pale proximally and dark distally forming distinctive stigma (Fig. 16.261); female with ovipositor projecting well beyond abdominal tip (Fig. 16.262); uncommon in Western and rare in Eastern North America

Megaleuctra Neave

71'.

Hind wing with 3-4 anal veins; forewing apical costal space pigment pattern similar to rest of wing; female without ovipositor, common 72

72(71').

Rs vein originates from R vein at or very near arculus and above origin of M vein (Fig. 16.263); female abdominal segment 10 complete ventrally (Fig. 16.264); Western North America Perlomyia Banks

72'.

Rs vein originates from R at a distance removed from the arculus by at least half arculus length (Fig. 16.265); female abdominal segment 10 interrupted ventrally (Fig. 16.266)

73

73(72').

Apex of abdomen with paraprocts modified into one or more long, upwardly curved probes (Fig. 16.267); vesicle usually present on abdominal sternum 9 Males

74

73'.

Apex of abdomen with simple, broadly triangular, flat paraprocts (Figs. 16.266); vesicle absent from abdominal sternum 9

Females

74(73).

80

A deep U or V-shaped cleft extends across abdominal tergum 9 (Fig. 16.268); Eastern North America

Zealeuctra Richer

74'.

Abdominal tergum 9 without deep U or V-shaped cleft

75

75(74').

Paraprocts consist of 2 or 4 closely appressed, long stylets (Figs. 16.182 and 16.269)

76

75'.

Paraprocts consist of single thick probe like structure (Fig. 16.267)

78

76(75).

Paraprocts consist of 4 stylets (Figs. 16.182 and 16.269); tergal lobes usually present on segments 7, 8, or both (Fig. 16.269); Eastern North America Leuctra Stephens

76'.

Paraprocts consist of 2 stylets (Fig. 16.270); tergal lobes absent from segments 7 and 8; Western North America, especially Pacific Northwest 77

77(76').

Vesicle absent from abdominal sternum 9; paraprocts gradually tapered to tip; cord crossvein located in costal space beyond end of Sc vein (Fig. 16.265); Western North America Despaxia Richer (one species, D. augusta (Banks))

77'.

Small vesicle present on abdominal sternum 9; paraprocts wide in lateral aspect near tip, but abruptly narrowed subapically (Fig. 16.271); cord crossvein connects end of Sc with R vein; Western North America Moselia Richer

78(75').

Posterior margin of abdominal tergum 9 modified as a sclerotized, multi-toothed plate, or with a pair of sclerotized lobes (Fig. 16.272); vein Cuj of hind wing without fork; CA,rare

78'.

79(78').

Calileuctra Shepard and Baumann

Posterior margin of abdominal tergum 9 unmodified; vein Cui of hind wing forked (Fig. 16.273)

79 Male cerci with thumb-shaped basoventral lobe (Fig. 16.274); mesothoracic furcasternum without median dark line; metathoracic presternum divided (Fig. 16.275); Western North America

Pomoleuctm Stark and Kyzar

Chapter 16 Plecoptera

477

joined A, & Figure 16.246

costal crossvein

Figure 16.248 sternum 9 apex

Figure 16.247

costal crossvein

Figure 16.251

Figure 16.249

subapical

spiny lobes

spine

Figure 16.250

Figure 16.252

Figure 16.246 Soyedina washingtoni(Nemouridae) forewing. Figure 16.247 Paranemoura perfecta (Nemouridae) forewing. Figure 16.248 Prostoia besametsa (Nemouridae)

Figure 16.253

Figure 16.250 Prostoia besametsa (Nemouridae) male adult termlnalia, ventral.

Figure 16.251 Lednia tumana (Nemouridae)forewing. Figure 16.252 Lednia tumana (Nemouridae) adult male termlnalia, lateral. Modified from Baumann (1975).

adult male termlnalia, dorsal.

Figure 16.253 Nemoura sp.(Nemouridae) adult male

Figure 16.249 Ostrocerca albidipennis (Nemouridae)

termlnalia, dorsal. Modified from Baumann (1975).

adult male termlnalia, dorsal.

v,;'.vv' sensiiia

■ "I

patches

Figure 16.317

■!'^ •■ •• K

rt.

:- 'r

Figure 16.319 Figure 16 .318

Figure 16.309 Rasvena terna (Chloroperlldae) adult female abdominal segments 7 and 8, ventral. Figure 16.310 Suwallia starki (Chloroperlldae) adult male terminalla, dorsal.

Figure 16.311

Sweltsa fidelis (Chloroperlldae) adult

male terminalla, lateral.

Figure 16.312

Sasquaperia hoopa (Chloroperlldae)

adult male terminalla, dorsal.

Figure 16.313

Sasquaperia hoopa (Chloroperlldae)

adult male mesosternum.

Figure 16.314

Bisancora rutriformis (Chloroperlldae)

adult mesosternum.

488

Figure 16.315

Bisancora rutriformis (Chloroperlldae)

adult male terminalla, dorsal.

Figure 16.316 Bisancora rutriformis (Chloroperlldae) adult male oblique lateral aspect epiproct. Figure 16.317 Paragnetina immarginata (Perlldae) adult male terminalla, dorsal.

Figure 16.318

Acroneuria coveili (Perlldae) adult

male terminalla.

Figure 16.319

Neoperia coosa (Perlldae) adult male

terminalla, dorsal.

gill remnants hemitergum hammer

ammer

Figure 16.323

Figure 16.321 Figure 16.320 hammer

hemitergum

Figure 16.322

Figure 16.324

Figure 16.326

it'®.ilysK#

...

..^.^.^^ sensilla row ^

^ Figure 16.325

Figure 16.327

...-a.-'.'. ... Figure 16.328

Figure 16.320 Claassenia sabulosa (Perlidae) adult

Figure 16.325 Doroneuria theodora (Perlidae) adult

male terminalia, ventral.

male terminalia, ventral.

Figure 16.321

male terminalia, lateral.

Figure 16.326 Perlinella drymo (Perlidae) forewlng. Figure 16.327 Perlinella drymo (Perlidae) adult aedeagal sclerite. Figure 16.328 Eccoptura xanthenes (Perlidae) adult

Figure 16.323 Anacroneuria litura (Perlidae) adult

male terminalia, dorsal.

Claassenia sabulosa (Perlidae) adult

male terminalia, dorsal.

Figure 16.322 Agnetina annulipes (Perlidae) adult

male abdominal sternum 9.

Figure 16.324 Hesperoperia pacifica (Perlidae) adult male terminalia, ventral.

489

490

122'.

Chapter 16 Plecoptera

Three ocelli

124

123(122). Subgenital plate covers half or more of abdominal sternum 9(Fig. 16.336); abdominal sternum 9 with distinctive pattern of setae and sclerites; egg spindle-shaped, chorion smooth; spermathecal stalk membranous; AZ,TX, Mexico Anacwneuria Klapalek 123'. Subgenital plate unproduced, or slightly produced as a small tab like structure only slightly extending to abdominal sternum 9(Fig. 16.337); abdominal sternum 9 unmodified; egg oval to barrel-shaped, chorion striate, punctate or smooth; spermathecal stalk lined with brown microsetae; Eastern North America Neopevla Needham 124(122'). Subgenital plate not covering abdominal sternum 9(Fig. 16.338) 125 124'. Subgenital plate extends over at least base of abdominal sternum 9(Fig. 16.339) 128 125(124). Posteromesal margin of abdominal sternum 8 bearing slight emargination surrounded by U or V-shaped pale area (Fig. 16.338); egg collar usually stalked (Fig. 16.340) .. Calineuria Ricker(one species, C. californica (Banks))

125'.

Posteromesal margin of abdominal sternum 8 with or without emargination, but if present, not surrounded by V-shaped membranous area; egg collar reduced to small rounded button (Fig. 16.341)

126

126(125') Mesal field of abdominal sternum 8 not clearly offset by lateral membrane (Fig. 16.342); abdominal sternum 10 with wide anteromesal field of microtrichia; vagina lined along lateral margins with brown microsetae, accessory glands present on anterolateral angles; egg oval with base and collar ends generally similar in width (Fig. 16.341) . .Doroneuria Needham and Claassen 126' Mesal field of abdominal sternum 8 clearly offset by lateral membrane (Fig. 16.343); abdominal sternum 10 without field of microtrichia; vagina with or without brown microsetal lining, if lining is present, anterolateral accessory glands are absent; egg spindle-shaped with base much narrower than collar end (Fig. 16.344)

127

127(126'). Dense microtrichia patch present in posterior membrane of abdominal sternum 9; vagina sparsely lined and anterolateral accessory glands present (similar to Fig. 16.345); most of egg chorion smooth but basal end covered with follicle cell impressions (Fig. 16.344) .... Claassenia Wu(one species, C. sabulosa (Banks))

127'.

Microtrichia patch absent from membrane of abdominal sternum 9; vagina completely lined with brown microsetae and anterolateral accessory glands absent; egg chorion entirely smooth Acroneuria Pictet(in part)

128(124'). Subgenital plate with a distinct notch (Fig. 16.346)

129

128'.

134

Subgenital plate truncate or entire

129(128). Egg collar stalked (Fig. 16.347)

130

129'. Egg collar absent, sessile or button like (Figs. 16.341 and 16.348) 131 130(129). Accessory glands present on anterolateral corners of vagina (Fig. 16.345); large species, forewing length 20-35 mm Paragnetina Klapalek 130'. Accessory glands absent from anterolateral corners of vagina; small to medium species, forewing length 8-16 mm Perlesta Banks (in part) 131(129'). Forewing length at least 20 mm

132

131'.

133

Forewing length at most 18 mm

132(131). Subgenital plate reaches beyond midpoint of abdominal sternum 9, notch as wide as lateral lobes

of plate (Fig. 16.349); pale yellow species

Eccoptum Klapalek

(one species, E. xanthenes(Newman))

132'.

Subgenital plate usually not reaching midpoint of abdominal sternum 9, notch variable, but not usually as wide as lateral lobes of plate; brown to dark brown species. . Acroneuria Pictet (in part)

133(131'). Subgenital plate reaches midpoint of abdominal sternum 9, its outline triangular (Fig. 16.339); egg lacking collar; Southern Appalachians or Piedmont

Beloneuria Needham and Claassen

paraprocts

hammer

.1

Figure 16.334 Figure 16.329

Figure 16.330 membranous

membranous

notch

iiSKiiP pMw

Figure 16.335

.'S'-*- ■■ ■ ■ ■ Figure 16.332 Figure 16.331

Figure 16.336

Figure 16.333

f/';" f»8

Si Figure 16.337

Figure 16.329

V

1^4? Figure 16.338

Beloneuria georgiana (Perlidae) adult

male termlnalia, ventral.

Figure 16.330

Beloneuria georgiana (Perlidae) adult

male termlnalia, dorsal.

Figure 16.331

Doroneuria theodora (Perlidae) adult

male termlnalia, dorsal.

Figure 16.332

t ./••■ - ivi: /

Caiineuria caiifornica (Perlidae) adult

Figure 16.339

Figure 16.335

Periineiia drymo (Perlidae) adult

female termlnalia, ventral.

Figure 16.336

Anacroneuria lltura (Perlidae) adult

female termlnalia, ventral.

Figure 16.337

Neoperia stewarti (Perlidae) adult

female termlnalia.

Figure 16.338

Caiineuria caiifornica (Perlidae) adult

male termlnalia, dorsal.

female termlnalia, ventral.

Figure 16.333 Figure 16.334

female termlnalia, ventral.

Neoperia ciymene (Perlidae) forewing. Hansonoperia appaiachia (Perlidae)

adult female termlnalia, ventral.

Figure 16.339

Beloneuria georgiana (Perlidae) adult 491

492

133'.

Chapter 16 Plecoptera

Subgenital plate usually covers only basal third of abdominal sternum 9; posterior margin of plate usually truncate, if triangular, then egg collar button like or short stalked; small to medium-sized species; widespread Perlesta Banks (in part)

134(128'). Subgenital plate with transverse mesal tubercle (Fig. 16.350); egg chorion completely covered with coarse follicle cell impressions except for smooth narrow annulus near each end (Fig. 16.348); note: in some eggs an anchor ring is attached covering one annulus; Eastern North America; rare Attaneuria Ricker (one species, A. ruralis(Hagen))

134'. Subgenital plate without tubercle; egg chorion with, at most, a single annulus,common 135 135(134'). Abdominal sternum 9 having large mesal membranous area enclosing a small microtrichia patch, paraprocts membranous on inner margins, subgenital plate often with dark sclerite imbedded in membrane plate surface (Fig. 16.351); egg body flanged around base of collar (Fig. 16.352); Western North America Hesperoperla Banks 135'. Abdominal sternum 9 lacking mesal membranous area and microtrichia patch; paraprocts uniformly sclerotized; subgenital plate without dark sclerite; egg without flange around base of collar

136

136(135'). Thoracic sterna 2-3 with transverse bands of dark pigment; vagina with anterolateral accessory glands and one or two pairs of internal sclerites; Eastern North America Agnetina Klapalek 136'. Thoracic sterna 2-3 pale yellow to brown; vagina without accessory glands or sclerites 137 137(136'). Forewing length . ■j ':-*\i.-'

Figure 16.374

Figure 16.373

Figure 16.376

paraproct

?f i.

f ■•.1'".I/-'.i-.i;//)-■-. ■,

^ yMt' Figure 16.375

W:W0M

m7 0m'mi f'j'* W • .&'-'0-M-'''-'-'-:'''f--0'i^ i

.>.3.-''v-' i"Vi >■•.•- •>>:■,V.

.w

I

Figure 16.378

Figure 16.377 spur vein

Figure 16.379

Figure 16.380

Y-arm •

Figure 16.381

Figure 16.373

Calliperia luctuosa (Perlodidae) adult

Figure 16.378

Diura knowltoni (Perlodidae) adult

male termlnalla, dorsal.

male termlnalla, dorsal.

Figure 16.374 Cascadoperia trictura (Perlodidae)

Figure 16.379 forewlng. Figure 16.380

adult male termlnalla, dorsal.

Figure 16.375 Cosumnoperia hypocrena (Perlodidae) adult male termlnalla, dorsal. Figure 16.376 Clioperia clio (Perlodidae) adult male termlnalla, dorsal. Modified from Szczytko and Stewart (1981). Figure 16.377 Diploperia robusta (Perlodidae) adult male termlnalla, ventral. 500

Oconoperia innubila (Perlodidae) Yugus bulbosus (Perlodidae) adult

male abdominal sternal 7 and 8. Modified from

Nelson (2001). Figure 16.381 Diploperia robusta (Perlodidae) adult male mesosternum.

;.i.

,~.vHC

liy[

Figure 16.383

lAm

Figure 16.384

Figure 16.382

paragenital

epiproct

lobe

epiproct

paragenital plate

coiled band

Figure 16.385

Figure 16.386

paragenital plate

basal fold

ii.*n

iPiiiii Figure 16.388 Figure 16.390

Figure 16.387 basal

intercubital crossveins

fold

intercubital crossveins

Figure 16.389

Figure 16.382 Malirekus hastatus (Perlodidae) adult

Figure 16.391

Figure 16.387 Kogotus modestus (Perlodidae) adult

head and pronotum.

male terminalia, dorsal.

Figure 16.383 Malirekus hastatus (Perlodidae) adult

Figure 16.388 Diura knowltoni(Perlodidae) adult

male termlnalia, lateral.

female sterna 8 and 9, ventral.

Figure 16.384 Cultus aestivalis (Perlodidae) adult head and pronotum.

Figure 16.389 Diura knowltoni(Perlodidae) forewing. Figure 16.390 Diura washingtoniana (Perlodidae)

Figure 16.385 Kogotus modestus (Perlodidae) adult

adult head and pronotum. Modified from Nelson and

male epiproct complex, lateral.

Nelson (2018).

Figure 16.386 Baumannella alameda (Perlodidae)

Figure 16.391 Isoperia fulva (Perlodidae) adult

adult male terminalia, dorsal.

female terminalia, ventral.

502

Chapter 16 Plecoptera

179'.

Suture lines behind ocelli no darker than background pigment of ocellar triangle; course of A2 vein of forewing relatively straight Yugus Ricker 180(177'). Egg triangular in cross section; Rs vein of forewing usually with basal spur (Fig. 16.379); Southern Appalachians, rare Oconoperla Stark and Stewart(one species, O. innubila(Needham and Claassen))

ISC'.

Egg circular in cross section (Fig. 16.393) or turtle-shaped (Fig. 16.396); Rs vein of forewing without basal spur

181

181(180'). Membranous folds at base of subgenital plate lie parallel to lateral margins of abdominal sternum 8(Fig. 16.391); egg generally circular in cross section, not turtle-shaped. . Isoperla Banks (in part) 181'. Membranous folds at base of subgenital plate angled toward center of abdominal sternum 8 (Fig. 16.397); egg turtle-shaped (Fig. 16.396) 182 182(181'). Head background color pale with pale to dark brown area completely covering ocellar triangle; forewing length 9-12 mm; Eastern North America Remenus Ricker

182'.

Head with wide, dark M-line extending across ocellar area, but ocellar triangle mostly pale (Fig. 16.384); forewing length 13-14 mm; widespread Cultus Ricker (in part) 183(172'). Subgenital plate a minute, mesoposterior lobe (Fig. 16.398); head pattern with an almost complete dark longitudinal band, darkest in ocellar region (Fig. 16.399); Western North America, mostly Pacific Northwest Cascadoperla Szczytko and Stewart(one species, C. trictura(Hoppe)) 183'. Subgenital plate at least half as wide as mesal sclerite of abdominal sternum 8; head pattern variable but without complete dark longitudinal band 184(183'). Subgenital plate notched, truncate or emarginate 184'. Subgenital plate rounded or triangular 185(184). Subgenital plate truncate (Fig. 16.388) 185'. Subgenital plate notched or emarginate (Fig. 16.400) 186(185). Membranous folds at base of subgenital plate angled toward center of abdominal sternum 8

186'.

184 185 190 186 187

(Fig. 16.388); usually at least 15 total median and intercubital crossveins in forewing (Fig. 16.389); dark pigment extends to, or near lateral pronotal margins (Fig. 16.390); fine setae on pale areas of thoracic sterna near coxae surrounded by obscure pale brown basally; Western and higher latitudes of Eastern North America Diura Billberg Membranous folds at base of subgenital plate lie parallel to lateral margins of abdominal segment 8(Fig. 16.391); usually less than 15 total median and intercubital crossveins in forewing; dark pigment on pronotum usually not approaching lateral margins(Fig. 16.401); fine setae on pale areas of thoracic sterna without basal brown rings Isoperla Banks(in part)

187(185'). Subgenital plate covers less than a third of abdominal sternum 9

188

Subgenital plate covers half or more of abdominal sternum 9(Fig. 16.402) 189 Head with striking pattern of dark brown and yellow including dark brown posterolateral margins and quadrangular dark ocellar spot with finger-shaped extensions lateral to posterior ocelli (Fig. 16.403); sternum 10 with large mesal membranous area; usually at least 15 total median and intercubital crossveins in forewing (as in Fig. 16.389); CA,OR, WA Calliperla Banks(one species, C. luctuosa (Banks)) 188'. Head pattern variable but not as above; sternum 10 without mesal membranous area; usually less than 15 total median and intercubital crossveins in forewing, widespread Isoperla Banks(in part) 189(187'). Membranous folds at base of subgenital plate angled toward center of sternum 8(Fig. 16.402); egg an oval, biconcave disc without collar (Fig. 16.404); CA ....Baumannella Stark and Stewart(one 187'.

188(187).

species, B. alameda(Needham and Claassen)

189'.

Membranous folds at base of subgenital plate extend parallel to lateral margins of abdominal

sternum 8 (Fig. 16.391); egg variable but not biconcave, collar present or absent 190(189'). Usually at least one radial crossvein beyond cord (Fig. 16.405); mesobasisternum with short median groove extending forward from transverse groove (Fig. 16.406); subgenital plate a

190

i Figure 16.393 Figure 16.394

Figure 16.392

Figure 16.395

Figure 16.396

Figure 16.397

!«¥i !iWi5VJ'-;t--^->.vfji>!.4i^ '-M ■■ ■•«

Figure 16.400 Figure 16.398

Figure 16.399

Figure 16.401

Figure 16.402

Figure 16.392 Clioperia clio (Perlodidae) adult head and pronotum. Modified from Szczytko and Kondratieff (2015). Figure 16.393 Isoperia fulva (Perlodidae) egg and inset of collar. Modified from Szczytko and Stewart (1979). Figure 16.394 Malirekus hastatus (Perlodidae) adult female terminalia, ventral.

Figure 16.395 Malirekus hastatus (Perlodidae) forewing. Figure 16.396 Cultus verticalis (Perlodidae) egg. Modified from Kondratieff (2004). Figure 16.397 Cultus aestivalis (Perlodidae) adult female terminalia, ventral.

Figure 16.403

Figure 16.398

Cascadoperla trictura (Perlodidae)

adult female terminalia, ventral.

Figure 16.399 Cascadoperla trictura (Perlodidae) adult head and pronotum. Modified from Szczytko and Stewart (1979). Figure 16.400 Calllperia luctuosa (Perlodidae) adult female abdominal sterna 8 and 9.

Figure 16.401 Isoperia fulva (Perlodidae) adult head and pronotum. Modified from Szczytko and Stewart (1979). Figure 16.402 Baumannella alameda (Perlodidae) adult female terminalia, ventral.

Figure 16.403

Calllperia luctuosa (Perlodidae) adult

head, dorsal.

503

504

Chapter 16 Plecoptera

radial crossvein

cord crossvein

Figure 16.405 r-m crossve n

Figure 16.404

Figure 16.410 median

groove

M.

Figure 16.409 Figure 16.406

Figure 16.407

Figure 16.404 Baumannella alameda (Periodidae) egg. Modified from Stark and Stewart (1985). Figure 16.405 Cosumnoperia hypocrena (Periodidae) forewing. Figure 16.406 Cosumnoperia hypocrena (Periodidae) adult mesosternum. Figure 16.407 Cosumnoperia sequoia (Periodidae) adult female termlnalia, ventral. Modified from Bottorff

(2007).

Figure 16.408

pigment bar

Figure 16.408 Kogotus modestus (Periodidae) adult female termlnalia, ventral.

Figure 16.409 Pictetlella expansa (Periodidae) egg. Modified from Baumann (1973). Figure 16.410 Pictetlella expansa (Periodidae) forewing.

Chapter 16 Plecoptera

505

narrow triangle extending to abdominal sternum 10(Fig. 16.407); egg green in life, about 0.8 mm long and without collar; CA Cosumnoperla Szczytko and Bottorff 190'.

Usually without radial crossveins beyond cord; mesobasisternum without median groove; egg variable but typically brown or pale brown in life; less than 0.5 mm long, with or without collar; widespread Isoperla Banks (in part)

191(184'). Membranous folds at base of subgenital plate lie parallel to lateral margins of abdominal sternum

8 (Fig. 16.391); egg generally circular in cross section (Fig. 19.393) Isoperla Banks (in part) 191'. Membranous folds at base of subgenital plate angled from plate base toward center of abdominal sternum 8 (Fig. 16.408); egg turtle-shaped(Fig. 16.396)or triangular in cross section 192 192(189'). Subgenital plate reaches midpoint or less of abdominal sternum 9(Fig. 16.408); base of plate with a pair of large, oblong pigment bars extending from membrane fold to basal area of abdominal segment 8; Western North America Kogotus Ricker 192'. Subgenital plate reaches beyond midpoint of sternum 9; base of plate usually without pigment bars as above; widespread 193 193(192'). Egg crudely 3-sided with keel, collar covered on one side by rim (Fig. 16.409); subgenital plate usually about as wide as abdominal sternum 8; forewing usually with a crossvein beyond r-m crossvein (Fig. 16.410); Western North America Pictetiella lilies 193'. Egg turtle-shaped without keel(Fig. 16.396); subgenital plate distinctly narrower than base of abdominal stemum 8(Fig. 16.397);forewing usually without crossvein beyond r-m crossvein 194

194(193'). Subgenital plate reaches beyond posterior margin of abdominal sternum 9; ocelli connected by narrow dark pigment bands, CA, NV, OR, WA

Rickera Jewett

(one species, R. sorpta (Needham and Claassen))

194'.

Subgenital plate not reaching beyond posterior margin of abdominal sternum 9(Fig. 16.397); ocelli usually connected by broad dark pigment bands (Fig. 16.384); widespread Cultus Ricker

ADDITIONAL TAXONOMIC REFERENCES General Needham and Claassen (1925), Claassen 1931), Ricker (1952), Stark et al.(1999), Stewart and Stark (2002), DeWalt et al. (2015), Plecoptera Species File (DeWalt et al. 2018) Perla, Annual Newsletter and Bibliography of the International Society of Plecopterologists publishes the world literature of the Plecoptera annually. All newsletters dating to 1974 are available from Plecoptera Species File (DeWalt et al. 2018, http;//plecoptera,speciesfile.org)

Regional faunas Alabama: Grubbs and Sheldon (2018) Alaska: Stewart and Oswood (2006)

Pacific Northwest: Jewett (1959).

Pennsylvania: Surdick and Kim (1976). Rocky Mountains: Baumann et al.(1977). Saskatchewan: Dosdall and Lehmkuhl (1979).

South Carolina: McCaskill and Brigham (1982), Stark (2017), Southeastern USA: Stark (2017).

Texas: Szczytko and Stewart(1977). Utah: Gaufin et al.(1966), Baumann et al.(1977), Baumann and Unzicker (1981), Houseman and Baumann (1997), Call and Baumann (2002). Washington: Hoppe(1938). Wisconsin: Hilsenhoff (1981), Yukon: Stewart and Ricker (1999).

Regional species lists

Armitage (2004). Florida: Stark and Gaufin (1979), Pescador et al.(2000). Illinois: Prison (1935), Webb (2002) Indiana: DeWalt and Grubbs(2011). Louisiana: Stewart et al.(1976). Minnesota: Harden and Mickel (1952). Montana: Gaufin et al.(1972), Gaufin and Ricker (1974). Nevada: Baumann et al.(2017).

Alabama: Stark and Harris(1986), Grubbs(2011). Alaska: Jewett(1971): Ellis(1975). California (Capniidae): Nelson and Baumann (1987a). Canadian Far North: Ricker (1944). Canadian Maritime Provinces: Ricker (1947), Brinck (1958), Kondratieff and Baumann (1994). Canadian Prairie Provinces: Ricker (1946). Colorado: Stark et al.(1973), Kondratieff and Baumann (2002), Zuellig et al.(2006), Delaware: Lake (1980). Florida: Berner (1948), Stark and Gaufin (1979). Georgia: Verdone et al.(2017). Idaho: Newell and Minshall (1976). Illinois: DeWalt et al.(2005), DeWalt and Grubbs(2011), Webb (2002). Indiana: Ricker (1945), Bednarik and McCafferty (1977), Grubbs (2004), DeWalt and Grubbs(2011). Iowa: Heimdal et al.(2004), Heimdal and Birmingham (2006). Kansas: Stewart and Huggins(1977).

North Carolina: McCaskill and Brigham (1982), Stark (2017).

Kentucky: Tarter et al. (1984), Pond (1999), Tarter and Chaffee

Alberta: Ricker (1946), Stewart and Oswood (2006), Dosdall and Giberson (2014a), Dosdall and Giberson (2014b). Black Hills: Huntsman et al.(1999) British Columbia: Ricker (1943), Ricker and Scudder (1975), Stewart and Oswood (2006), Baumann and Stark (2010), Dosdall and Giberson (2014a), Dosdall and Giberson (2014b). California: Jewett(1956, 1960). Canada: Ricker (1964) Connecticut: Hitchcock (1974).

Eastern North America: Stark and Armitage (2000), Stark and

Ozark and Ouachita Mountains: Poulton and Stewart (1991).

(2004), Tarter et al.(2006), Tarter et al.(2015).

506

Chapter 16 Plecoptera

Maine: Mingo (1983). Manitoba: Burton (1984). Maryland: Grubbs(1997), Duffield and Nelson (1990). Michigan: Grubbs and Bright(2001), Grubbs et al.(2012). Minnesota: Lager etal.(1979).

Mississippi: Stark (1979^ Stark and Hicks(2003). Mount Rainier National Park: KondratiefT and Lechleitner(2002). New Jersey: Earle (2009). Nebraska: Rhodes and KondratiefT (1996). Nevada: Gather et al.(1975), Baumann et al.(2017).

New Mexico: Stark et al.(1975), Jacobi and Baumann (1983), Jacobi et al.(2005). New York: Myers et al.(2011).

North Carolina: KondratiefT et al.(1995), Stark (2017). North Dakota: KondratiefT and Baumann (1999). Nova Scotia: Ogden et al. 2018

Ohio: Walker (1947), Gaufin (1956), Tkac and Foote (1978), DeWalt et al.(2012), DeWalt et al.(2016).

Oklahoma: Stark and Stewart(1973). Ontario: Harper and Ricker (1994). Ozark and Ouachita Mountains(Neoperla): Ernst et al.(1986), Poulton and Stewart (1991).

Pennsylvania: Lager et al.(1979), Masteller (1996), Grubbs (1996), Earle (2004). South Carolina: McCaskill and Prins (1968), KondratiefT et al. (1995).

South Dakota: Huntsman et al.(2001). Southwestern United States: Stewart et al.(1974). Virginia: KondratiefT and Voshell (1979), KondratiefT and Kirchner (1987), KondratiefT et al.(2017). Western Intermountain Area: Gaufin (1964), Logan and Smith (1966).

West Virginia: Tarter and Kirchner(1980), Tarter and Nelson(2006)

Taxonomic treatments at the family and generic levels(N = nymphs;A = adults) Capniidae: Allocapnia(Harper and Hynes 1971b-A, N, Ross and Ricker 1971-A, Stark and Lacey 2005-A, N, Stark and KondratiefT 2012-A; Webb 2002-A, N); Anapnia (Muranyi et al. 2014-A, Nelson and Baumann 1989-A); Bohhecapnia (Baumann and Potter 2001-A); Capnia(Muranyi et al. 2014-A, Nelson and Baumann 1989-A); Capnura(Nelson and Baumann 1987b-A); Eucapnopsis(Baumann et al. 1977-A); Isocapnia (Zenger and Baumann 2004-A); Mesocapnia(Baumann and Gaufin 1970-A); Nemocapnia (Stark et al. 2016-A); Paracapnia (Stark and Baumann 2004-A); Sierraeapnia (Bottorff and Baumann 2015-A); Utacapnia(Nebeker and Gaufin 1965-A).

Chloroperlidae: Ala.tkaperla (Stewart et al. 1991-A, N); Alloperla (Baumann and KondratiefT 2009-A, Hitchcock 1968-A, 1974-A, Lyon and Stark 1997-A, Stark and KondratiefT 2010-N, Stark 2017-N, Surdick 2004-A, Willett and

Stark 2009-A); BLtancora (Surdick 1981-A, Stewart and Stanger 1985); Haploperla (Hitchcock 1974-A, Surdick 2004-A ); Kathroperla (Stark et al. 2015-A); Neaviperla (Alexander and Stewart 1999-A, Baumann and Lee

2014-A); Paraperla (Stark et al. 2013-A, N); Plumiperla (Surdick 1985-A); Rasvena (Surdick 2004-A); Sa.squaperla Stark and Baumann 2001-A, N); Suwallla (Alexander and Stewart 1999-A, Surdick 2004-A); Sweltsa(Nye and Stark 2010-A, Surdick 1995, Surdick 2004-A); Triznaka (Baumann and KondratiefT 2008, KondratiefT and

Baumann 2012-A); C/tapcr/a(Baumann etal. 1977-A, N; Harper and Roy 1975-A, Surdick 2004-A). Leuctridae: Calileuctra (Shepard, W. D. and R. W. Baumann. 1995-A, Stewart et al. 2013-N); Despaxia(Baumann et al. 1977-A, KondratiefT and Lechleitner 2002-A); Leuctra (Grubbs 2015-A, Harper and Hynes 1971-N; Harper and Harper 2003-A Hitchcock 1974-A,, Harrison and Stark 2010-A, N); Megaleuctra(Baumann and Stark 2013-A); Mo.selia(Stark and Harrison 2016-A); Paraleuctra (Stark and Kyzar 2001-A); Perlomyia(Baumann et al. 1977-A,

Nelson and Hanson 1973-A); Pomoleuctra (Stark and Kyzar 2001-A); Zealeuctra(Grubbs et al. 2013). Nemouridae: Amphinemura(Baumann and Gaufin 1972.-A, Harper and Hynes 1971d-N, Hitchcock 1974-A; Baumann 1996-A, Boumans and Baumann 2012-A); Lednia(Baumann and KondratiefT 2010-A); Malenka(Ricker 1952-A; Jewett 1959-A, Baumann et al. 1977); Nanonemoura(Baumann and Fiala 2001-A, N); Nemoura (Hitchcock 1974-A, Stewart and Oswood 2006-A); Ostrocerca(Harper and Hynes 1971d-N, Hitchcock 1974-A; Young etal. 1989-A); Paranemoura(Baumann 1996-A); Podmosta(Jewett 1959-A, Baumann etal. 1977-A, Hitchcock 1974-A, Stewart and Oswood 2006, Stewart and Stark 2011 -N); Prostoia (Grubbs era/2014-A); Shipsa(Harper and Hynes 1971d-N, Hitchcock 1974-A); Soyedina (Ricker 1952-A; Baumann and Grubbs 1996-A; Grubbs 2006-A); VLsoka(Baumann et al. 1977-A); Zapada(Baumann et al. 1977-A; Grubbs et al. 2015-A). Peltoperlidae: Peltoperla (Stark 2000-A); Sierraperla (Stark etal. 2015-A); Soliperla (Stark and GustaTson 2004-A); Tallaperla (Stark 2000-A); Viehoperla (Stark 2000-A), Yoraperla (Stark and Nelson 1994-A). Pteronarcyidae: Pteronarcella(Baumann et al. 1977-A, N); Pteronarcys(Baumann et al. 1977-A, N, Nelson 2000-A, Myers and KondratiefT 2017-N).

Perlidae: Aewneuria (Stark 2004-A); Agnetina(Stark 2004-A); Anacroneuria (Stark and KondratiefT 2004-A); Belonewia (Stark 2004-A); Calineuria(Baumann et al. 1911-A, Stark and Gaufin 1974-A); Claassenia(Baumann etal. 1977-A; Stark and Sivec 2010-A); Doroneuria(Baumann et al. 1911-A, Stark and Gaufin 1974-A); Eecoptura (Stark 2004-A); Hansonoperla(KondratiefT and Kirchner 1996-A; Stark 2004-A); Hesperoperla(Baumann et al. 1977-A; Baumann and Stark 1980-A, N); Neoperla(Stark 2004-A); Paragnetina (Stark 2004-A); Perlesta (Stark 2004-A); Perlinella (KondratiefT et al. 1988-A, N; Stark 2004-A). Perlodidae: Arcynopteryx (Teslenko 2012-A); Baumannella (Stark and Stewart 1985-A, N); Calliperla (Szczytko and Stewart 1984-A, N); Cascadoperla (Szczytko and Stewart 1979-A, N); Chernokrilus(KondratiefT et al. 2007-A, N); Clioperla(Szczytko and KondratiefT 2015-A); Cosumnoperla (Szczytko and Bottorff 1987-A, N;Bottorff 2001-A, N); Cultu.s(Baumann et al. 1977-A, Stark et al. 1988-A; Myers and KondratiefT. 2009-N); Diploperla (KondratiefT 2004A); Diura(Baumann et al. 1977-A, KondratiefT 2004-A, Nelson and Nelson 2018-A); Frisonia (Jewett 1959-A); Helopicm (KondratiefT 2004-A); Hydroperla (KondratiefT 2004-A); Isogenoide.s(Sandberg and Stewart 2005-A, N); Isoperla(Szczytko and Stewart 1979-A, N;Szczytko and Stewart 2002 A,N,Szczytko and Stewart 2004 A, N,Sandberg 2011-N; Sandberg and KondratiefT 2013-A -A; Szczytko and Kondratiefl" 2015-A); Kogotus(Baumann et al. 1977-A); Malirekus(KondratiefT 2004-A); Megarcys(Van Weiren et al. 2001-A; Stewart and KondratiefT 2012-N); Oconoperla(KondratiefT 2004-A); Oroperla (Siegfried et al. 1977-A, Stark et al. 2017); O.wbenus(Sandberg et al. 2015-A); Perlmodes(Stark and Stewart 1982-A, N); Pictetiella(Baumann et al. 1977; Stark and KondratiefT 2004-A, N); Remenus(KondratiefT 2004-A); Rickera (Szczytko and Stewart 1984-A, N); Salmoperla (Stark and Baumann 2006; Verdone and KondratiefT 2016-A); Setvena (Stewart and Stanger 1985-A, N,Stewart and Stark 2002bA); Skwala(Baumann et al. 1977-A); Siisulus(Bottorff et al. 1989); Yugus(Nelson 2001 -A, N, KondratiefT 2004-A). Taeniopterygidae: Bolotoperla (Stewart 2000-A, Stark et al. 2016-A); Dodd.sia(Baumann et al. 1977-A); Oemopteryx (Baumann et al. 1977-A; Stewart 2000-A, Baumann and KondratiefT 2009-A); Strophopteryx (Stewart 2000-A, Earle and Stewart 2008-N); Taenionema (Stanger and Baumann 1993-A; Stewart 2000-A, Stewart 2009-N); Taeniopteryx (Harper and Hynes 1971c-N, Fullington and Stewart 1980-N, Stewart 2000-A).

o

Family

Lotic—erosional

Generally clingers— sprawlers

Clingers— sprawlers

Clingers— sprawlers

Generally dingers— sprawlers

Habit

**Emphasis on trophic relationships

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic

litter)

depositionai (leaf

Generally lotic— erosional and

Roach Stoneflies

depositionai (logs, leaf litter)

and

Peltoperlidae (24)-

Pteronarcys(8)

depositionai (logs, leaf litter)

and

Lotic—erosional

depositlonal (debris jams. leaf packs)

and

Generally

Habitat

lotic— erosional

Species

Salmonflies

Pteronarcella (2)

Genus

Pteronarcyidae(IO)-

Plecoptera - Stoneflies

Order

species in parentheses)

(number of

Taxa

DeWalt etal. 2018.)

detritivores

Generally shredders—

scrapers

(macroalgae); facultative predators (engulfers); facultative

herbivores

detritivores and

Shredders—

(macroalgae); facultative predators (engulfers) Widespread

West, Southwest

and herbivores

1.7

2.0

0.0

0.0

NW

MA*

(continued)

1887

2374, 2660, 2835, 2966, 2801, 3179, 4599, 5019, 1991,

1433

318, 655, 1272, 2011, 2012, 2374, 2694, 2695, 3747, 3935, 4054, 4262, 4285, 3484, 5006, 5404, 5720, 1347, 1893, 5477, 774, 1972,4122, 2366, 4162,4626, 5559, 4718, 5410,6888, 1489,4717,

1426

87, 655, 2011, 2012, 4587, 4624, 5720, 5743, 3118, 6328,

1249, 1272, 1988, 2213, 2374, 2660, 2827, 2835, 2846, 2966, 4240, 4599, 5006, 5019, 5404, 5720, 1887, 2958

PLECOPTERA

2.2

M

Ecological UM

References** SE

American

Tolerance Values

Distribution

Shredders—detritivores

predators

and facultative

facultative scrapers

herbivores: some

detritivores and

Generally shredders—

Trophic Relationships

of values within a region was 3); table prepared by K. W. Cummins, R. W. Merritt, K. W. Stewart, M. B. Berg, and P. P. Harper. Numbers of species updated from

Tables 6A-6C; tolerance values are taken from Barbour ef a/.(1999) and represent either the mean (when the range

Table 16B Summary of ecological and distributional data for Plecoptera (stoneflies). (For definition of terms see

) )) ) ) ) ))> 3 J )))) ) ) 3 3 3 ) 3 3 3 ) ) 3

Ul

(11)

Taeniopteryginae

(coarse sediments,

Winter Stoneflies

debris jams, leaf packs) and depositional at margins

Generally lotic—erosional

(leaf litter)

Taeniopterygidae

Yoraperla (4)

Viehoperia

collectors—gatherers and scrapers

ciingers

Generally shredders— detritivores; facultative

sprawlers;

Generally

(mountain, intermountain)

facultative—scrapers

detritivores;

West

Shredders—

Appalachians

East

detritivores (leaf litter)

Northwest

Lotic—erosional Ciingers— and sprawlers depositional

West

East

(Nevada, Oregon, California)

Shredders—

detritivores (leaf litter)

Shredders—

Distribution

Lotic—

Ciingers— sprawlers

Habit

erosional

(leaf litter)

Lotic—erosional and depositional

Habitat

North American

Tallaperia (7)

ada

Species

Trophic Relationships

Soliperia (8)

Sierraperla (2)

Peltoperia (2)

Genus

(36)- Willowflies,

Family

Continued

*SE = Southeast, DM = Upper Midwest, M : Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

species in parentheses)

(number of

Taxa

Table 16B

1.4

2.0

2.0

2,0

NW

MA*

Tolerance Values

Ecological

1987,2374,2660, 2835, 2966, 4240, 4599, 5019, 201, 202, 1991, 1887

979, 5654, 5720, 4442

5720

1662,2660, 6277, 6719, 2409, 2410, 4411, 5720, 5777

5720

5720

5720, 1893, 2240, 4411, 1433

2660, 4057, 5192,

References**

) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) > ) ) )) J ) ) ) )

00

o

o

'Jt

2.5

3.0

2.0

UM

M

2.0

NW

2.0

MA*

Ecological 318, 417, 1077, 1087, 1820, 1987, 2021, 2403, 3200, 2409, 5006, 6298, 6469, 5398, 5720, 6585, 3527, 1433

References**

Amphinemurinae(29)

Forestflies

Nemouridae (80) -

occidentalis

Doddsia

Generally erosional (coarse sprawlers; sediments, wood. clingers leaf packs)and depositional (leaf

Generally lotic—

Generally shredders—

litter); lentic— erosional

**Emphasi5 on trophic relationships

collectors—gatherers

detritivores; facultative

Widespread

Appalachians)

Northwest, East (1 in

West,

Taenionema(13)

(continued)

2270

688, 979, 2390, 3667, 1639, 4077,4285, 6770, 3735, 5720, 6880, 168, 2095, 2240, 1991, 2521, 1887

2410, 4624, 5720, 5741, 6328

PLECOPTERA

2.0

2.0

5398, 5720 2403, 2410, 2801, 3591, 5720, 1433

California East

2409, 2410, 4297,

5720

5720

East, Central,

West

East

Strophopteryx(5)

Oemopteryx(5)

rossi

Bobtoperia

5720

New Mexico)

6.3

1.4-

East (1 in Northwest;

collectors— gatherers; Texas, facultative scrapers Colorado,

detritivores; facultative

Shredders—

SE

Distribution

2827, 3666, 5006,

Sprawlers— clingers

Habit

Generally scrapers;

packs)and depositional at margins

(coarse sediments. wood, leaf

Lotic—erosional

Habitat

facultative shredders

Species

North American

Brachypterainae

Taeniopteryx 0 1)

Genus

Trophic Relationships

Tolerance Values

(25)

Family

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic

Order

species in parentheses)

(number of

Taxa

Table 16B

Nemourinae (51)

Family

Continued

detritus)

Lotic—erosional and depositional (in

Habitat

detritivores

detritivores;

(facultative scrapers)

and

depositional

West, East

Northeast Shredders—

Lotic—erosional

detritivores

Prostoia (5)

Northeast

West,

Shredders—

East, Northwest

Podmosta (5)

(macroalgae)

Paranemoura (2)

streams

Shredders—

Lotic—

Northeast

herbivores

West, North,

detritivores and

River Gorge

Columbia

Northwest

West

Widespread

Distribution

American

North

Shredders—

temporary

collectors—gatherers

detritivores; facultative

Shredders—

Trophic Relationships

Ostrocerca (6)

Sprawlersdingers

Sprawlersdingers

Habit

Lotic—erosional (detritus)

wahkeena Spring seeps

Species

Nemoura (4)

Nanonemoura

Lednia (4)

Malenka (13)

Amphinemura(16)

Genus

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

species In parentheses)

(number of

Taxa

Table 16B

6.1

3.4

2.0

3.0

2.0

2.0

2.0

NW

MA*

To erance Values

Ecological

1087, 2390, 2843, 5404, 1704, 2410, 2454, 2414, 5720, 6328, 1433

6529

2408, 3472, 5720,

5720

2392, 2410, 3640, 5720, 1441, 1433

413,418, 2390, 5404, 3525, 6469, 5741, 5720, 6598, 1703, 3527, 1433

353

356, 5720

1470, 3484, 5000, 6328, 4442

1087, 2390, 3640, 3666, 1396, 3667, 5404, 6770, 1704, 2410, 6338, 3029, 3524, 5720, 1, 6529, 1433, 1677, 3010

References**

) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) > )) ) ) ) ) ) )

o

(Jt

'Ji

Northeast

Leuctrinae (59)

Megaleuctrinae

Megaleuctrinae (5)

Megaleuctra (5) Springs, seeps

debris jams, leaf packs) and depositional

(coarse sediments,

Generally sprawlersdingers

Generally lotic erosional

streams

detritivores

Generally shredders—

detritivores (leaf litter) and herbivores (moss)

Shredders—

Appalachians

Northwest,

West, East

West

Northwest

detritivores

temporary

East,

Shredders—

Lotic—spring outflows and

depositional

North,

detritivores,(scrapers)

Distribution

Shredders—

Leuctridae (64) -

Zapada (10)

North American

Lotic—erosional

Habit

Trophic Relationships

and

Habitat

Needleflies

cataractae

rotunda

Species

Sprawlersdingers

Visoka

Soyedina (12)

Shipsa

Genus

Lotic—erosional (detritus)

Family

Continued

*SE = Southeast, DM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

species In parentheses)

(number of

Taxa

Table 16B

0.3

0.0

0.0

2.0

1.0

2.0

MA*

Ecological

(continued)

5720

2240, 2390, 3023, 5209, 1991, 1887

5436, 5720, 5001, 5741, 6328, 4804, 6529, 4442

934, 2442, 3120, 4860, 5438, 4235, 5000,

1433

2410, 3120, 3640, 6770, 5720, 5850, 5854, 5845, 5720,

318, 2390, 5404, 5720

References*

PLECOPTERA

2.0

NW

Tolerance Values

collectors— gatherers

depositional

Stoneflies, Snowflies

Allocapnia (47)

packs)and depositional Clingers

Generally shredders—

detritivores

Shredders—

detritivores

East

Texas

2.8

0.7

SE

3.0

UM

M

1.0

0.0

0.0

0.0

0.0

NW

3.0

1.0

0.0

MA*

Tolerance Values

Ecological

417, 1087, 1820, 1821, 1823, 1824, 2403, 3640, 4836, 4883, 1433, 1677, 3010

1987, 1991, 2403, 4442, 1887, 574, 1677

5548, 5720, 1433

5659

5720, 6643

5720, 6328, 1433

5720

413, 2390, 5404, 6467, 3527, 5631, 1704, 2392, 3029, 6843, 3523, 3526, 5720, 6653, 6882, 6883, 2521, 5720, 1636, 2801, 1321, 6184, 3383, 5195, 1433, 2270

5001, 5720

5424

References**

)) ) > ) ) ) ) )))) ) ))) ) > )) ) ) ) )))

Mid-Atlantic

Generally lotic— erosional (coarse sediments, wood, leaf

Capniidae(169)

detritivores

East, Central,

(some intermittent)

Northwest Shredders—

Small streams

Pomoleuctra (2)

Widespread

Northwest

Zealeuctra (11)

detritivores

Shredders—

detritivores; facultative

Pacific

East, North Central

Shredders—

West

California

Distribution

Lotic—erosional

Generally sprawlers— dingers

Habit

and

Spring outflows

(some intermittent)

Small streams

Habitat

North American

West

augusta

Species

Trophic Relationships

Perlomyia (2)

Paraleuctra (8)

Moselia (2)

Leuctra (31)

Despaxia

Calileuctra (2)

Genus

- Small Winter

Family

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = **Emphasis on trophic relationships

Plecoptera

Order

parentheses)

(number of species in

Taxa

Table 16B

u>

and lentic—

Summer Stoneflles

erosional

Generally lotic

Lotic and lentic

Utacapnia (11)

large rivers

Lotic, small to

Hyporheal

Tahoe)

Lotic (1 pelagic sp. in Lake

alpine lakes

Small streams,

Habitat

Lotic, erosional

Carolina

brevicauda

Species

Sierracapnia (7)

Paracapnia (7)

Nemocapnia

Mesocapnia (16)

Isocapnia (12)

Eucapnopsis

Capnura (7)

Capnia (45)

Bolshecapnia (7)**'

Arsapnia (8)

Genus

Perlldae (91)-

Family

Continued

Clingers

Clingers

Habit

*SE = Southeast, DM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships ***See additional genera in Broome etal. 2019. Illiesia 15: 1-26.

Order

parentheses)

(number of species in

Taxa

Table 16B

North

Predators (engulfers)

Nevada

West, East

California, detritivores

West, East

East

West

West

West

West, East(1)

West, North,

Shredders—

detritivores

Shredders—

detritivores

Shredders—

detritivores

Shredders—

West

Northwest, West

Detritivores

Distribution

American

Shredders—

Trophic Relationships

0.2

SE

M

1.0

1.0

1.0

1.0

NW

1.0

MA*

Ecological

(continued)

674, 1249, 1988,2213, 2374, 2660, 2827, 2835, 2846, 2966, 3179, 4240, 4599, 861, 5019, 6602, 1991, 5720, 6370, 5339, 1887, 2958

5720, 6328

2242, 2403, 2409, 2394, 1396, 5720, 1433

2220, 4305, 5720

4305, 5720

5628, 5629, 5720

6328

1470, 3120, 5720,

2403

252, 1470,3120,3133, 3472, 5438, 5436, 5960, 6328, 697, 3527, 4602, 2409, 3524, 5720, 6643

References**

PLECOPTERA

1.0

UM

Tolerance Values

) ) )))) )

Acroneuriinae (67)

Perlinae (24)

Family

Continued

Attaneuria

Anacroneuria (2 north of Mexico, 32 in Mexico)

Acroneuria (18)

ruralis

erosional

lentic—

Lotic and

Lotic—erosional

Paragnetina (5)

Lotic—erosional

Habitat

Lotic—erosional

sabulosa

Species

Neoperia (15)

Claassenia

Agnetina (3)

Genus

Clingers

Clingers

Clingers

Ciingers

Clingers

Habit

tUnpublished data, K. W. Cummins, Kelloggw

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

species In parentheses)

(number of

Taxa

Table 16B

Ephemeroptera, Plecoptera)

Chironomidae, Trichoptera,

Predators (engulfers;

Predators(engulfers; Diptera, Ephemeroptera, Hydropsychidae)

Predators(engulfers)

Predators(engulfers; Trichoptera, Ephemeroptera, Chironomidae, Simuliidae)

Ephemeroptera, Trichoptera)

Chironomidae,

Predators (engulfers;

Trophic Relationships

Midwest

East, Midwest, Upper

Arizona

Texas,

Widespread

East

Southwest

East,

West, North

East

Distribution

American

North

1.5

1.8

1.6

0,0

0.0

1.0

2.3

2.1

3.1

3.0

NW

0.0

2.0

MA*

Tolerance Values

Ecological

4565, 5720, 1433

5742, 5720

3001, 6298, 6299, 4420, 6658, 6659, 6753, 1610, 2220, 4585, 1704, 3029, 5720, 1433, 3010

4565, 4587, 2564,

1077,2499,3001,

1077, 2389, 2498, 3001, 5404, 2009, 5720, 5888, 1347, 2013, 6753, 1772, 4779, 6658, 6659, 1558, 6, 2890, 1433, t

1705,1704,5742,6158, 3029,5687, 5720,6,1433

83,318, 2011,2012, 5720, 1035, 2454, 5377, 6328, 73, 1057

1077,2389,3001,3280, 5404, 1704, 3029, 5720, 2009, 6585, 2013,4585, 1433, t

References**

) ) ) ) ) > ) ) ) ) ) ) ) ) ) ) > >) ) ) ) ) ) ) )

Ih

crt

h-

cn

Family

Continued

Habitat Habit

Predators (enguifers;

Perlinella (3)

Perlesta(31)

Hesperoperla (2)

Hansonoperia (3)

xanthenes

Lotic—erosional

detritus)

depositional (in

and

Clingers

1.0

1.0

2,0

NW

MA*

Ecological

1640, 2012,2021,

5720

93, 4565, 5720, 1433

3484, 5720

979, 3120, 4240, 5405, 5441, 3484, 5440, 5720, 5410

2801, 5720

References**

gatherers (especially in early instars)

facultative collectors—

Ephemeroptera, Trichoptera);

Chironomidae, Simuliidae,

Predators (enguifers;

East

Widespread

0.0

(continued)

3195, 5720, 74, 1433

5404, 5549, 5720, 1775, 6, 4883, 1433, t

PLECOPTERA

1.0

5.0

5410

4.5

M

Ephemeroptera) 0-4.9 5.0

UM

5720, 88, 89, 4095, 3002, 4096, 5075,

West

4.1

SE

Tolerance Values

Chironomidae, Trichoptera,

Predators(enguifers;

Appalachian

East, Central

Northwest

Eccoptura

Ephemeroptera)

Chironomidae, Trichoptera,

Appalachian

Southern

American Distribution

Trophic Relationships

Northwest

californica Lotic—erosional Clingers

Species

North

> )) ) ) ) ) ) )

Doroneuria (2)

Calineuria

Beloneuna (3)

Genus

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships tUnpublished data, K. W. Cummins, Kelloggw

Order

species in parentheses)

(number of

Taxa

Table 16B

JJ3 >))) 3J3 ) ))) > ) >

Perlodinae (64)

misnomus

Chernokrilus(1)

Predators (engulfers;

Flelopicus(3)

Frisonia

Predators (engulfers)

Predators (engulfers); facultative scrapers

Chironomidae, Simuliidae)

Predators (engulfers)

Clingers

Predators (engulfers)

gatherers)

and collectors—

Generally predators (engulfers);(some facultative scrapers

Diploperla (5)

Lotic—erosional

Generally clingers

Habit

Trophic Relationships

DIura (3)

picticeps

alameda

Baumannella

Cultus(6)

dichroa

Arcynopteryx

Arctic, alpine

Lotic and lentic-

erosional

Generally lotic

Habitat

and lentic—

Species

Perlodidae(153)

Genus

- Spring Stoneflies

Family

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

species in parentheses)

(number of

Taxa

Table 16B

) ) ) ) } ) ) )) )))) ))) V

a\

h-»

(71

North

0.4

2.0

1.6

SE

UM

M

2.0

2.0

2.0

2.0

NW

2.0

MA*

Ecological

4071, 5720

2255, 5720

252, 5208, 5366, 5406, 6079, 1775, 3698, 4212, 634, 6528

225, 5720, 1433, 3010

2011,2410, 4077, 4078, 1347, 5684, 5720, 6328, 1433

5720

5720

3440, 4119, 5406, 5720, 3524, 5741

5366, 1991, 5720, 6370, 6602, 6225, 6492, 1887

2966, 4599, 5019,

References**

> )) ) ) ) ) ) )

East

Northwest

West,

East

West, North,

East

West, East

California, Oregon

California

York, New Hampshire

Superior), New

Michigan (Lake

West, Northwest, Far North,

Distribution

American

Tolerance Values

h-

-4

Family

Continued

innubila barbara

yakimae

Oroperia Osobenus

Species

Oconoperia

Megarcys(5)

Malirekus(2)

Kogotus(2)

Isogenoides(8)

Hydroperla (4)

Genus Predators (engulfers;

Lotic—erosional

Clingers

Predators (engulfers; Ephemeroptera, Trichoptera, Diptera—

Lotic—erosional

Simuliidae, Chironomidae)

Predators (engulfers; Ephemeroptera, Trichoptera, Diptera—

facultative scrapers

Simuliidae, Chironomidae); some

Chironomidae)

and

depositional

Ephemeroptera)

Simuliidae, Chironomidae,

Predators,(engulfers; Diptera, especially Clingers

Clingers

Habit

Trophic Relationships

Lotic erosional

and erosional

depositional

Lotic—

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

species In parentheses)

(number of

Taxa

Table 16B

North

Ecological

Northwest

Nevada

PLECOPTERA

(continued)

4860, 5720

5720

83, 933, 4587, 5720, 6222, 1118, 4095,4566, 4569,4570,4571,4572, 4573, 1704, 4574,4575, 4579,4415,4590, 4582, 4584, 4096,4589,4585, 6328, 4577, 3948

5720, 1433

4590, 4583, 4584, 4586, 1705,4585, 6223, 6225, 5727, 6774

4570, 4572, 1704, 4574, 4575, 4579,

83, 5412, 5720, 5991, 6222, 4566, 4569,

318, 5404, 5720, 5266, 5267, 4883

4138, 4414, 5720

References**

California,

2.0

2.0

2.0

2.0

MA*

5663,5720

1.4

0.0

NW

East

West

East

West

West

East, North,

East

Distribution

American

Tolerance Values

) ) ))))) ) J )} ) ) ) )) ^ > ) > ) ) ) ) ) )

C/l

Isoperlinae (89)

Family

Continued

Cllngers

luctuosa

trictura

Calliperia

Cascadoperia

Yugus(4)

Lotic—erosional

Clingers; sprawlers

Chironomidae);

streams

Predators (engulfers)

gatherers

facultative collectors

Predators (engulfers. especially

Lotic— temporary

Predators (engulfers)

West

West

West

(Appalachian)

East

(California)

Predators (engulfers)

Susulus venustus

West

West

California

West

Predators (engulfers)

Predators (engulfers)

Southeast

East,

Northwest

West,

Northwest

west,

Distribution

Skwala (2)

Lotic—erosional Generally and cllngers; depositional sprawlers

Lotic—eroslonal

Habit

North American

Predators (engulfers) (facultative scrapers?)

sylvanica

Habitat

Trophic Relationships

Setvena (3)

sorpta

Rickera

aurea

Species

Salmoperla

Remenus(4)

Pictetiella (2)

Perlinodes

Genus

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

species in parentheses)

(number of

Taxa

Table 16B

0.0

2.0

2.0

2.0

2.0

2.0

2.0

NW

MA*

Tolerance Values

Ecological

5720

5720

5720, 6719

3523, 5720

5720, 3002

2454, 5412, 5435,

5720, 5991

2409, 5720

5421, 5720, 5991

4860, 5412

References**

) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) >> )) ) ) ) ) ) )

oe

LO

Paraperlinae (7)

(107)-Sallflies

Chloroperlidae

Family

Continued

Utaperia (2)

Paraperia (2)

Kathroperla (3)

Isoperia (84)

Cosumnoperia (2)

Clioperia

Genus clio

Species

Hyporheal

Generally lotic— erosional

depositionai, large cold lake

and

Lotic—erosional

intermittent

Lotic—

Lotic—erosional

Habitat

Generally clingers

Clingers; sprawlers

Clinger

Habit

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

species In parentheses)

(number of

Taxa

Table 16B

Collectors—gatherers: facultative scrapers

Generally predators (engulfers); facultative scrapers and collectors—gatherers

Ephemeroptera, Plecoptera): facultative collectors—gatherers

Chironomidae, SImullidae,

Predators (engulfers;

Predator

Trophic Relationships

North

East, West

West

West

Widespread

West

East

Distribution

American

1.0

1.0

0.5

1.0

2.0

NW

2.0

MA*

Ecological

(continued)

5720

5628, 5720, 6328, 5619

5720

1077, 1656, 1739, 1988, 2011, 2389, 4345, 5720, 1991, 2012, 6370, 1638, 1887, 3235

318, 1739, 1988,2011, 2223, 3640, 4077, 4078, 6753, 1640, 1703, 5366, 5404, 3527, 3698, 6585, 1347, 2410, 2454, 3030, 4433, 5741, 5720, 1775, 3286, 3524, 1704, 3700, 4192, 2389, 3702, 2408, 6658, 5265, 5268, 6528, 4442, 6643, 1433, 3010

5868

1775, 2389, 4077, 5720, 1433, 3010

References*

PLECOPTERA

0-5.6 2.0

4.8

M

Tolerance Values

j ; i I } ) ) J )))) J ) ) ) > ) > J 3 3 )))3

Ul

O

»

(100)

Chloroperlinae

Family

Continued

hoopa

Triznaka (3)

Sweltsa (36)

Suwallia (12)

terna

Rasvena

forcipata

Sasquaperla

Plumiperia (2)

Neaviperia

Haploperia (6)

streams

i

West(Alaska)

Distribution

West

East, West

East, West

Northwest

East

Northwest

West,

Northwest

East, West

California, Oregon

0.0

0.0

1.3

1.4

1.0

1.0

1.0

1.0

1.0

1.0

NW

MA*

Tolerance va ues

Ecological

6328

2012, 2454, 5720,

5720, 4602, 6328, 1433, 3010

2389, 3640, 2801,

1296, 1470, 2012,

5728

2012, 5720, 6328,

5670

5720

5720, 6643

318, 2394, 3029, 5404, 5720, 1433, 3010

5720

5720, 4883, 1433

5740

References*

i ) ) ) )) ) ) ) ) ) )) ) > )) ) ) ) I ))

Chironomidae, Simuliidae)

Predators (engulfers;

collectors— gatherers (scavengers)

Predators,(engulfers; Chironomidae, Simuliidae); facultative

Lotic—

temporary

Chironomidae, Simuliidae)

Predators (engulfers;

gatherers and scrapers

facultative collectors—

Chironomidae);

Predators (engulfers;

Predator (engulfers)

collectors—gatherers

and facultative

Generally facultative predators (engulfers)

Relationships

Mexico (Baja),

Habit

Bisancora (2)

Lotic—erosional

Habitat

North American

East, West

ovibovis

Species

Trophic

Alloperla (34)

Alaskaperia

Genus

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

parentheses)

species in

(number of

Taxa

Table 16B

AQUATIC AND SEMIAQUATIC HEMIPTERA Dan A. Polhemus

Bishop Museum, Hawaii

HETEROPTERA INTRODUCTION

Sixteen of the 19 major families of Heteroptera associated with the aquatic habitats are represented in the North American insect fauna, with only the pantropical Helotrephidae,the Old World Aphelocheiridae, and the Neotropical Potamocoridae absent. Seven families are totally aquatic, leaving the water only to mate or migrate. Three other families of the water striders live on the surface film and adjacent banks, and the remaining six groups live at aquatic margins, although some are often found on the water surface as well. All of these families belong to three subordersin theorder Heteroptera:Leptopodomorpha, Gerromorpha, and Nepomorpha. It should be noted that although Heteroptera were considered to be an order by Henry and Froeschner(1988)in their catalog of North American fauna, other authors have treated them as a suborder within Hemiptera. In

addition

to

the

above, several

other

heteropteran families occur in North America whose members are essentially terrestrial, but have species that are occasionally found near water. They are the Ceratocombidae, Dipsocoridae, and Schizopteridae (in the infraorder Dipsocoromorpha), and the intro duced Leptopodidae (related to Saldidae). These families are not treated further in this work except to list ecological and taxonomic references for the Dipsocoromorpha that provide an entry to the liter ature.

Over 4,700 species of aquatic and semiaquatic Heteroptera are now considered to occur worldwide

(Polhemus and Polhemus 2008). This current state of knowledge contrasts with the 2,900 species previously estimated by Jaczewski and Kostrowicki in 1969, a total which excluded the Saldidae. Presently 68 genera and 424 species are recognized from North America. The fauna, at this point, is well-documented and no new native species have been described for many

years(Polhemus and Polhemus 2007),although intro duced, non-native species are sporadically added. Dispersal of most aquatic Hemiptera occurs by flight, but there is also evidence for hurricane transport (Herring 1958). Our Nearctic fauna has been enriched by the northward dispersal of Neotropical forms, whereas transverse mountain ranges have blocked such northward invasions into the Palaearctic region, which has a comparatively poor fauna by compari son. Holarctic species are known only in the coldadapted Corixidae, Gerridae, and Saldidae, the latter family having 14 such species. A discussion of the world distribution of water bugs was given by Pol hemus and Polhemus(2008), building on the previous work of Hungerford (1958) and of an analysis for Gerromorpha by Andersen (1982). The local faunal composition and zoogeography of North American water bugs has been discussed by Menke (1979) for California, Slater (1974) for Connecticut, and Epler (2006)for Florida. The aquatic and semiaquatic Heteroptera are remarkable for their diversity of body forms, reflect ing adaptions to a wide variety of niches. They occupy many varied habitats, including small streams, large rivers, lakes,swamps,saline marshes, hot springs,and high mountain beaver ponds, where they occur on the water surface, underwater, and along the damp margins. A habitat key and other significant data on biology were provided by Hungerford (1920) in his classic work on water bug ecology; this was subse quently updated by Usinger (1956b) and further by Menke et al. (1979), these latter two works having particular reference to California and adjacent states. The characterizations of habitat provided for each group in Table 15A should be considered only rough generalizations due to the many exceptions that occur. Water bugs are for the most part predators, and in addition many species seem to be relatively resistant to vertebrate predation, which is generally attributed to their possession ofthoracic scent glands, a defining

521

522

Chapter 17 Aquatic and Semiaquatic Hemiptera

character for the order. Only the Corixidae differ sig nificantly in regard to diet, with many genera being primarily collectors, feeding on detritus and diatoms, and being heavily preyed upon by fish and birds (Table 15A). Some aquatic Heteroptera are of recognized economic importance, and the role of additional forms is presently being investigated. Certain genera, including Notonecta, Belostoma, and Lethocems are attracted to lights and may be a nuisance in swimming pools,because they can inflict painful bites. Let/jocerw^, a very large belostomatid, can also be a nuisance at fish hatcheries (Wilson 1958) where they prey on fry. Corixidae have long been a relished food item in Mexico under the name "ahuautle" and are also used

extensively as food for pet fish and turtles, and Belostomatidae are considered a delicacy in Asia, where they are fried and served on wooden skewers. Corixidae have been shown to be good indicators of lentic water quality (Jansson 1977). Both surface and aquatic bugs can be important predators of mosquito larvae and adults(Jenkins 1964; Collins and Washino 1985; Table 15A), with Notonecta undulata Say pre ferring mosquito larvae over other foods (Ellis and Borden 1970; Toth and Chew 1972b). These preced ing authors and Laird(1956)have urged further study of aquatic Hemiptera as biological control agents of such pests. Although there are exceptions, most aquatic Heteroptera lay their eggs in the spring, develop during the warmer months,over-winter as adults, and repeat the cycle. Some saldids overwinter as eggs, many gerrids are bivoltine, and in southern regions a number of species breed throughout the year. Eggs are of various forms and are laid in a wide variety of places, either glued to a substrate or inserted in the earth or plants. They have a wide variety of shapes: spindle-shaped, oval, or occasionally stalked. The chorion is tough and often hexagonally reticulate, and successful hatching usually takes place submerged or in damp habitats(see Chapter 5,Table 5A). A splendid review of heteropteran eggs and embryology is given by Cobben (1968). All but a few species have five nymphal instars; the exceptions have four and include some Mesovelia sp., Microvelia sp., Macrovelia sp., and Nepa species. Modes oflocomotion are variable throughout the aquatic Heteroptera. Some of the subaquatic bugs (Pleidae, Notonectidae, Naucoridae, Corixidae)swim with synchronous oar-like strokes of the hind legs, whereas others(Nepidae, Belostomatidae)move using synchronous strokes of both the middle and hind pairs of legs, those on each side working alternately. When swimming vigorously the latter families stroke both

pairs in unison. The belostomatids are the strongest, the nepids the weakest, and the corixids the most agile of the underwater swimmers.

The water striders (Gerromorpha) live on the water surface, where they are supported by surface tension of the water and an unwettable hydrofuge pile present on their tarsi and sometimes their tibiae. These bugs are able to easily glide over the water because of the low resistance, and can use their wettable claws or other pretarsal structures to pene

trate the surface for "traction," although in the more specialized families such as Gerridae and Veliidae these can also be retracted when necessary. Forward thrust during rowing is created by the surface film "packing up" behind the tarsi during the backward power stroke. These animals usually steer by using their hind legs as parallel rudders, combined with unequal strokes of the middle legs (Menke 1979; Andersen 1982). Exploiting these surface-dwelling capabilities has allowed species of the gerrid genus Halobates to colonize the open pelagic ocean, the only insects to do so(Cheng 1985; Cheng and Frank 1993). Other surface-dwelling Gerromorpha "walk" over the surface film with tripodal locomotion, each leg alternating movement with its opposite. Some veliids are also able to move very rapidly when alarmed by using "expansion skating," discharging saliva from their beak that causes a localized lower

ing of the surface tension, and then being carried forward on the contracting surface film (see review in Andersen 1982). Most of the subaquatic bugs (Nepomorpha) breathe by means of an air store carried dorsally

between the wings and abdomen,combined with an exposed thin bubble (or physical gill) on the ventral surface held in place by a fine pile of hydrofuge hairs (Chapter 4). The methods of air store replen ishment are often distinctive for each family, e.g.,

through tubes (Nepidae), air straps (Belostomati dae), the pronotum (Corixidae), or the tip of the abdomen (Notonectidae and most Naucoridae). A few naucorid bugs can remain submerged indefi nitely utilizing plastron respiration (Chapter 4; Menke 1979).

Several types of communication have been documented in aquatic Heteroptera. Sound produc tion in the Gerromorpha has long been reported for the Veliidae (Leston and Pringle 1963); however, a few Gerridae also possess stridulatory mechanisms. In addition, Saldidae and most Leptopodidae (Leptopodomorpha)also possess evident stridulatory mechanisms (Polhemus 1985; Pericart and Polhemus 1990), as do some, if not most, of the families of Nepomorpha. Stridulatory mechanisms have been

Chapter 17 Aquatic and Semiaquatic Hemiptera

described in the Old World Helotrephidae, the New World Naucoridae and Nepidae, and the cosmopolitan Gelastocoridae, Notonectidae, and Corixidae (Polhemus 1994b). Corixid "songs" have been studied in detail (Jansson 1976), with multiple species often inhabiting the same pond and producing acoustically distinctive, species-specific signals. Male water striders (Gerridae) also signal on the surface using ripple communication to identify potential mates and ward off other competing males (Wilcox and Spence 1986).

External Morphology of Nymphs and Adults

The nymphs of aquatic and semiaquatic Heteroptera have one-segmented tarsi, a useful char acteristic in separating them from adults, especially the apterous gerromorphans(water striders),in which adults always have at least two tarsal segments on one or more pairs oflegs. Nymphs of Hemiptera are hemimetabolous and resemble the adults, but the body parts have different proportions and the developing wings are present as wing pads (Fig. 17.4) in the ulti mate and penultimate instars. Keys to the nymphs of North American families and subfamilies of Heterop tera, based in a large part on the trichobothria (specialized sensory hairs) and dorsal abdominal scent gland openings (Fig. 17.4) have been published (DeCoursey 1971; Herring and Ashlock 1971; Yonke 1991). A key and synopsis of the families and genera of adult North American Heteroptera has been published by Slater and Baranowski (1978), but dif fers somewhat in arrangement from this work; e.g., the genera Aquarius and Limnoporus are included under Gerris (Gerridae); the family Macroveliidae is included under the Mesoveliidae; and the genera Platyvelia and Steinovelia are included under Paravelia (Veliidae). The interpretations followed in the present chapter are those now generally accepted. With the exception of a few families in which the head and thorax are fused or closely conjoined (e.g., Pleidae, Naucoridae), the head,thorax, and abdomen

are generally well-defined in aquatic and semiaquatic Hemiptera. Head: The eyes are usually prominent and well-developed. Ocelli may be present but are lacking in many aquatic families, or present only in winged

forms of some semiaquatic species (e.g., Mesovelia). Antennae are three-, four-, or five-segmented and are ordinarily long and quite conspicuous in the semiaquatic bugs, but short and hidden in the subaquatic species. Two semiaquatic families (Ochteridae, Gelastocoridae), which inhabit the margins of

523

fresh waters and are closely allied to the other subaquatic Nepomorpha, have short antennae that are largely or entirely hidden. The rostrum, or beak, varies in length, and has either three or four visible segments (except for Corixidae; Fig. 17.17). It attaches to the apex of the head and is directed pos teriorly underneath (Cobben 1978). The ventral region between the base of the beak and the collar, called the gula (Fig. 17.100), is one of the primary characters used in separating the HemipteraHeteroptera from the Homoptera (leaf hoppers; see Homoptera section). Thorax:The three-segmented thorax bears the legs and wings, and due to fusions or extra sutures, the segments are often difficult to discriminate except in the wingless forms (Fig. 17.145). The metasternum usually bears one or more scent glands(Fig. 17.81)and sometimes lateral scent channels(Fig. 17.145). The legs are variable in form and reflect striking adaptations of aquatic Heteroptera to their environment(Figs. 17.80, 17.143, and 17.146). Leg segments are variable in length; however, each leg always consists of a coxa articulating with the body followed by a trochanter joining the coxa and femur. The femur and tibia are usually the longest segments, with the tarsi being one-, two-, or three-jointed and bearing the claws. Alary (wing) polymorphism in water bugs is a common phenomenon, although a few genera, such as the marine water striders in Halobates and the

freshwater macroveliid Oravelia, are known only in the wingless state. The forewings of Heteroptera, or hemelytra, have a leathery basal section divided into a clavus and corium, and a thin, membranous

posterior portion (Fig. 17.2), which may appear to be lacking in some aquatic species (e.g., Pleidae). The hind wings are covered by the forewings and are uniformly thin, membranous and translucent; in some cases the forewings may be present but the hind wings may be absent. Winged forms are more common in southern regions, and wingless or short-winged forms are more common among the surface-dwelling Gerromorpha, although the subaquatic Naucoridae may also exhibit significant wing polymorphism in genera such as Cryphocricos. The most extensive studies of alary polymorphism and its mechanisms have involved the Gerridae (Brinkhurst 1959, 1960; Vepsalainen 1971a,b; Vepsalainen 1974; Andersen 1973). It appears that in this family temperature, photoperiod, population density, resource availability and habitat stability may all play a role in determining whether winged or wingless forms are produced. Abdomen: The abdomen bears the spiracles and genitalia. The first visible segment ventrally is actually

524

Chapter 17 Aquatic and Semiaquatic Hemiptera

the second, and the first seven segments are usually similar in form. The eighth through tenth segments form the genitalia and may or may not be distinguish able. In some families of Nepomorpha (e.g., Ochteridae, Gelastocoridae, and Corixidae) the last few abdominal segments are asymmetrical in males but symmetrical or nearly so in females. The male genita lia usually consist of a cup-like pygophore ventrally, which encloses a complex aedeagus, overlain by a lid like proctiger dorsally, and flanked by a pair of parameres laterally. The shapes and sclerotization of these structures are often very diagnostic at the indi vidual species level, and dissection of them may be required for definitive identifications to species level in certain genera. HOMOPTERA INTRODUCTION

The suborder Homoptera is currently being divided, however the classification is not yet settled (see Schaefer 1996), so the suborder "Homoptera" is retained here for practical reasons. The Homoptera have not adapted to truly aquatic life as have some Heteroptera. However, some species of Homoptera can be considered marginally semiaquatic as they have a more or less permanent association with the margins of both the intertidal zone and fresh water, where they feed on aquatic plants. Homopteran species commonly found in semiaquatic habitats belong to a half dozen or more families. Draeculacephala spp. (Cicadellidae) are found along stream margins,and the transcontinental Helochara communis Fitch (Cicadellidae) is appar ently always associated with low marshy grasses. Adults and nymphs of Megamelus davisi Van Duzee (Delphacidae) feed on all emergent parts of water lilies {Nuphar sp.)(Wilson and McPherson 1981). Although few, if any, of the homopterans are subjected to significant aquatic inundation, the eggs of Prokelisia marginata (Van Duzee)(Delphacidae) are deposited in Spartina sp., an intertidal grass, which is sometimes completely submerged. Many intertidal Homoptera (Delphacidae, Cicadellidae,

Issidae) inhabit salt marsh vegetation (Denno 1976) that is occasionally inundated, but the adults of these species move up the culms of vegetation to escape rising tides (Davis and Gray 1966). Conversely, Cameron (1976) has suggested that certain Homoptera can withstand long periods of submergence in California marshes, which typically have a steeper littoral gradient than those studied by Davis and Gray (1966) in North Carolina. Certain of the Auchenhorrhyncha may locate in an air bubble trapped in leaf or blade axils during inundation; the bubble could function as a physical gill. EXTERNAL MORPHOLOGY

Although the inclusion of Homoptera in a work on aquatic insects is problematic, collectors of semi aquatic insects will potentially encounter them and have the need for identifications. Homoptera differ morphologically from Heteroptera primarily in that their posteriorly-directed beak appears to arise from the underside of the thorax rather than the head

(Fig. 17.1). The form of the forewing is also different, with the corium varying from uniformly leathery to hyaline or membranous, rather than exhibiting the consistently thickened basal section seen in the heteropteran wing as described above (see Figs. 17.2, 17.100, and 17.116). Within the Sternorrhyncha, Haliaspis spartinae (Comstock)(Diaspididae), Eriococcus sp. (Eriococcidae), and various pseudococcids inhabit intertidal vegetation and are at least occasionally inundated by tides. In Britain, the aphid Pemphigus trehernei Foster occurs on the roots of an intertidal aster

(Foster 1975). For the most part, the Homoptera have not evolved the sophisticated respiratory mechanisms

found in aquatic Heteroptera. They can be considered no more than semiaquatic,although some scale insects and mealybugs approach an aquatic existence. The respiratory adaptations of salt marsh homopterans have been reviewed by Foster and Treherne(1976; see also Chapter 4).

Chapter 17 Aquatic and Semiaquatic Hemiptera

525

KEY TO THE FAMILIES OF AQUATIC AND SEMIAQUATIC HEMIPTERA

1.

Head without a gula; gular region hidden by posteriorly directed head

1'.

Head with a gula; gular region not hidden

(Fig. 17.1)

2(1').

2'.

3(2).

3'.

(Fig. 17.100)(Suborder Heteroptera) Antennae shorter than head,inserted beneath eyes, not plainly visible from above (Fig. 17.100) except in Ochteridae (Fig. 17.5); aquatic or semiaquatic bugs (at margins of standing- or running-water habitats Antennae longer than head,inserted forward of eyes, plainly visible from above (Figs. 17.2 and 17.3); bugs on surface or at aquatic margins Beak triangular, very short, unsegmented (sometimes transversely striated), appearing as apex of head (Fig. 17.15); front tarsus with a single segment, scoop-like, fringed with stiff setae forming a rake (Fig. 17.22) Beak cylindrical, short to long, 3- or 4-segmented (Fig. 17.100); front tarsus not scoop-like or fringed with stiff setae

4.

Scutellum exposed, covered by pronotum only at anterior angles

4'. 5(3'). 5'.

(Fig. 17.21) Scutellum concealed (Fig. 17.28) Apex of abdomen with respiratory appendages(Figs. 17.12-17.111) Apex of abdomen without respiratory appendages

6(5).

Apex of abdomen with a pair of flat, retractile air straps

(Fig. 17.12) 6'.

7(5').

(Suborder Homoptera) 2

3 11

4

5

MICRONECTIDAE(p. 538) CORIXIDAE(p. 527) 6 7 BELOSTOMATrDAE(p. 527)

Apex of abdomen with cylindrical breathing tube (siphon)composed of 2 slender, non-retractile filaments (Fig. 17.111) NEPIDAE(p. 538) Middle and hind legs without fringe-like swimming hairs; ocelli present(Fig. 17.64)

except in Nerthra rugosa Desjardins; semiaquatic bugs at margins of aquatic habitats

8

7'.

Middle and hind legs with fringe-like swimming hairs (Fig. 17.115); ocelli absent;

8(7).

aquatic bugs 9 Front legs raptorial (grasping), femora broad (Figs. 17.60 and 17.61); rostrum short, not reaching

8'.

hind coxae; antennae not visible from above (Fig. 17.64) GELASTOCORIDAE(p. 532) Front legs not raptorial, femora not broad (Fig. 17.6); rostrum long, reaching, or extending beyond hind coxae; tips of antennae usually visible from above

9(7').

(Figs. 17.5 and 17.69) OCHTERIDAE—DcAferMS Front legs raptorial, femora broad; weakly convex dorsally (Fig. 17.100). . NAUCORIDAE(p. 538)

9'.

Front legs slender, femora not broad; body strongly convex dorsally

(Figs. 17.95-17.116) 10(9').

10

Body form ovoid; 3 mm or less in length (Fig. 17.95); all legs similar; hind tarsus with 2 well-developed claws(Fig. 17.97) PLEIDAE(p. 540) 10'. Body form elongate, 5 mm or more in length (Fig. 17.116); hind legs long, oar-like; claws of hind tarsus inconspicuous(Fig. 17.115) NOTONECTIDAE(p. 538) 11(2'). Membrane of wing with 4 or 5 distinct similar cells (Fig. 17.2); hind coxae large, transverse, with broad coxal cavity (Fig. 17.3) SALDIDAE(p. 540) 11'. Membrane of wing without distinct similar cells (Fig. 17.139); hind coxae small, cylindrical, or conical; coxal cavity socket-like (Fig. 17.145) 12 12(11'). Claws of at least front tarsus inserted before apex (Fig. 17.144) 13 12'. Claws of all legs inserted at tips of tarsi (Fig. 17.92) 14

526

Chapter 17 Aquatic and Semiaquatic Hemiptera

ocelli

collar

-antenna

callus rostrum

pronotum

pronotum

Figure 17.1

posterior lobe pronotal furrow claval suture

Figure 17.5

tarsus scutellum

wing pads

clavus

scent gland

veins of the cerium

ostiole corium

embolar fracture

subiateral cell

membrane

Figure 17.6

Figure 17.4 Figure 17.2 antenna rostrum

coxae

hypocostal ridge-

trochanter metasternum

t— femur

female subgenltal plate

hemelytra tibia

Figure 17.3

Figure 17.7

Figure 17.8

Figure 17.1 Lateral view of adult Cicadellidae (Homoptera).

Ochteridae.

Figure 17.2 Dorsal view of adult Saldula sp.

Figure 17.6 Leg of Ochteridae.

Figure 17.5 Dorsal view of head and pronotum of

(Saldldae).

Figure 17.7 Lethocerus americanus (Leidy), dorsal

Figure 17.3 Ventral view of adult Saldula sp.

view (Belostomatidae; from Usinger 1956). Figure 17.8 Belostoma baker! Montandon, dorsal view (Belostomatidae; from Usinger 1956).

(Saldldae).

Figure 17.4 Dorsal view of Saldldae nymph.

Chapter 17 Aquatic and Semiaquatic Hemiptera

527

13(12).

Hind femur short, distally either scarcely or not surpassing apex of abdomen; metasternum with a pair of lateral scent grooves terminating on pleura in front of hind coxae (Fig. 17.145); dorsum of head usually with median longitudinal sulcus or glabrous(smooth)stripe; mid legs inserted about midway between front and hind legs, except Trochopus and Rhagovelia, which have feather-like structures on the middle tarsus (Fig. 17.146), and Husseyella, which has blade-like structures instead of claws on middle tarsus (Fig. 17.143) VELIIDAE (p. 544)

13'.

Hind femur long, distally greatly exceeding apex of abdomen (scarcely in Rheumatobates females); metasternal region with single median scent gland opening (omphalium; Fig. 17.81), lateral scent grooves absent; dorsum of head without median groove or line except in Rheumatobates-, mid legs inserted closer to hind legs than forelegs GERRIDAE(p. 534)

14(12').

Body long, slender; head as long as or longer than combined length of pronotum and scutellum (Fig. 17.94) HYDROMETRIDAE—//prfro/netra

14'.

Body stout; head length not greater than combined length of pronotum and scutellum, or pronotum alone in wingless forms(Fig. 17.90)

15

15(14'). Tarsi two-segmented; head ventrally with deep longitudinal channel for reception of rostrum (Fig. 17.86) HEBRIDAE(p. 535)

15'.

Tarsi 3-segmented; head ventrally without longitudinal channel(Fig. 17.91)

16

16(15'). Inner margins of eyes converging anteriorly; femora with at least 1 or 2 black spines on dorsum distally; winged forms with exposed

bilobed scutellum (Fig. 17.90) 16'.

MESOVELIIDAE—Mesovelia

Inner margins of eyes arcuate, not converging anteriorly (Fig. 17.88); femora without black spines; winged forms with scutellum concealed by pronotum

(Fig. 17.88)

MACROVELIIDAE(p. 535)

KEYS TO THE GENERA OF AQUATIC AND SEMIAQUATIC HEMIPTERA Belostomatidae

1.

Tibia and tarsus of hind leg strongly compressed, thin, much broader than middle tibia and tarsus (Figs. 17.7 and 17.11); basal segment of beak about half length of 2nd; length 40 mm or more (LETHOCERINAE)2

r.

Tibia and tarsus of middle and hind leg similar (Figs. 17.8-17.10); basal segment of beak subequal to 2nd; length 37 mm or less

2(1). 2'. 3(1'). 3'.

Inner pad of setae offorefemur with two furrows Inner pad of setae of forefemur without any trace of a furrow Membrane of hemelytron reduced (Figs. 17.9 and 17.12) Membrane of hemelytron not reduced (Figs. 17.8 and 17.13); length 26 mm or less

3 Lethocems Mayr Benacus Stal Abedus StM

Belostoma Latreille

Corixidae

1. r.

Rostrum (beak) without transverse striations; nodal furrow absent (Fig. 17.16) (CYMATIINAE) CymatiaFlor Rostrum with transverse striations (Figs. 17.15 and 17.17); nodal furrow present (Fig. 17.19) (CORIXINAE) 2

2(1').

Foretarsus with rather thick, well-developed apical claw; pala of both sexes narrowly digitiform (finger-like)(Figs. 17.24 and 17.28) (GRAPTOCORIXINI) 3

2'.

Foretarsus with spine-like apical claw usually resembling spines along lower margin of palm; pala not digitiform (Figs. 17.22 and 17.35)

4

528

Chapter 17 Aquatic and Semiaquatic Hemiptera

postocular space middle tibia

Figure 17.14

hind tibia

Figure 17.9

Figure 17.10

Figure 17.11

Figure 17.15

membrane

air strap

Figure 17.13

Figure 17.12

Figure 17.16

Figure 17.9 Abedus indentatus (Haldeman), dorsal view (Belostomatidae; from Usinger 1956). Figure 17.10 Middle and hind legs of Belostoma sp. (Belostomatidae). Figure 17.11 Middle and hind legs of Lethocerus sp. (Belostomatidae). Figure 17.12 Hemelytra of yAdedus sp. (Belostomatidae).

Figure 17.13 Hemelytra of Belostoma sp. (Belostomatidae). Figure 17.14 Side view of head of Glaenocorisa sp. (Corixidae). Figure 17.15 Side view of head of Hesperocorixa sp. (Corixidae). Figure 17.16 Left hemelytron of Cymatia sp. (Corixidae; from Brooks and Kelton 1967).

Chapter 17 Aquatic and Semiaquatic Hemiptera

529

pronotal disk

.prothoracic lobe lablum

clavopruina

conum

embolium

-corioprulna

clavus metasternum

postnodal

pruina

metaxyphus

nodal furrow

Figure 17.19

membrane

\

interocular space

abdominal strlgil

Figure 17.17

Figure 17.18

peg row

width of eye tarsus (pala)

upper palar row of setae

apical claw

Figure 17.20

Figure 17.25

lower palar row of setae

femur

Figure 17.22

Figure 17.26

Figure 17.27

Figure 17.23 Figure 17.24

Figure 17.17 Ventral view of Corisella sp.(Corixidae; from Menke etal. 1979). Figure 17.18 Dorsal view of Corisella sp. (Corixidae; from Menke et al. 1979). Figure 17.19 Enlarged portion of hemelytron of Corisella sp.(Corixidae) showing pruinose areas (from Menke etal. 1979). Figure 17.20 Dorsal view of head of Palmacorlxa sp. (Corixidae). Figure 17.21 Dorsal view of Tenagobia sp. (Micronectldae).

Figure 17.21

Figure 17.22 Male foreleg of Cenocorlxa sp. (Corixidae; from Menke et al. 1979). Figure 17.23 Abdominal tergites of Neocorlxa sp. (Corixidae). Figure 17.24 Male foreleg of Graptocorixa sp. (Corixidae; from Menke etal. 1979). Figure 17.25 Metaxyphus of Dasycorixa sp.(Corixidae). Figure 17.26 Metaxyphus of Glaenocorisa sp. (Corixidae). Figure 17.27 Abdominal tergites of Graptocorixa sp. (Corixidae; from Menke et al. 1979).

530

Chapter 17 Aquatic and Semiaquatic Hemiptera

3(2).

Tergal lobes on left side of male abdomen produced posteriorly (i.e., abdomen sinistral) (Fig. 17.23); strigil absent; female abdomen slightly asymmetrical; female face slightly concave, densely pilose (hairy) Neocorixa Hungerford

3'.

Tergal lobes on right side of male abdomen produced posteriorly (i.e., abdomen dextral) (Fig. 17.27); strigil present on right side; female abdomen asymmetrical; female face not concave, not densely pilose Gmptocorixa Hungerford

4(2').

Eyes protuberant(produced above surface) with inner anterior angles broadly rounded; face depressed in both sexes, with dense hair covering; postocular space broad, head transversely depressed behind eyes (Fig. 17.14) (GLAENOCORISINI) 5 Eyes not protuberant, inner anterior angles acute (Fig. 17.15); face of females usually not densely hairy or depressed; if postocular space is broad, then head is not transversely depressed behind eyes (CORIXINI) 6

4'.

5(4). 5'.

6(4').

6'.

7(6').

7'.

8(7'). 8'. 9(8).

Pronotum and clavus strongly rastrate (with longitudinal scratches); metaxyphus(Fig. 17.17) broadly triangular (Fig. 17.26) Glaenocorisa Thomson Pronotum and clavus not strongly rastrate; metaxyphus(Fig. 17.17) narrowly triangular (Fig. 17.25) Dasycorixa Hungerford Apices of hemelytral clavi not or scarcely surpassing a line drawn through the costal margins at the nodal furrows; abdominal asymmetry of males sinistral, strigil on left; foretibia of male produced over base of pala (Figs. 17.29 and 17.34); species smaller than 5.9 mm in length rr/cAoconxa Kirkaldy Apices of hemelytral clavi clearly surpassing a line drawn through the costal margins at the nodal furrows(Figs. 17.19 and 17.33); abdominal asymmetry of males dextral, strigil on right (Fig. 17.27); foretibia of male not produced over base of pala (greatly expanded distally in Centrocorisa sp. but not over base of pala; Fig. 17.37); species usually longer than 5.9 mm 7 Pruinose (frosted) area at base of claval suture (clavopruina) short and broadly rounded, usually about one-half to two-thirds as long as postnodal pruinose area (postnodal pruina) (Figs. 17.30 and 17.32); clavus rastrate Hesperocorixa Kirkaldy Pruinose area at base of claval suture (clavopruina) either broadly rounded, narrowly rounded or pointed distally, subequal to or longer than postnodal pruinose area (postnodal pruina)(Fig. 17.33)(except two-thirds in Corisella, which has smooth clavus) 8 Body short, broad, more than one-third as broad as long (width measured across pronotum) 9 Body elongate, distinctly less than one-third as broad as long 10 Pronotum and clavus only faintly rugulose (wrinkled); male foretarsi expanded distally (Fig. 17.37); male without strigil or pedicel (Fig. 17.38); none of the species have a longitudinal groove on the ventral surface of middle femora

9'.

Centrocorisa Lundblad

Pronotum and clavus distinctly rastrate; male foretarsi not expanded distally in United States species (Fig. 17.35); male with strigil or at least pedicel; some species with a longitudinal groove on the ventral surface of middle femora of both sexes

(Fig. 17.31) Morphocorixa iaczewski 10(8'). Palar claw of both sexes minutely serrate (saw-like) at base (Fig. 17.46); upper surface of male pala deeply incised (Fig. 17.45); vertex of male usually acuminate (tapering to a point)(Fig. 17.42); usually with hemelytral pattern indistinct or obscure Ramphocorixa Abbott 10'. Palar claw not serrate; upper surface of male pala not deeply incised; vertex of male not acuminate (Fig. 17.43); hemelytral pattern usually distinct 11 11(10'). Posterior margin of head sharply curved, embracing very short pronotum (Fig. 17.43); interocular space much narrower than width of an eye (Fig. 17.20); median lobe of male abdominal tergite VII with a hook-like projection (Fig. 17.44) Palmacorixa Abbott

Chapter 17 Aquatic and Semiaquatic Hemiptera

Figure 17.28

Figure 17.30

Figure 17.29

Figure 17.35

length of clavopruina'

531

Figure 17.34

length of postnodal ■ prulna

Figure 17.37

'f-V-Vi pedicel

W Figure 17.31

Figure 17.32

Figure 17.33

Figure 17.36

Figure 17.38

Figure 17.28 Graptocorixa californica (Hungerford)(Corixidae; from Usinger 1956). Figure 17.29 Trichocorixa reticulata (Guerin-Menevilie)(Corixidae; from Usinger 1956). Figure 17.30 Hesperocorixa vulgaris (Hungerford) (Corixidae; from Usinger 1956). Figure 17.31 Dorsal view of middle femur of Morphocorixa sp. (Corixidae). Figure 17.32 Left hemelytron of Hesperocorixa sp.(Corixidae; from Brooks and Kelton 1967). Figure 17.33 Left hemelytron of Corisella sp. (Corixidae; from Menke etal. 1979).

Figure 17.34 Male foreleg of Trichocorixa sp. (Corixidae; from Brooks and Kelton 1967). Figure 17.35 Male foreleg of Morphocor/xa sp. (Corixidae). Figure 17.36 Abdominal tergltes of Morphocorixa sp. (Corixidae). Figure 17.37 Male foreleg of Centrocorisa sp. (Corixidae). Figure 17.38 Abdominal tergltes of Centrocorisa sp. (Corixidae).

532

Chapter 17 Aquatic and Semiaquatic Hemiptera

1 r.

Posterior margin of head not sharply curved, pronotum longer (Fig. 17.40); interocular space at least subequal to width of an eye; median lobe of male abdominal tergite VII without a hook-like projection

12

12(11'). Pronotum and clavus smooth and shining, at most faintly rugulose (wrinkled) (Fig. 17.39)

12'.

Coraef/fl Lundblad

Part or all of pronotum and clavus rough, either rastrate (with longitudinal scratches) or rugulose, or both (Fig. 17.40)

13

13(12').

Markings on clavus and corium narrow and broken, usually open reticulate (covered with network of fine lines) with much anastomosing (merging)(Fig. 17.55); pronotal carina (ridge) distinct on at least anterior one-third (Fig. 17.41)

14

13'.

Markings on clavus transverse, those of corium transverse, longitudinal or reticulate (Figs. 17.56-17.59); pronotal carina absent or faintly expressed on anterior border at most (Figs. 17.40 and 17.54)

15

14(13).

Median carina of pronotum well-defined on anterior two-thirds or more; male forepala usually digitiform with rather evenly curved row of pegs (Fig. 17.47); peg row interrupted and pala broadened basally only in A. chancea Hungerford (Fig. 17.48). .. Arctocovisa Wallengren

14'.

Median carina of pronotum well-defined only on anterior one-third (Fig. 17.41); male forepala broadened medially, peg row sharply curved (Fig. 17.22) or disjunct medially (Fig. 17.51) Ccnoconxa Hungerford Male strigil present; palar pegs usually in 1 row (Fig. 17.50) with notable exceptions (Fig. 17.49); ground color usually yellowish, dark pattern on hemelytra strongly contrasting (Figs. 17.54, 17.56-17.58) 5/jfarfl Fabricius Male strigil absent; palar pegs in 2 rows(Fig. 17.52); ground color greenish yellow, dark pattern on hemelytra weakly contrasting (Figs. 17.40 and 17.59); in all except C. audeni, posterior first tarsal segment with black spot(Fig. 17.53) Callicorixa White

15(13').

15'.

Gelastocoridae

1.

Foretarsus articulating with tibia, 1-segmented with 2 claws in nymphs and adults; forefemur only moderately enlarged at base, twice as long as basal width, not subtriangular (Fig. 17.61); closing face of forefemur flat and bordered by 2 rows of short spines; beak clearly arising at front of head, directed posteriorly; dorsal aspect of body as in Figure 17.62 (GELASTOCORINAE) Gelastocoris Kirkaldy

1'.

Foretarsus fused with tibia and terminated by a single claw (adults) or 2 claws(nymphs); forefemur very broad at base, about as long as broad, subtriangular (Fig. 17.60); closing face of forefemur with a dorsal flange-like extension that projects over tibia when it is closed against femur; beak appearing to arise from the back of the head, L-shaped; dorsal aspect of body as in Figure 17.63 (NERTHRINAE)Nerthra Say

Gerridae

1. r.

Inner margins of eyes sinuate or concave behind the middle (Figs. 17.75-17.77); body comparatively long and narrow (Fig. 17.79) Inner margins of eyes convex (Fig. 17.78), body comparatively short and broad (Figs. 17.66-17.68)

(GERRINAE) 2 6

2(1).

Pronotum shiny

3

2'.

Pronotum dull

4

3(2).

Forelobe of pronotum with pair of long, pale lines (Fig. 17.76)

3'. 4(2').

Forelobe of pronotum with single, median, pale spot(Fig. 17.77) Neogenis Matsumura Antennal segment I not less than nine-tenths of the combined lengths of II and III (Fig. 17.71) 5

Limnogonus Stal

Chapter 17 Aquatic and Semiaquatic Hemiptera

533

Figure 17.41 Figure 17.39 Figure 17.40

Figure 17.46

tergife VII

Figure 17.42

Figure 17.43

I

Figure 17.46 Figure 17.49

Figure 17.44

Figure 17.47

Figure 17.51

Figure 17.50

Figure 17.53 Figure 17.48

Figure 17.52

Figure 17.39 Corisella decolor (Uhler)(Corixidae; from Usinger 1956b). Figure 17.40 Callicorixa vulnerata (Uhler)(Corixidae; from Usinger 1956b). Figure 17.41 Cenocorixa kuiterti Hungerford (Corixidae; from Usinger 1956b). Figure 17.42 Dorsal view of Ramphocorixa sp. (Corixidae). Figure 17.43 Dorsal view of Palmacorixa sp. (Corixidae). Figure 17.44 Abdominal tergites of Palmacorixa sp. (Corixidae). Figure 17.45 Male foreleg of Ramphocorixa sp. (Corixidae). Figure 17.46 Palar claw of Ramphocorixa sp. (Corixidae). Figure 17.47 Male foreleg of Arctocorisa sutliis (Uhler), semi-diagrammatic view (from Brooks and Kelton 1967).

Figure 17.48 Male foreleg of Arctocorisa chanceae (Hungerford), semi-diagrammatic view (from Brooks and Kelton 1967). Figure 17.49 Male foreleg of Sigara faiienoidea (Hungerford), semi-diagrammatic view (from Brooks and Kelton 1967). Figure 17.50 Male foreleg of Sigara mathesoni (Hungerford), semi-diagrammatic view (from Brooks and Kelton 1967). Figure 17.51 Male foreleg of Cenocorixa sp. (Corixidae), semi-diagrammatic view (from Brooks and Kelton 1967). Figure 17.52 Male foreleg of Caiiicorixa sp. (Corixidae), semi-diagrammatic view (from Brooks and Kelton 1967). Figure 17.53 Posterior tarsus of Caiiicorixa sp. (Corixidae).

534

Chapter 17 Aquatic and Semiaquatic Hemiptera

Figure 17.58 Figure 17.57

Figure 17.55

Figure 17.59

Figure 17.56

Figure 17.54

I;

femur

Figure 17.60 Figure 17.62

Figure 17.63 ocellus

antenna

pronolum

Figure 17.61

pronotun

Figure 17.65

Figure 17.54 Dorsal view of Sigara mckinstryi Hungerford (Corixidae; from Usinger 1956b). Figure 17.55 Left hemelytron of Arctocorisa sp. (Corixidae; from Brooks and Kelton 1967). Figure 17.56 Left hemelytron of Sigara decoratella (Hungerford)(Corixidae; from Brooks and Kelton 1967). Figure 17.57 Left hemelytron of Sigara muiiettensis (Hungerford)(Corixidae; from Brooks and Kelton 1967). Figure 17.58 Left hemelytron of Sigara iineata (Forster)(Corixidae; from Brooks and Kelton 1967). Figure 17.59 Left hemelytron of Cailicorixa audeni Hungerford (Corixidae; from Brooks and Kelton 1967).

Figure 17.64

Figure 17.60 Foreleg of Nerthra sp.(Gelastocoridae). Figure 17.61 Foreleg of Gelastocoris sp. (Gelastocoridae). Figure 17.62 Dorsal view of Gelastocoris ocuiatus (Fabricius)(Gelastocoridae; from Brooks and Kelton 1967). Figure 17.63 Dorsal view of Nerthra martini Todd (Gelastocoridae; from Usinger 1956). Figure 17.64 Dorsal view of head and pronotum of Nerthra sp.(Gelastocoridae). Figure 17.65 Dorsal view of head and pronotum of Ochterus sp.(Ochteridae).

Chapter 17 Aquatic and Semiaquatic Hemiptera

535

4'.

Antennal segment I not more than eight-tenths of the combined lengths of II and III (Fig. 17.70) Limnoporus StkX

5(4).

Hind tibia at least 4.0 times as long as first tarsal segment; larger species, with total length at least 11 mm., with prominent connexival spines (Fig. 17.151) Aquarius Schellenberg Hind tibia not over 3.2 times as long as first tarsal segment; usually smaller species, with total length less than 10 mm., without prominent connexival spines (Fig. 17.152) Gems Fabricius Tibia and first tarsal segment of middle leg with fringe of long hairs (Fig. 17.80); always apterous(without trace of wing pads); the meso- and metanotum fused, without trace of a dividing suture; marine forms (HALOBATINAE). Halobates Eschscholtz

5'.

6(1').

6'.

7(6').

Tibia and first tarsal segment of middle leg without a fringe of long hairs; dimorphic(have both apterous and long-winged forms); the meso- and metanotum of apterous forms with a distinct dividing suture 7 Antennal segment 111 with several stiff bristles that are at least as long as diameter of segment(Fig. 17.74); length of antennal segment I much shorter than remaining 3 taken together; abdomen as long as remainder of body (Fig. 17.67) Rheumatobates Bergroth

7'.

Antennal segment III with fine pubescence or tuft of short, stiff bristles, but these not as long as diameter of segment; abdomen shorter than remainder of body (Fig. 17.68), or if subequal then length of antennal segment I about equal to remaining 3 taken together (Fig. 17.73) (TREPOBATINAE) 8

8(7').

Length of antennal segment I subequal to remaining 3 taken together (Fig. 17.73)

8'.

Length of antennal segment I much shorter than remaining 3 taken together (Fig. 17.72)

Metrobates \Jh\Qv Trepobates \Jh\ev

Hebridae

1.

Antennae distinctly shorter than greatest width of pronotum; antennal segments stout, 4th segment subequal in length to 1st segment(Fig. 17.83) Merragata White

1'.

Antennae distinctly longer than greatest width of pronotum; antennal segments slender, 4th segment much longer than 1st segment (Figs. 17.84 and 17.85) 2

2(1').

Antennal segment IV without a constriction (false joint structure) in the middle (Fig. 17.84)

2'.

Antennal segment IV with a constriction (false joint structure) in the middle, appearing 5-segmented (Figs. 17.82 and 17.85)

Lipogomphus Berg

Hebrus Curtis

Macroveliidae

1.

1'.

Ocelli absent; apterous; posterior margin of pronotum arcuate (arched), scutellum exposed (Fig. 17.89); antennal segments I-lII each longer than head width across eyes Oravelia Drake and Chapman Ocelli present, well developed (Fig. 17.88); macropterous or brachypterous; posterior margin of pronotum angular, concealing scutellum (Fig. 17.88); antennal segments I-III each shorter than head width across eyes (Fig. 17.87) Macrovelia Uhler

536

Chapter 17 Aquatic and Semiaquatic Hemiptera

Figure 17.68 Figure 17.67

Figure 17.66

Figure 17.71 Figure 17.69

Figure 17.70

Figure 17.66 Dorsal view of Mefrohafes frux infuscatus Usinger (Gerrldae; from Usinger 1956b). Figure 17.67 Dorsa\ \j\e\N of Rheumatobates rileyi Bergroth (Gerrldae; from Brooks and Kelton 1967). Figure 17.68 Dorsal view of 7repobafes beck; Drake and Harris (Gerrldae; from Usinger 1956b). Figure 17.69

Dorsal view of Ochterus barberi Schell

(Ochteridae; from Usinger 1956b).

Figure 17.72

Figure 17.70 Figure 17.71 Figure 17.72 Figure 17.73 Figure 17.74 (Gerrldae).

Figure 17.73 Figure 17.74

Antenna of Limnoporus sp.(Gerrldae). Antenna of Aquarius sp. (Gerrldae). Antenna of Trepobates sp.(Gerrldae). Antenna of Metrobates sp.(Gerrldae). Antenna of Rheumatobates sp.

Chapter 17 Aquatic and Semiaquatic Hemiptera

Figure 17.77

537

Figure 17.78

Figure 17.76 Figure 17.75

Figure 17.80

coxa

metasiernunn

omphalium

Figure 17.79 Figure 17.81

Figure 17.75 Dorsum of head and thorax of Aquarius sp.(Gerridae). Figure 17.76 Dorsum of head and thorax of Limnogonus sp.(Gerridae). Figure 17.77 Dorsum of head and thorax of Neogerris sp.(Gerridae).

Figure 17.78 Dorsum of head and thorax of Trepobates sp. (Gerridae). Figure 17.79 Dorsal view of Aquarius remigis (Say) (Gerridae; from Usinger 1956b). Figure 17.80 Hind leg of Halobates sp.(Gerridae). Figure 17.81 Ventral view of thorax of Gerridae.

538

Chapter 17 Aquatic and Semiaquatic Hemiptera

Micronectidae

1.

r. 2(F). 2'.

Head with a longitudinal elliptical carina (Fig. 17.153); foretibia and tarsus fused (Fig. 17.157) Synaptonecta L\xadh\2Ld Head without a longitudinal elliptical carina; foretibia and tarsus not fused (Fig. 17.156) 2 Pronotum crescent-shaped (Fig. 17.21); strigil absent in males Tenagobia Bergroth Pronotum lenticular (Fig. 17.154); strigil present in males(Fig. 17.155) Micronecta Kirkaldy

Naucoridae

1.

Anterior margin of pronotum straight or slightly concave behind interocular space

r. 2(1).

Anterior margin of pronotum deeply concave behind interocular space (Fig. 17.104) 3 Inner margins of eyes diverging anteriorly (Figs. 17.103 and 17.106); meso- and metasterna bearing prominent longitudinal carinae (keels) that are broad and foveate (with a deep impression) along middle; body broadly oval, sub-flattened; embolium may be dilated (Fig. 17.103), or produced outward and backward as an arcuate (arched), acute spine (Fig. 17.106) (LIMNOCORINAE)Limnocoris StM Inner margins of eyes converging anteriorly (Fig. 17.105); meso- and metasterna without longitudinal carinae at middle; body weakly convex above, the embolium rounded; embolium not dilated, or produced as an acute spine .. (NAUCORINAE)Pelocom Stil Posterior part of prosternum covered by plate-like extensions of propleura which are nearly contiguous at midline (Fig. 17.102); abdominal venter densely pubescent (hairy),

(Figs. 17.103, 17.105-17.106)

2'.

3(1').

2

except glabrous (shining) around spiracles, each spiracle also with a transverse row of small

3'.

glabrous areas behind; macropterous (Fig. 17.104) (AMBRYSINAE)Ambrysus Stal Prosternum completely exposed, separated from flattened pleura by simple sutures (Fig. 17.101); abdominal venter bare and with a perforated disk-like area near each spiracle; dimorphic, the brachypterous forms with hemelytra truncate (shortened and squared-off) at apices, about half as long as abdomen (CRYPHOCRICINAE)Cryphocricos Signoret

Nepidae

1.

1'.

2(1'). 2'.

Anterior lobe of pronotum not wider than head; body long, slender, cylindrical (Fig. 17.107); abdominal sterna of adult undivided; adult female subgenital plate laterally compressed, keel-like (RANATRINAE)Ranatra Fabricius Anterior lobe of pronotum wider than head (Fig. 17.110); body flattened; abdominal sterna of adult divided longitudinally into median and parasternites (Figs. 17.108 and 17.109); adult female subgenital plate broad, flattened (NEPINAE) 2 Median length of 6th sternite twice median length of 5th (Fig. 17.109) ... (NEPINl) Nepa Linnaeus Median length of 6th sternite about equal to length of 5th (Fig. 17.108)...(CURICTINI) Curicta Stil

Notonectidae

1.

r.

Hemelytral commissure with a definite hair-lined pit at anterior end (Figs. 17.113 and 17.117); antennae 3-segmented Buenoa Kirkaldy Hemelytral commissure without a definite hair-lined pit at anterior end (Figs. 17.112and 17.114); antennae 4-segmented

2(1').

not foveate (with a deep impression)

2'.

2

Eyes not holoptic (touching), separated dorsally (Fig. 17.114); intermediate femur with anteapical (before apex) pointed protuberance; anterolateral margins of prothorax Notonecta Linnaeus

Eyes holoptic, contiguous dorsally (Fig. 17.112); intermediate femur without anteapical pointed protuberance; anterolateral margins of prothorax foveate ... Martarega White

Chapter 17 Aquatic and Semiaquatic Hemiptera

539

Figure 17.86

Figure 17.83

Figure 17.84

Figure 17.85

ocellus

pronotum

Figure 17.82

pronotum

Figure 17.88

' scutellum

Figure 17

Figure 17.90

77777777 claws

rostrum

tarsus

Figure 17.92

Figure 17.91

Figure 17.87

Figure 17.82 Dorsal view of Hebrus sobrinus Uhler (Hebridae; from Usinger 1956b). Figure 17.83 Antenna of Merragata sp.(Hebridae). Figure 17.84 Antenna of Upogomphus sp. (Hebridae). Figure 17.85 Antenna of Hebrus sp.(Hebridae). Figure 17.86 Lateral view of head of Hebridae. Figure 17.87 Dorsal view of Macrovelia hornii Uhler (Macrovellldae; from Usinger 1956b).

Figure 17.88 Dorsum of head and pronotum of Macrovelia sp.(Macroveliidae). Figure 17.89 Dorsal view of Oraveiia pege Drake and Chapman (Macroveliidae). Figure 17.90 Dorsal view of Mesovelia sp. (Mesoveliidae). Figure 17.91 Lateral view of head of Mesovelia sp. (Mesoveliidae). Figure 17.92 Tarsus and claws of Macroveliidae.

540

Chapter 17 Aquatic and Semiaquatic Hemiptera

Pleidae

1. r.

Anterior tarsi each with 2 segments (Fig. 17.98); abdominal carinae (keels) on ventrites 2-6 Pamplea Esaki and China Anterior tarsi each with 3 segments (Fig. 17.99); abdominal carinae on ventrites 2-5 Neoplea Esaki and China

Saldidae

1.

Hemelytra with long embolar fracture reaching forward at least to level of posterior end ofclaval suture (Fig. 17.121) (CHILOXANTHINAE) 2

r.

Hemelytra with short embolar fracture, not reaching forward more than half-way from beginning of fracture on costal margin to level of posterior end of claval suture (Figs. 17.2 and 17.128) (SALDINAE) 3 Sublateral cell of membrane short, only half as long as lateral cell (Fig. 17.127) Chiloxanthus Reuter

2(1). 2'.

Sublateral cell of membrane subequal in length to lateral cell (Figs. 17.121 and 17.133)

Pentacom Renter

3(1').

Pronotum with 2 prominent conical tubercles on anterior lobe (Fig. 17.126)

3'. 4(3'). 4'. 5(4').

Pronotum without prominent tubercles on anterior lobe (Fig. 17.2) 4 Lateral margins of pronotum concave, humeral (posterolateral) angles produced (Fig. 17.132) Lampracanthia Router Lateral margins of pronotum straight or convex, humeral angles rounded (Fig. 17.131) 5 Hypocostal ridge simple, secondary hypocostal ridge absent (Fig. 17.137) 6

5'.

Hypocostal ridge complex, secondary hypocostal ridge present (Figs. 17.134-17.136)

6(5).

Innermost cell of membrane produced anteriorly one-half its length beyond base of 2nd cell (Fig. 17.120); 1st and 2nd antennal segments of male flattened, oval in cross section, the flattened sides glabrous(shiny) Calacanthia Renter Innermost cell of membrane produced anteriorly only slightly, not more than one-third its length beyond base of 2nd cell (Fig. 17.118); 1st and 2nd antennal segments of male not flattened, round in cross section, evenly pubescent (hairy) or pilose over entire surface Rupisalda Polhemus Second segment of tarsi usually nearly half again as long as 3rd (Fig. 17.129); innermost cell of membrane short, usually reaching only four-fifths the distance to apex of adjacent cell; outer corium with large, pale spots; clavus with yellow spot on each side in velvety black area (Fig. 17.131); secondary hypocostal ridge present, oblique, meets or projects to

6'.

7(5').

costal margin

7'.

8(7').

Saldoida Osborn

Teloleuca Renter

Second segment of tarsi subequal or slightly longer than 3rd segment(Fig. 17.130); innermost cell of membrane long, usually reaching almost to apex of adjacent cell (Figs. 17.119 and 17.128); outer corium with or without pale spots; clavus with or without yellow spot on each side in velvety black area; secondary hypocostal ridge present, may or may not meet or project to costal margin Males longer than 5.5 mm,females longer than 6 mm or, if shorter, then innermost cell of membrane produced two-fifths to one-half its length anteriorly beyond base of 2nd (Fig. 17.119), and dorsal surface unicolorous or with a few small, pale spots on corium and membrane (Fig. 17.125); secondary hypocostal ridge does not meet or project to costal margin (Fig. 17.134)

8'.

7

8

Salda Fabricius

Males shorter than 5.5 mm,females shorter than 6 mm;innermost cell of membrane

produced anteriorly only slightly beyond base of 2nd (Fig. 17.128) or, if inner cell is produced more strongly anteriorly, then dorsum has more or less extensive pale markings; secondary hypocostal ridge meets or projects to costal margin (Figs. 17.135 and 17.136)

9

Chapter 17 Aquatic and Semiaquatic Hemiptera

pronotum

541

Figure 17.95

scuteilum

Figure 17.96 tarsus c aws

Figure 17.97 Figure 17.93 Figure 17.98

prosternum

Figure 17.94

propleuron

Figure 17.99 embohum

Figure 17.101

Figure 17.103

antenna

gula rostrum

Figure 17.100 propleuron

Figure 17.102

-s

Figure 17.93 Dorsal view of Hydrometra australis Say (Hydrometrldae; from Usinger 1956b). Figure 17.94 Dorsal view of Hydrometra (Hydrometrldae). Figure 17.95 Lateral view of adult Pleidae. Figure 17.96 Dorsal view of Neoplea striola (Fleber) (Pleidae; from Brooks and Keiton 1967). Figure 17.97 Hind tarsus of Pleidae, Figure 17.98 Foretarsus of Paraplea sp.(Pleidae).

Figure 17.99 Foretarsus of Neoplea sp.(Pleidae). Figure 17.100 Lateral view of adult Naucoridae. Figure 17.101 Ventral view of liead and thorax of Cryphocricos sp.(Naucoridae). Figure 17.102 Ventral view of head and thorax of Ambrysus sp.(Naucoridae). Figure 17.103 Dorsal view of Limnocoris sp. (Naucoridae).

542

Chapter 17 Aquatic and Semiaquatic Hemiptera

Figure 17.105

Figure

Figure 17.104

abdominal

j;

sternltes abdominal

sternltes ^ °

Figure 17.109 Figure 17.108

Figure 17.110

3 siphon abdomen

Figure 17.107 Figure 17.111

Figure 17.104 Dorsal view of Ambrysus mormon Montandon (Naucorldae; from Menke etal. 1979). Figure 17.105 Dorsal view of Pelocoris shoshone LaRlvers (Naucorldae; from Usinger 1956). Figure 17.106 Dorsal view of Limnocoris moapensis (LaRlvers)(Naucorldae; from Usinger 1956b). Figure 17.107 Dorsal view of Ranatra brevicollis Montandon (Nepldae; from Usinger 1956b).

Figure 17.108 Ventral view of abdomen of Curlcta sp. (Nepldae). Figure 17.109 Ventral view of abdomen of Nepa sp. (Nepldae). Figure 17.110 Dorsal view of fiead and pronotum of Curicta sp.(Nepldae). Figure 17.111 Dorsal view of breathing tube of Nepa sp.(Nepldae).

Chapter 17 Aquatic and Semiaquatic Hemiptera

543

Figure 17.113 Figure 17.112 tarsus

claws

Figure 17.116 swimming hairs

Figure 17.115

rostrum

innermost cell of membrane hair-lined

Figure 17.117 Figure 17.119 Figure 17.118

Figure 17.121 Figure 17.120

Figure 17.112 Dorsal view of Martarega mexicana Truxal (Notonectidae; from Menke etal. 1979). Figure 17.113 Dorsal view of Buenoa scimitra Bare (Notonectidae; from Usinger 1956b). Figure 17.114 Dorsal view of Notonecta unifasciata Guerin-Meneviile (Notonectidae; from Usinger 1956b). Figure 17.115 Hind ieg of Notonectidae. Figure 17.116 Lateral view of adult Notonectidae. Figure 17.117 Dorsal view of Buenoa sp. (Notonectidae).

Figure 17.118 Left hemeiytron of Rupisalda sp. (Saididae). Figure 17.119 Left hemeiytron of Salda sp. (Saididae). Figure 17.120 Right hemeiytron of Calacanthia sp. (Saididae). Figure 17.121 Right hemeiytron of Pentacora sp. (Saididae).

544

Chapter 17 Aquatic and Semiaquatic Hemiptera

9(8').

Antennae relatively thick, the 3rd and 4th segments thicker than the distal end of the 2nd segment(Fig. 17.123); secondary hypocostal ridge (hrs) meets or projects to the costal margin at approximately three-fourths distance from base to embolar fracture; strigil (file) present on hrs (Fig. 17.136), plectrum (rasp) on distal portion of hind femur loscytus Reuter

9'.

Antennae relatively slender, the third and fourth segments not thicker than the distal

end of the second segment(Figs. 17.122 and 17.124); secondary hypocostal ridge meets

10(9'). 10'.

or projects to costal margin at three-fifths or less distance from base to embolar fracture (Fig. 17.135); strigil and plectrum absent 10 Veins of corium more or less distinct (Fig. 17.122); body usually more than 3.5 mm long; if less, then anterior margin of pronotum wider than collar Saldula Van Duzee

Veins of corium obsolete (Fig. 17.124); body usually less than 3.5 mm long; ahterior margin of pronotum usually narrower than collar

Micracanthia Reuter

Veliidae

1.

1'. 2(1'). 2'. 3(2). 3'. 4(2'). 4'.

Middle tarsi deeply cleft, with leaf-like claws and plumose (plume-like) hairs arising from the base of cleft (Fig. 17.146); hind tarsi 2-segmented (Fig. 17.141) or 3-segmented (Fig. 17.144) (RHAGOVELIINAE) Mayr Middle tarsi not deeply cleft and without plumose hairs arising from the base of cleft 2 Tarsal formula 1:2:2 (Fig. 17.142) (MICROVELIINAE) 3 Tarsal formula 3:3:3 (VELIINAE) 4 Middle tarsi with 4 leaf-like blades arising from cleft(Fig. 17.143) Hmseyella Herring Middle tarsi with narrow claws arising from cleft (Fig. 17.142) Microvelia Westwood Body broad (Fig. 17.147); metasternum with lateral tubercles meeting mesoacetabulae (Fig. 17.150) Polhemus and Polhemus Body narrow (Fig. 17.148); metasternum with lateral tubercles meeting mesocoxae (Fig. 17.149)

Steinovelia Polhemus and Polhemus

Chapter 17 Aquatic and Semiaquatic Hemiptera

545

Figure 17.124

Figure 17.123

Figure 17.122

.sublateral cell

Figure 17.125

Figure 17.128

tarsal segments

Figure 17.129

Figure 17.126

Figure 17.130 Figure 17.127

Figure 17.122 Dorsal view of Saldula pexa Drake (Saididae; from Usinger 1956b), Figure 17.123 Dorsal view of loscytus politus (Uhler) (Saididae; from Usinger 1956b). Figure 17.124 Dorsal view of Micracanthia quadrimaculata (Champion)(Saididae; from Usinger 1956b).

Figure 17.125 Dorsai view of Salda buenoi (McDunnough)(Saididae; from Usinger 1956b).

Figure 17.126 Lateral view of head and thorax of Saldoida sp.(Saididae). Figure 17.127 Right hemelytron of Chiloxanthus sp. (Saididae). Figure 17.128 Right hemelytron of Saldula sp. (Saididae). Figure 17.129 Tarsus of Teloleuca sp. (Saididae). Figure 17.130 Tarsus of Salda sp.(Saididae).

546

Chapter 17 Aquatic and Semiaquatic Hemiptera

Figure 17.132 Figure 17.133

Figure 17.131

secondary

secondary hypocostal ridge

hypocostal ridge

-hypocostal ridge

costal margin

Figure 17.137

Figure 17.135

Figure 17.134 Figure 17.136

Figure 17.131 Dorsal view of Teloleuca bifasciata Thomson (Saldldae; from Brooks and Kelton 1967). Figure 17.132 Dorsal view of Lampracanthia crassicornis (Uhler)(Saldidae; from Brooks and Kelton 1956b). Figure 17.133 Dorsal view of Pentacora signoreti (Guerin-Menevllle)(Saldidae; from Usinger 1956b). Figure 17.134 Ventral view of hemelytron of Salda sp.(Saldidae).

Figure 17.135 Ventral view of hemelytron of Saldula sp.(Saldidae). Figure 17.136 Ventral view of hemelytron of loscytus sp.(Saldidae). Figure 17.137 Ventral view of hemelytron of Rupisalda sp.(Saldidae).

Chapter 17 Aquatic and Semiaquatic Hemiptera

547

pronotum

—I—membrane Figure 17.138

Figure 17.140

Figure 17.139

Figure 17.141 Figure 17.142

tarsus

claws

Figure 17.144

Figure 17.143

scent groove coxae

Figure 17.145

Figure 17.146

Figure 17.138 Dorsal view of Rhagovelia distincta Champion (Veliidae; from Usinger 1956b). Figure 17.139 Dorsal view of Microvelia sp. (Veliidae). Figure 17.140 Dorsal view of Microvelia beameri McKinstry (Veliidae; from Usinger 1956b). Figure 17.141 Dorsal view of hind tarsus of Trochopus sp. (Veliidae). Figure 17.142 Lateral view of middle tarsus of Microveiia sp. (Veliidae).

Figure 17.143 Lateral view of middle tarsus of Husseyelia sp. (Veliidae). Figure 17.144 Lateral view of hind tarsus of Rhagoveiia sp. (Veliidae). Figure 17.145 Lateral view of thorax of Microveiia sp. (Veliidae). Figure 17.146 Middle tarsus of Rhagovelia sp. (Veliidae) showing swimming plume.

548

Chapter 17 Aquatic and Semiaquatic Hemiptera

Figure 17.147

Figure 17.148

Figure 17.151

Figure 17.150

Figure 17.152

Figure 17.149

Figure 17.147 Platyvelia brachialis (St§l), dorsal view (Velildae).

Figure 17.148 Steinovelia sp., dorsal view (Veliidae). Figure 17.149 Ventral view of thorax of Steinovelia sp. (Veliidae). Figure 17.150 Ventral view of thorax of Platyvelia sp. (Veliidae).

Figure 17.151 Lateral view of abdominal terminalia of Aquarius sp.(Gerridae). Figure 17.152 Lateral view of abdominal terminalia of Gerrls sp.(Gerridae).

Chapter 17 Aquatic and Semiaquatic Hemiptera

549

Figure 17.154 Figure 17.153

Figure

Figure 17.157

Figure 17.156

Figure 17.153 Synaptonecta issa (Distant), dorsal view (Corixidae; from Polhemus and Rutter 1997). Figure 17.154 Micronecta ludibunda Breddin, dorsal view (Corixidae; from Lundblad 1933). Figure 17.155 Sixth abdominal tergite of Micronecta sp.(Corixidae), sfiowing strigil at right (from Hutchinson 1940).

Figure 17.156 Male foreleg of Micronecta iudibunda Breddin (Corixidae; from Lundblad 1933).

Figure 17.157 Male foreleg of Synaptonecta issa (Distant)(Corixidae), showing fused tibio-tarsus (from Hutchinson 1940).

550

Chapter 17 Aquatic and Semiaquatic Hemiptera

ADDITIONAL TAXONOIVIIC REFERENCES

Taxonomic treatments at the family and generic

General VanDuzee (1917); Hungerford (1920, 1958, 1959); Parshley (1925); Blatchley (1926); China (1955); Usinger tl956a); China and Miller (1959); Lawson (1959); Polhemus(1966, 1973); Brooks and Kelton (1967); Cobban (1968); Ruhoff (1968); Jaczewski and Kostrowicki (1969); DeCoursey (1971); Herring and Ashlock (1971); Miller (1971); Bobb (1974); Stys and Kerzhner (1975); Cheng (1976); Slate and Baranowski(1978); Andersen (1981a, 1982); Henry and Froeschner (1988); Stys and Jansson (1988); Yonke (1991); Spence and Andersen (1994). Dolling (1991); Mahner (1993); Schuh and Slater (1995); Schaefer(1996); Polhemus(1997); Poole and Gentili (1997); D. Polhemus

levels Belostomatidae: Menke (1958, 1960, 1963); Lauck and Menke

and J. Polhemus (2002).

Regional faunas Arizona; Polhemus and Polhemus(1976); Blinn and Sanderson (1989); Stevens and Polhemus(in press) Arkansas: Kittle (1980); Farris and Harp (1982). British Columbia: Scudder (1977). California: Usinger (1956a); Menke et al.(1979). Central Canada: Strickland (1953); Brooks and Kelton (1967). Connecticut: Britton (1923). Florida: Herring (1950, 1951a); Chapman (1958); Epler (2006). Idaho: Harris and Shull(1944). Illinois: Lauck (1959); Tinerella et al. (2009). Kansas: Slater (1981).

Louisiana: Ellis (1952); Gonsoulin (1973a,b,c, 1974, 1975). Minnesota: Bennett and Cook (1981). Mississippi: Wilson (1958); Lago and Testa (1989). Missouri: Froeschner (1949, 1962). Montana: Roemhild (1976). New Jersey: Chapman (1959). North Carolina: Sanderson (1982a). Oklahoma: Schaefer (1966); Schaefer and Drew (1964, 1968). Quebec: Chagnon and Fournier (1948). Rhode Island: Reichart (1976, 1977, 1978). South Carolina: Sanderson (1982a). South Dakota: Harris(1937). Texas: Millspaugh (1939). Virginia: Bobb (1974).

Wisconsin: Hilsenhoff(1981, 1984, 1986, 1995). Yukon: Scudder(1997)

(1961); Lauck (1963, 1964).

Corixidae: Hungerford (1948); Sailer (1948); Lansbury (1960); Hilsenhoff (1970); Applegate (1973); Scudder (1976); Nieser (1977); Jansson (1978, 1981); Dunn (1979); Jansson and Polhemus (1987); Lundblad (1933); Hutchinson (1940); Polhemus and Rutter (1997); Chordas and Armitage (1998); Chordas and Hudson (1999); Tinerella and Gundersen (2005); Polhemus and Golia (2007). Dipsocoridae: McAfee and Malloch (1925); Stys (1970). Gelastocoridae: Martin (1928); Todd (1955, 1961); Polhemus and Lindskog (1994). Gerridae: Anderson (1932); Drake and Harris(1932, 1934); Deay and Gould (1936); Kuitert(1942); Hussey and Herring (1949); Hungerford (1954); Hungerford and Matsuda (1960k Matsuda (1960); Herring (1961); Cheng and Fernando (1970); Scudder (1971b); Calabrese (1974); Kittle (1977b,c, 1982); Stonedahl and Lattin (1982); Cheng (1985); Spangler, Froeschner; and Polhemus(1985); Polhemus and Spangler(1989); Andersen (1990, 1991); Andersen and Spence (1992); Polhemus and Polhemus (1993b, 1995); Gallant and Fairbairn (1996); J. Polhemus and D. Polhemus(2002); Andersen and Cheng (2004). Hebridae: Porter (1950); Drake and Chapman (1954, 1958a); Polhemus and Chapman (1966); Andersen (1981b); Polhemus and McKinnon (1983). Hydrometridae: Torre-Bueno (1926); Hungerford and Evans (1934); Drake and Lauck (1959). Leptopodidae: Schuh, Galil, and Polhemus(1987). Macroveliidae: McKinstry(1942). Mesoveliidae: Jaczewski (1930); Andersen and Polhemus(1980). Naucoridae: Usinger (1941); LaRivers(1949,1951, 1971, 1974, 1976); Polhemus and Polhemus (1994); Polhemus and Sites (1995); Sites and Polhemus (1995); Davis (1996). Nepidae: Hungerford (1922c); Polhemus (1976b); Sites and Polhemus (1994); Keffer (1997). Notonectidae: Hungerford (1933); Hutchinson (1945); Truxal (1949, 1953); Scudder (1965); Reichart(1971); Voigt and Garcia (1976); Zalom (1977). Ochteridae: Schell (1943); Polhemus and Polhemus(1976); Polhemus and Polhemus(2016). Pleidae: Drake and Chapman (1953); Drake and Maldonado (1956).

Saldidae: Hodgden (1949a,b); Drake (1950, 1952); Drake and Hoberlandt 0950); Drake and Hottes(1950); Drake and Chapman (1958b); Chapman (1962); Polhemus (1967, 1976c, 1985, 1994); Schuh (1967); McKinnon and Polhemus(1986); Schuh, Galil, and Polhemus(1987); Lindskog and Polhemus(1992). Schizopteridae: Emsley (1969). Veliidae: Drake and Hussey (1955); Bacon (1956); Polhemus (1974, 1976a); Smith and Polhemus (1978); Smith (1980); Polhemus and Polhemus(1993a); D. Polhemus (1997).

cn

Family

hornii

pege

Oravelia

Species

Macrovelia

Hydrometra (9)

Genus

Predators

(piercers)

on surface

film)

Climbers—

sprawlers (rarely in or

Lotic—margins (semiaquatic)

Predators

(piercers)

Predators

(piercers)

Climbers—

sprawlers

Climbers—

sprawlers

and depositional margins (in protected areas) Lotic—margins

on the water) Lotic—erosional

**Emphasis on trophic relationships

ostracods)

Predators

(piercers)(adult arthropods, dead or live, especially mosquito larvae and pupae, and

"Skaters"

(slow walkers

margins

Trophic Relationships

Lentic—limnetic, surface (in littoral zone); lotic—

Habitat Habit

HEMIPTERA

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic

Bugs

- Macroveliid Shore

Macroveliidae (2)

Water Measurers

- Marsh Treaders,

Hydrometridae (9)

Hemiptera - True Bugs

Order

of species in parentheses)

Taxa(number

Merritt.)

(For definition of terms see Tables 6A-6C; table prepared by J. T. Polhemus, K. W. Cummins and R. W.

Table 17A Summary of ecological and distributional data for Hemiptera (aquatic and semiaquatic bugs).

North

West

West

Widespread

American Distribution

SE UM

M NW

3995

3956, 3995, 6103, 3972

135, 536, 2786, 3179, 3956, 4599,6103

1435, 2628, 2776, 3179, 3676, 3995, 4151,4538, 3386, 4932, 5614, 6704

135, 536, 565, 2786, 2828, 3179, 3362, 3403, 4599, 5274, 6103, 6423

Ecological References**

(continued)

MA*

To erance Values

) ) ) ) ) )) J > ))) J )3 J :) > )) 3 D ))))3

Lentic—limnetic;

Steinovelia

surface

depositional

lotic—

Lentic—limnetic;

bays)

Lotic—erosional

surface; lentic— limnetic (saltwater

surface

depositional

lotic—

Lentic—limnetic;

surface

depositional

lotic—

Rhagovelia (10)

stagnalis

Lotic and lentic—

surface (brackish water)

(=Trochopus)

Platyvelia (3)

Microvelia (20)

tumalis

lotic—surface

Water Striders

Husseyella

Generally lentic—

Habitat limnetic surface;

Species

Broad-Shouldered

Genus

Veiiidae (35)-

Family

Continued

Skaters

Skaters

Skaters

Skaters

Skaters

Skaters

Habit

**Emphasis on trophic relationships

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW == Northwest, MA = Mid-Atlantic

Order

of species in parentheses)

Taxa(number

Table 17A

North

South, East, Midwest

Predators

Florida coast

southern

Widespread,

South, Southwest, East, Midwest

Widespread

Florida

Southern

Distribution

American

(piercers)

(piercers) (scavengers)

Predators

(piercers)

Predators

(piercers)

Predators

(piercers)

Predators

Generally predators (piercers)(live and dead arthropods)

Trophic Relationships SE

UM

M

NW

6.0

MA*

Tolerance Values

Ecological

1435, 2684, 3995, 6103

1435, 3995, 4151, 4538, 4920, 5061, 6624, 140, 3995

1001, 1421,

1435, 3995, 6103, 573

1435, 1469, 1981,412,4151, 4918, 4920, 6012, 2536, 2537, 6014, 6103, 2538, 4088, 4252, 6039, 5919, 6778, 573

140, 3995

2828, 3179, 3362, 3403, 3995, 4599, 4932, 5274, 6103, 716, 6423

2776, 2786,

135, 140, 536, 565, 2553, 2628,

References**

) ) ) ) I ) ) ) ) ) )) ) ) ) )) ) )) ))) ) ) ) 1

'J\ 'Ji

(J\ 'J\

**Emphasis on trophic relationships

HEMIPTERA

Midwest, NW = Northwest, MA = Mid-Atlantic

surface

depositional

lotic—

Generally lentic— limnetic surface;

Generally predators (piercers) (scavengers)

Florida

bispina

Schizoptera

Gerridae (46)-

Florida

arenarius

Nannocoris

Louisiana

major

Tennessee,

Georgia

California,

Corixidea

Skaters

Borrowers

Southeast

New Mexico

New Mexico

East, California,

Distribution

burrowers

(active)

edge)

North American

West,

margins(under stones at stream

Trophic Relationships

Generally

Lotic—erosional

Generally lotic— erosional margins

Lentic

Habitat

Cryptostemma

latipennis

Species Habit

(3)

Leptonannus

(3)

Ceratocombus

Genus

Water Striders

Schizopteridae (3)

Dipsocoridae (4)(= Cryptostemmatidae)

Ceratocombidae (4)

Family

Continued

*SE = Southeast, UM = Upper Midwest, M :

Order

of species in parentheses)

Taxa (number

Table 17A

SE

UM

M

5.0

NW

Ecological

2828, 3179, 3362, 3403, 3995, 4599, 4907, 5032, 5061, 5274, 5598, 5601, 6103, 6423, 2172, 2453, 6366, 2707

135, 140, 536, 565, 2553, 2628, 2776, 2786,

1663

1663

6103, 6101

5274, 6103

1663

References**

(continued)

MA*

Tolerance Values

) ) ) )) ))) j j j ) j j j ))))) :> ) ) )) j

)

)

(J\ (Jt ■u.

Gerrinae (22)

Continued

Lentic—limnetic

hesione

Neogem's

Lentic—limnetic

littoral surface

limnetic and

surface; lentic-

depositional

Lotic—

littoral surface

limnetic and

depositional surface; lentic-

Lotic—

Habitat

Lentic—limnetic

frandscanus

Species

Limnoporus (4)

Limnogonus

Gem's (10)

Aquarius (6)

Genus

)

)

)

)

**Emphasis on trophic relationships

)

)

)

)

)

)

)

Habit

Skaters

Skaters

Skaters

Skaters

Skaters

)

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic

Order

of species in parentheses)

Taxa (number

Table 17A

)

Trophic

)

(piercers)

Predators

(piercers)

Predators

)

)

South, East

Widespread

South

(primarily tropical)

Widespread

Widespread

Predators

)

American Distribution

(piercers)

(piercers) (scavengers)

Predators

(piercers) (scavengers)

Predators

Relationships

North

)

SE

)

UM

M

)

5.0

NW

)

MA*

Tolerance Values

Ecological

)

)

)

563, 837, 838, 1435, 2939, 2940,4151, 4538, 6152, 6167

837, 838, 847, 2939, 2940, 3995, 4151, 4538, 4932, 5595, 6103, 6152, 3041, 3263, 4403, 5597, 5599, 6167, 998, 1008, 2828, 3995, 4158, 1204, 2959

2940, 3293, 3995, 4151, 4220, 4538, 4920, 4087, 6103,6152, 6551, 3925, 4495, 6686, 2959, 2958

2030, 2939,

753, 837, 838,

References**

)

t/i ui

in

Belostomatidae (19) - Giant Water Bugs

Trepobatinae (14)

Rhagadotarsinae (8)

Halobatinae (2)

Family

Continued

Trepobates(8)

Metrobates(6)

littoral

detritus); lentic—

hydrophytes and

(vascular

Generally lotic— depositional

surface

surface; lotic— depositional

Lentic—limnetic

surface (usually large rivers)

Lotic—erosional

Lotic and lentic—

surfaces

protected reefs)

ocean and

Marine (open

Habitat

Rheumatobates

Species

(8)

Halobates(2)

Genus

**Emphasis on trophic relationships

Predators

(piercers)

East, West, South

(piercers)

East)

Widespread (especially

Predators

(piercers)

Predators

East, Central, South

Predators

(piercers)

East and West Coasts

Distribution

Predators

Climbers—

HEMIPTERA

North American

(piercers) (scavengers)

Trophic Relationships

swimmers

Skaters

Skaters

Skaters

Skaters

Habit

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW ; : Northwest, MA = Mid-Atlantic

Order

of species in parentheses)

Taxa (number

Table 17A

SE

UM

M

10.0

NW

Ecological

536, 565, 1244, 2553, 2628, 2776, 2786, 2828, 3178, 3362, 3995, 3999,4151, 4599, 4907, 5274, 1525, 5813, 6103, 6423, 5252, 5914, 6161, 675

3995, 4538, 6103

4538, 6103, 2959, 2958

3995,4151,

5451

140, 3995, 4538,

140, 2555, 3995, 6103, 2535

References**

(continued)

MA*

Tolerance Values

) ) ))))) j > :j )) D ))))>)) ) ))) ) )

Nepidae (13)Water Scorpions

Continued

Lethocerus(4)

Benacus

Belostoma (8)

Abedus(6)

Genus

gnseus

Species

**Emphasis on trophic relationships

Trophic

swimmers)

(vascular

littoral

detritus); lentic—

hydrophytes and

Predators

(piercers)

Climbers

(poor

(piercers)

Predators

(piercers)

Predators

(piercers)

Predators

(piercers)

Predators

Relationships

Generally lotic— depositional

lentic—littoral

Climbers— swimmers

Lotic—

depositional;

lentic—littoral

Climbers— swimmers

Lotic—

depositional;

lentic—littoral

Climbers— swimmers

Lotic—

depositional;

Climbers— swimmers

Lotic—

Habit

depositional

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic

Order

of species in parentheses)

Taxa (number

Table 17A

Widespread

East

Widespread

Southeast

Southwest,

Distribution

American

North

) ) ) ) ) ) ) ) ))) ) ) ) ) ))))

in o\

9.(

SE

M

NW

MA*

Ecological

536, 565, 2553, 2776, 2779, 2786, 2828, 4599, 3179, 3362, 3403, 3995, 4907, 5274, 6103

1240, 1435, 2828, 3995, 2204, 4869, 5061, 6103,

5531, 5700, 1115, 6103, 6349, 1222, 3086, 3121, 572, 805, 5205, 924, 992, 2195, 6778, 1204, 1901

2183, 3260, 3995, 4538,

1240, 1468,

2808, 3995, 5530, 5532, 6103, 6349, 5533, 6159, 574

References**

) ^ )) )) 1

DM

Tolerance Va ues

Ul

Backswimmers

Pleidae (5) - Pygmy

Family

Continued

Paraplea (2)

Neoplea (3)

Ranatra (10)

Nepa

Curicta (2)

Genus

apiculata

Species

Predators

(piercers)

Climbers

(poor swimmers)

Lotic—

depositional hydrophytes);

hydrophytes (especially dense

climbers

hydrophytes (especially dense stands)

SwimmersLentic—^vascular

stands)

Swimmersclimbers

Lentic—^vascular

lentic—littoral

**Emphasis on trophic relationships

HEMIPTERA

microcrustacea)

(piercers) (especially

Predators

microcrustacea)

(piercers) (especially

Predators

Predators

(piercers)

Climbers

(poor swimmers)

Ecological

(continued)

1507

Southeast

548, 1359, 1362, 1435, 2041, 2628, 3995, 4151, 4041, 4538, 4861, 5061, 6011, 1070, 1941, 4087, 6423, 4872, 5202, 5912, 638, 6162, 1204, 1901

3995, 6103

3995, 4599, 6561

References**

301, 367, 536, 565, 1435, 1651, 2138, 2140, 2388, 2776, 2786, 3179, 3324, 4599, 1096, 4920, 4991, 6103, 5876, 3968, 1901

7.5

Tolerance Values

Widespread

Widespread

Central, East

Southwest, South

Predators

(piercers)

Distribution

Climbers

Lentic—vascular

(vascular

North American

(poor swimmers)

Habit

Trophic Relationships

hydrophytes

Lentic—littoral

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = : Northwest, MA == Mid-Atlantic

Order

of species in parentheses)

Taxa (number

Table 17A

00

Pelocoris(4)

Climbers— swimmers

Lentic—^vascular

Clingers

Clingers

hydrophytes

warm springs)

Lotic—erosional

(in sediments,

(sediments)

Lotic—erosional

hydrophytes)

Limnocoris(2)

hungerfordi

vascular

(sediments and

Clingers— swimmers

Lotic and lentic—

lentic—littoral

erosional;

erosional

(=Usingenna)

Cryphocricos

Ambrysus 05)

Habit

Generally

Habitat

lotic—

Species

Naucoridae (22)

Genus

- Creeping Water Bugs

Continued

**Emphasis on trophic relationships

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic

Order

of species in parentheses)

Taxa(number

Table 17A

North

Belostomatidae)

Predators (piercers)(Diptera,

(piercers)

Predators

(piercers)

Predators

East, Central, Southwest

Texas, Nevada

Texas

West, Southwest

Predators

(piercers)

Predators

Distribution

American

(piercers)

swimmers

Generally clingers;

Relationships

Trophic

5.0

7.0

Tolerance Va ues

Ecological

565, 1435, 2388, 2781, 3407, 3727, 3995, 3971, 4151, 4538, 5061, 6010, 6103

3995, 4532, 5779, 6103, 5484, 5789, 6676

5481, 5484, 5789, 6624

4532, 4533, 5779, 6103,

293, 753, 3995, 6102, 5789, 6103, 6624, 5480, 5484, 5485

3403, 3995, 4599,4881, 5274, 6103, 6423

536, 809, 2553, 2776, 2786, 2832, 3179,

References*

vo

Lentlc

Lentlc

Arctocorisa (5)

Calllcorixa (6)

hydrophytes)

(vascular

hydrophytes: lotlc—deposltional

Generally swimmers

Habit

Generally lentlc—

Habitat vascular

Species

Corixidae (128)

Genus

-Water Boatmen

Continued

**Emphasis on trophic relationships

HEMIPTERA

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = : Northwest, MA = Mid-Atlantic

Order

of species in parentheses)

Taxa (number

Table 17A

(piercers)

Predators

scrapers

predators (engulfers and piercers) or

herbivores; some

Generally piercers—

Trophic Relationships

North

North

Distribution

American

North

9.0

10.0

5.0

Tolerance Values

(continued)

3995, 4490, 4491, 4493, 4494, 4495, 4496, 5256, 4871, 1204

4495, 4496, 5700, 4871

4490, 4491, 4493, 4494,

536, 565, 753, 809, 1207, 2137, 2244, 2553, 2628, 2773, 2776, 2783, 2786, 2828, 2861, 2948, 2949, 3179, 3362, 3401, 3403, 3610, 3614, 3616, 3995, 4151, 4491, 4599, 4644, 4834, 4907, 5032, 5061, 5274, 5384, 5738, 5813, 5835, 6013, 6103, 6423, 2486, 6802, 2952, 6009, 6492

References**

Ecological

) ) ))) )

Family

Continued

Neocorixa

(2)

**Emphasis on trophic relationships

snowi

Lentic—littoral

Midwest, NW = Northwest, MA = Mid-Atlantic

Ramphocorixa

Palmacorixa (4)

(engulfers)

Predators

herbivores

Widespread (except West)

extreme North)

Widespread (except

Southwest

Southwest

Predators

(piercers); piercers—

Morphocorixa (3) (=Pseudocorixa)

Extreme

Widespread

Southwest

herbivores

Piercers—

Krizousacorixa

Swimmers; climbers

Lotic—

depositional

West

(piercers)

(3){=Ahautlea)

Hesperocorixa (19)

Southwest,

Predators

North

Graptocorixa (6)

propinqua

North

West, North

Florida, Texas

Northwest

American Distribution

Claenocoris

lakes)

water and saline

Predators

(piercers)

(piercers)

Predators

Trophic Relationships

Lentic—littoral

Habit

(including brackish

Lentic

Habitat

North

americana

nigripennis

Species

North

Dasycorixa (3)

Cymatia

Corisella (4)

Centrocorisa

Cenocorixa (8)

Genus

*SE = Southeast, UM = Upper Midwest, M :

Order

of species in parentheses)

Taxa (number

Table 17A

SB

UM

M

8.0

NW

5.0

MA*

Tolerance Values

2244, 3995, 4151

4151

3995, 4643

564, 1435, 3995, 4151, 1901

1947, 3995

1700,2530,4471

2962, 3402, 4914

3995,4151, 6103, 6349, 1901

2954, 4975, 4976

Ecological References*

) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) > ) ))))))) 1

as

o\

Micronectidae (3)

Family

Continued

ludibunda

issa

mexicana

Synaptonecta

Tenagobia

Species

Micronecta

Trichocorixa (11)

Sigara (50)

Genus

**Emphasis on trophic relationships

HEMIPTERA

(primarily tropical)

Florida

Florida

Southwest,

herbivores

Piercers—

Lotic—

Swimmers

Florida

(primarily tropical)

Southwest, herbivores

(primarily tropical)

Southwest, Florida

Widespread

Widespread

Piercers—

herbivores

Piercers—

North

American Distribution

depositional

Lentic—littoral

Lentic—littoral

Swimmers

gatherers (as early

sea)

Lentic—littoral

Oligochaeta); some collectors—

instars)

chironomid larvae,

including intertidal pools and offshore

(piercers) (especially

Predators

gatherers

collectors—

herbivores;

Piercers—

Trophic Relationships

saltwater

Swimmers

climbers

brackish or

Swimmers;

Lentic—littoral

Swimmers; climbers

Habit

(freshwater, some

depositional

Lotic—

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic

Order

of species in parentheses)

Taxa (number

Table 17A

SE

UM

8.0

M NW

5.0

9.0

3498, 4770

2949, 3498, 4772, 4770

2949, 3498, 4772, 4770

1901

860, 1364, 1365, 2290, 2828, 3104, 3995, 4151,4538, 5274, 6103, 6773, 4471, 4920, 32, 1119,

1435, 2774, 2776, 2783, 3271, 3995, 4151,4920, 4471, 5846, 1901

Ecological References"

(continued)

MA*

Tolerance values

- Backswimmers

Notonectidae (32)

Family

Continued

Martarega

Buenoa (14)

Genus

mexicana

Species

(rest at

depositional

open water)

surface in

Swimmers

Lotic—

'*Emphasis on trophic relationships

(piercers)

Predators

submerged in hydrostatic

depositional balance)

Predators

(piercers)

Swimmers

(rest

Lentic—littoral;

Habit

Trophic Relationships

lotic—

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic

Order

Taxa (number of species in parentheses)

Table 17A

North

Southwest

Widespread

Distribution

American

SE

UM

M

NW

MA*

Tolerance Values

Ecological

2756, 3995

1901

4151, 5738, 6018, 6103, 1115, 1117, 6349, 6836, 4,

1435, 2141, 2828, 3995,

2776, 2777, 2782, 2828, 3179, 3362, 3403, 3456, 3616, 4599, 4907, 4991, 5274, 5813, 6103, 4567, 5913, 6423, 4091

367, 536, 565, 2553, 2632,

References**

) ) ) > ) ) ) ) ) ) ) ) > )) ) 1 )))) ) ) ) ) ) 1

Ul 0\

U)

OS

'Ji

Family

Continued

Notonecta (17)

Genus

Species

depositlonal

climbers (rest (piercers) submerged or (including at surface) cannibalism)

**Emphasis on trophic relationships

HEMIPTERA

Predators

Swimmers—

Lentic—littoral;

Habit

Trophic Relationships

lotic—

Habitat

*SE = Southeast, UM = Upper Midwest, M : : Midwest, NW = Northwest, MA = Mid-Atlantic

Order

of species in parentheses)

Taxa (number

Table 17A

North

Widespread

American Distribution SE

UM

M NW

301, 753, 1362, 1435, 1654, 1710, 1947, 1948, 2116, 2137, 2140, 2628, 3293, 3324, 3966, 3995, 4538, 4991, 5061, 5444, 5700, 5786, 5791, 5917, 6017, 6103, 2538, 4756, 6349, 6836, 1009, 1096, 1115, 1117, 1210, 1313, 1459, 1941, 2113, 2486, 2389, 2657, 2962, 4087, 4214, 4215, 4216, 4466, 5445, 5488, 5787, 5788, 5789, 5968, 5790, 213, 4, 2187, 537, 538, 668, 4311, 5707, 1204, 1901

Ecological References**

(continued)

MA*

Tolerance Values

brevis

**Emphasis on trophic relationships

"burrowers"

(under stones at water's

edge)

vascular

hydrophytes (emergent zone and sediments at

(under stones at water's

edge)

hydrophytes (emergent zone and sediments at

water's edge)

Climbers— "burrowers"

Lentic—littoral

mats of floating algae) vascular

Skaters— climbers

Lentic—littoral (on

water's edge)

Climbers—

Midwest, NW = Northwest, MA = Mid-Atlantic

Lipogomphus

Merragata (2)

Generally climbers (at shore. semiaquatic)

Lentic—littoral

and detritus)

hydrophytes (emergent zone

Generally lentic—

marshes)

climbers (or sprawlers at

water's edge, semiaquatic)

Skaters—

Lentic—vascular

Habit

hydrophytes (emergent and floating zone including salt

Habitat

littoral vascular

Hebrus(M)

Species

Hebridae (15)-

Mesovelia (3)

Genus

Velvet Water Bugs

- Water Treaders

Mesoveliidae (3)

Continued

*SE = Southeast, UM = Upper Midwest, M :

Order

of species in parentheses)

Taxa(number

Table 17A

(piercers)

Predators

(piercers)

Predators

(piercers)

Predators

Generally predators (piercers)

(piercers) (scavengers)

Predators

Trophic Relationships

South

Widespread

Widespread

Widespread

American Distribution

North

SE

UM

M

NW

MA*

Tolerance Va ues

1502, 2388, 3995, 4599

2776, 3995, 6103

6103, 6423

3999, 4599, 4788, 5274,

3179, 3995,

135, 536, 565, 2776, 2786,

135, 536, 565, 1435, 2678, 2775, 2776, 2780, 2786, 2828, 3104, 3179, 3362, 3995, 4151, 4538, 4599, 5061, 5498, 5700, 6103, 2537, 6423, 5918, 5919, 1204

Ecological References**

) )) ) ) ) ) > ) ) ) ) ) ) ) ) ) )) ) )) ) ) ) ) )

in o\

ifi

OS 'Ji

Saldidae (59) ■ Shore Bugs

Family

Continued

**Emphasis on trophic relationships

northern

(piercers)

and freshwater;

Pentacora (6)

Rupisalda (3)

margins

Lentic and lotic—

depositional

surfaces)

vertical rock

Clingers(on wet or dry

HEMIPTERA

carnivores

Piercers—

Beaches—marine lotic—

Predators

(piercers) (scavengers)

Climbers

(semiaquatic)

meadows)

semiaquatic)

Predators

(piercers)

Climbers

(at shore,

Lentic (marshy

Micracanthia

(10)

Arizona, Idaho

Widespread

Widespread

boreal zone)

Arctic

(widespread in

Predators

(piercers)

Climbers

Lampracanthia

(semiaquatic)

California, Southwest

Predators

(piercers) (scavengers)

Climbers

Canada

Alaska,

Predators

(at shore, semiaquatic)

semiaquatic)

zone)

northern

Canada

Alaska,

Predators

(piercers)

larvae)

Lentic (including

(at shore,

margins (tundra

Predators

(piercers) (scavengers, especially Diptera

Distribution

alkaline marshes); lotic—margins

Climbers

semiaquatic)

Climbers (at shore,

Lentic and lotic—

hydrophytes (emergent zone)

Lentic—vascular

freshwater

beach zone—

depositional;

lotic—

(at shore,

hydrophytes (emergent zone); semiaquatic)

Generally climbers

Generally lentic—

Habit

vascular

Habitat

North American

meadows)

crassicornis

trybomi

Species

Trophic Relationships

Lentic (marshy

Isocytus(7)

Chlloxanthus(2)

Calacanthia

Genus

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic

Order

of species in parentheses)

Taxa (number

Table 17A

> )}3J

SE

UM

M

10.0

NW

Ecological

4755

3995, 4754, 4755

3995, 4755

3995, 4755

3995, 4754, 4755

4754, 4755

4755

3995, 4599, 4754, 4755, 5274, 6103

253,536,565, 2776, 2786,

References**

(continued)

MA*

Tolerance Values

Gelastocoris(2)

freshwater

beaches—

(water's edge);

Lentic—littoral

freshwater

beaches—

depositional;

lotic—

hydrophytes (emergent zone);

vascular

Lotic—margins

(also salt marshes)

lotic—shorelines

Sprawlers (jumpers)

Generally sprawlers at shore, (semiaquatic)

Climbers (at shore, semiaquatic)

Climbers (at shore, semiaquatic)

meadows)

Lentic—littoral;

Climbers

(semiaquatic)

Lentic (marshy

(marshy meadows)

Climbers

(semiaquatic)

Beach zone—

Habit

freshwater: ientic

Habitat

-Toad Bugs

Species

Generally lentic—

Teloleuca (2)

Saldula (26)

Saldoida (3)

Salda (8)

Genus

Gelastocoridae (7)

Family

Continued

**Emphasis on trophic relationships

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic

Order

of species In parentheses)

Taxa (number

Table 17A

(piercers)

Predators

Generally predators (piercers)

(piercers) (scavengers)

Predators

(piercers) (scavengers)

Predators

(piercers) (scavengers)

Predators

(piercers) (scavengers)

Predators

Trophic Relationships

Widespread

mountains

Arctic, midlatitude

Widespread

Kansas

Michigan,

Texas,

East, South,

Widespread

Distribution

American

North SE

UM

M

10.0

NW

MA*

Tolerance Values

Ecological

565, 1418, 2778, 3653, 3995, 4538,4918, 5061, 6103, 749

536, 2776, 2786, 3179, 3995, 4599, 5274, 6103, 6423

3995, 4755

5738, 5751, 2235, 4958, 6103, 6560

4755, 5061,

3995, 4754,

2776,3537,

4754, 4755

3995, 4754, 4755, 6560

References**

) } ) )) ) ) ) ) ) ) > ) ) ) ) > ) ) ) ) ) V) ) )

a\ o\

iJl

Ul ON

Velvety Shore Bugs

Ochteridae (6)-

Family

Continued

Ochterus(6)

{=Mononyx)

Nerthra (5)

Genus

Species

Lentic—^vascular

surfaces faces)

hydrophytes; lotic—margins and seeps on rock

**Emphasis on trophic relationships

North

North to

Nebraska and

HEMIPTERA

South and

Great Lakes

Southwest,

vertical rock)

(piercers)

Widespread, particularly

Southeast

Predators

Southwest,

Predators

American Distribution

(piercers)

Trophic Relationships

semiaquatic); clingers(on seeping

Climbers (at shore,

under logs, etc.)

water's edge and terrestrial

beaches—

freshwater and

"Burrowers"

(in mud at

Lentic—littoral

Habit

(water's edge);

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic

Order

of species in parentheses)

Taxa (number

Table 17A

SE

UM

Wl NW

MA*

Tolerance Values

562, 565, 3995, 3685, 5274, 5311, 6103

3995, 4754,6103

Ecological References**

MEGALOPTERA AND AQUATIC NEUROPTERA David E. Bowles

Atilano Contreras-Ramos

Missouri State University, Springfield

Institute de Biologia, Universidad Nacional Autonoma de Mexico, Mexico City

INTRODUCTION

The Megaloptera (alderflies, dobsonflies, fishflies, hellgrammites) and aquatic Neuroptera (spongillaflies) constitute a small worldwide fauna of probably less than 400 species, representing three families (Sialidae, Corydalidae, and Sisyridae). This group of holometabolous aquatic insects contains some of the largest and most spectacular species. The aquatic larvae are predaceous and inhabit both lotic and lentic environments in tropical and temperate climates; however, all eggs, pupae, and adults are terrestrial. Large numbers of adults are seldom seen in nature because they are short-lived, secretive, and many species are nocturnal. Larval sialids are usually abundant in streams, rivers, or lakes where the substrate is soft and detritus

is abundant. Larvae usually burrow into the substrate and feed nonselectively on small animals,such as insect larvae, annelids,crustaceans,and mollusks,in the hab

itat. Sialids pass through as many as 10 instars during a 1- to 2-year life cycle. Prior to pupation, larvae leave the stream, river, or lake and pupate in an unlined chamber dug 1-10 cm deep in shoreline soil and litter. Adults (alderflies) usually emerge from late spring to early summer and are active during warm midday hours. Flight is brief and infrequent, and most individ uals stay in the same general area where the larvae occur. Apparently,the adults do not feed. Eggs are laid in masses primarily on leaves or branches overhanging the aquatic habitat, on large rocks overhanging or projecting from the water, or on bridge abutments. Larval corydalids (sometimes called hellgram mites) occur in a wide variety of habitats including spring seeps, streams, rivers, lakes, ponds, swamps, temporarily dry streambeds and even tree holes.

The life cycle is 1-5 years long with the larvae passing through 10-12 instars. As with sialids, corydalid lar vae feed on a wide variety of small aquatic inverte brates. Pupation occurs mostly in chambers in the soil adjacent to the larval habitat. However, some species pupate in dry streambeds, and others prefer soft, rot ting shoreline logs or stumps. Adults (dobsonflies, fishflies) emerge from late spring to midsummer. Most species of adult corydalids are nocturnal and may fly considerable distances; many are attracted to lights. Diurnal species are usually found resting or flying near the larval habitats. Oviposition habits are similar to those of adult sialids. Observational data in

captivity show some adults are capable of actively licking or ingesting soft fruit. Larvae of the sisyrids are usually found associ ated with freshwater sponges and bryozoans. They occur on the surface or in the cavities of the host, and

pierce the sponge or bryozoan cells sucking the fluids with their elongated mouthparts. Larvae pass through three instars, and some species have several genera tions each year. Just before pupation,the larvae leave the water,climb onto shoreline plants or other objects, and spin a silk cocoon in which to pupate. Sites cho sen for pupation are frequently somewhat secluded and usually near shore, but larvae may migrate inland up to 20 m before pupating. The cocoon is usually double walled, i.e., composed of an inner, closemeshed enclosure and an additional, more loosely

constructed outer envelope. No feces are voided by the larvae prior to pupation; this is typical of many neuropterans but unique among aquatic insects. Adults (spongillaflies) appear to be primarily noctur nal. Some CUmacia are diurnal and may be seen for aging on streamside flowers.

569

570

Chapter 18

Megaloptera and Aquatic Neuroptera

EXTERNAL MORPHOLOGY

Megalopteran eggs are quite distinct and can be separated by the size and appearance of the egg mass, and the size, color, sculpturing, and shape of the micropylar process of individual eggs. The eggburster, left with the hatched egg by a newly emerged larva, is also diagnostic. Identification of larval Megaloptera is based primarily on the number of abdominal filaments, the presence or absence of ven tral abdominal gill tufts, and the appearance and loca tion ofthe eighth abdominal spiracles. Setae and color patterns are also used, particularly for separating spe cies, although intraspecific variation may exist. Pupal identifications of sialids and corydalids can be made based on size and color patterns. Familial and generic identification of adult Megaloptera relies mainly on wing venation; species identification is based primar ily on male and female genitalic characters. Eggs of the two genera of aquatic Neuroptera have not been studied adequately to provide diagnos tic characters. Larval sisyrids are best separated at the generic level by the presence or absence and the loca tion of certain setae, and by the presence or absence ofspines associated with setae. The labium is useful in separating pupal sisyrids, and there are possibly some differences in the appearance of the cocoons of the two genera. Adult aquatic Neuroptera are identified mainly by wing venation and genitalia.

Megaloptera (Slalidae, Corydalidae) Eggs: Sialid eggs are laid in even rows of about 200-900 eggs, in compact, more or less quadrangular masses with the eggs vertical (Fig. 18.1) or horizontal (Fig. 18.2) to the substrate. Each egg is cylindrical, about 0.2 by 0.6 mm in size, with rounded ends; the outer surface is partially or completely covered with small, very short, shield-shaped projections. The micropylar process is cylindrical or slightly fusiform (Fig. 18.16). The egg-burster is V-shaped and sharply toothed (Fig. 18.3). Corydalid egg masses of 300-3,000 eggs are com pact,rounded to quadrangular in shape,and have 1-5 layers (Figs. 18.18, 18.19, and 18.20); sometimes the eggs have a white or brown protective covering (Fig. 18.18). Individual eggs, in general, appear similar to sialid eggs; sometimes the eggs are covered with shield-shaped processes (Fig. 18.17), but usually they are relatively smooth. The size ofeach egg is about 0.5 by 1.5 mm. The position of the eggs relative to the substrate is as in the Sialidae, and the micropylar pro cess is apically enlarged. Egg-bursters are elongate, rounded, or ridged apically, and toothed (Fig. 18.4).

Larvae: Terminal instar sialids (Figs. 10.106 and 18.7) reach a maximum length of approximately 25 mm, including the unsegmented, median caudal filament. Mouthparts consist of a labrum, two welldeveloped mandibles (for grasping and engulfing the prey), two maxillae, and a labium. Antennae are four-segmented. The quadrate head is patterned, as is the 10-segmented abdomen, which ranges in color from purplish or reddish brown to yellow. Thoracic legs have two claws. Abdominal segments 1-7 bear four- or five-segmented lateral filaments (Fig. 18.7). Corydalid larvae (Figs. 18.21, 18.26-18.27) are larger than sialids, reaching 30-65 mm or more in length when full grown. Mouthparts are similar to those of sialid larvae but the mandibles are usually more robust. Antennae are four- or five-segmented. Thoracic legs have two claws. The head and thorax may be of a uni form color or patterned. Abdominal segments 1-8 bear two-segmented (a short, basal segment and a long, distal segment) lateral filaments (Figs. 18.21, 18.2618.27) and the abdomen terminates in a pair of anal prolegs. Each proleg bears paired claws and a dorsal filament. The last pair of spiracles on the abdomen (segment 8)are sometimes modified with regard to size and location (Figs. 18.28-18.35). Pupae: Both sialid (Fig. 18.8) and corydalid (Fig. 18.9) pupae are exarate (appendages free, not fastened to body), decticous (mandibles articulated, functional), and range in length from 10 to 12 mm and from 30 to 60 mm,respectively. Adults: Adults of Sialidae are approximately 10-15 mm in length. Their bodies are black, brown,or yellowish orange with similarly colored wings (Figs. 10.110 and 18.12). The head lacks ocelli, and the fourth tarsal segment is dilated (Fig. 18.15). Corydalid adults are 40-75 mm long, with black, brown, or gray bodies; many species have pale smoky wings mottled with brown (Figs. 10.109 and 18.13). Some species (Nigronia spp.) have darker, almost black, wings with white markings (Figs. 10.108 and 18.36). The head has three ocelli and males of some species {Corydalus spp.) have very long mandibles (Figs. 10.109, 18.44-18.45), which may vary in length with body size. All tarsal segments are simple (Fig. 18.14).

Aquatic Neuroptera (Sisyrldae) Eggs: Masses are of 2-5, or occasionally as many

as 20, oval, whitish to yellowish eggs covered with a web of white silk. Each egg is about 0.1 by 0.3 mm in size with a short micropylar process(Fig. 18.47). The egg-burster is elongate (Fig. 18.46).

Chapter 18 Megaloptera and Aquatic Neuroptera

Larvae: Terminal instars (Fig. 18.48) are small (4-8 mm in length), stout, and with conspicuous setae (Fig. 10.107). Body color varies from yellowish brown to dark green. Mouthparts are modified into elon gate, unsegmented stylets (usually separated in pre served specimens). Antennae are relatively long and legs are slender and bear a single claw. Second and third instars bear two- or three-segmented,transparent

571

ventral gills, which are folded medially and posteri orly on abdominal segments 1-7. Pupae: All species are exarate (exposed) and housed in hemispherical, usually double-walled, silken cocoons (Fig. 18.52). Adults: Spongillaflies lack ocelli, have brown bodies with brown or mottled wings(Figs. 10.111 and 18.57), and are 6-8 mm in length.

KEYS TO THE FAMILIES AND GENERA OF MEGALOPTERA

1

Eggs in masses of approximately 15 mm diameter (Figs. 18.1-18.2); egg-burster V-shaped (Fig. 18.3). Larvae with 7 pairs of 4- to 5-segmented lateral filaments on abdominal segments 1-7 and a single long caudal filament(Fig. 18.7); anal prolegs absent; 4th tarsomere enlarged and wider than remaining segments; ocelli absent; 25 mm or less when full grown. Pupae 10-12+ mm (Fig. 18.8). Adults less than 25 mm in length (many 10-15 mm)(Fig. 18.12); ocelli absent; 4th tarsal segment dilated (Fig. 18.15) SIALIDAE

1'.

Eggi in masses of 20+mm diameter (Figs. 18.18, 18.19-18.20); egg-bursters elongate, apically rounded, or ridge-like and toothed (Fig. 18.4). Larvae with 8 pairs of 2-segmented lateral filaments on abdominal segments 1-8, and a pair of 1-segmented filaments on abdominal segment 10(Fig. 18.21); apex of abdomen with 2 anal prolegs, each bearing a pair of claws; 4th tarsomere cylindrical and about the same width as other segments; three ocelli; 30-65 mm when full grown. Pupae greater than 30 mm in length (Figs. 18.9-18.11). Adults over 25 mm in length (Figs. 18.13, 18.37); ocelli present; 4th tarsal segment simple (Fig. 18.14) CORYDALIDAE

Siaildae Larvae

1. 1'.

Mandibles with three subapical teeth; basal tooth usually about one-half the size of the other two teeth (Fig. 18.5) Mandibles with two subapical teeth (Fig. 18.6)

Protosialis Weele Sialis Latreille

Corydalidae 1. 1'. 2(1'). 2'. 3(2).

Egg mass 3-layered with a thick, white chalky covering (Fig. 18.18) Egg mass not as above Egg mass single layered, may have a thin coating Egg mass with more than one layer Egg chorion with peltate (shield-shaped) processes on dorsum (Fig. 18.17)

3'.

Egg chorion smooth; a thin coating may cover egg mass

Corydalus Latreille 2 3 4 Chauliodes Latreille

Neohermes Banks, Protochauliodes Weele

4(2').

Egg mass 3-5 layered; western United States and Canada (Fig. 18.19)

5

4'. 5(4).

Egg mass with up to 50 eggs in a second layer; eastern and central United States ... .Nigronia Banks Egg approximately 2.0 mm long Dysmicohermes Munroe

5'.

Egg approximately 1.0 mm long

Orohermes Evans

572

Chapter 18

Megaloptera and Aquatic Neuroptera

Figure 18.3

Figure 18.2

Figure 18.1

Figure 18.4

Figure 18.5

Figure 18.6

labrum

mandible antenna

lateral filament

caudal filament

Figure 18.7

Figure 18.1 Eggs of Sialis rotunda Ross (Sialidae). Figure 18.2 Eggs of Sialis hamata Ross (Sialidae). Figure 18.3 Lateral view of V-shaped egg-burster of Sialis sp.(Sialidae)(length 90 pm). Figure 18.4 Lateral view of corydalid egg-bursters (Corydalidae); A, Orohermes crepusculus (Chandler) (length, 115 pm); B, Corydalus sp.(length, 160 mm). Figure 18.5 Mandible of Protosiaiis larva.

Figure 18.8

Figure 18.9

Figure 18.6 Mandible of Sialis larva. Figure 18.7 Dorsal view of larva of Sialis rotunda Ross (Sialidae). Figure 18.8 Lateral view of pupa of Sialis cornuta Ross (Sialidae)(after Leischner and Pritchard 1973). Figure 18.9 Lateral view of Neohermes sp. pupa (Corydalidae); length 30+ mm.

Chapter 18 Megaloptera and Aquatic Neuroptera

Figure 18.12

Figure 18.11

Figure 18.10

573

Figure 18.14

\ segment IV

Figure 18.13

segment IV

Figure 18.15 chalky covering.

3rd layer

micropylar process

/ micropylar process 2nd layer—

/

)stlayer

Figure 18.17 Figure 18.16

Figure 18.10 Lateral view of Corydalus cornutus pupa (Corydalldae). Figure 18.11 Ventral view of Corydalus cornutus pupa (Corydalldae). Figure 18.12 Adult of Sialis californica Ross (Slalldae). Figure 18.13 Adult of Orohermes crepusculus (Cfiandler)(Corydalldae).

Rgure 18.14 Distal part of tibia and tfie tarsus of Orohermes crepusculus (Chandler) with a simple 4th tarsal segment (Corydalldae).

Figure 18.18

Figure 18.15 Distal part of tibia and the tarsus of Sialis sp. with a dilated 4th tarsal segment (Slalldae). Figure 18.16 Egg of Sialis hasta Ross (Slalldae) (length, 0.6 mm). Figure 18.17 Egg of Chauliodes pectinicornis (L.) (Corydalldae)(length 1.0 mm). Figure 18.18 Section of Corydalus egg mass (Corydalldae); covering partially removed.

574

Chapter 18 Megaloptera and Aquatic Neuroptera

Larvae

1.

Abdominal segments 1-7 with ventral gill tufts at base of lateral filaments

r.

Abdominal ventral gill tufts absent

2(1').

Last pair of abdominal spiracles(segment 8)at the apex of 2 long dorsal

(Figs. 10.105 and 18.21)

Corydalus Latreille 2

respiratory tubes extending beyond prolegs (Fig. 18.22)

Chauliodes Latreille

2'.

Last pair of abdominal spiracles not at apex of long respiratory tubes

3

3(2').

Larval head not conspicuously patterned

4

3'. 4(3).

Larval head with a conspicuous pattern (Figs. 18.26-18.27) 6 West coast of United States and Canada; last pair of abdominal spiracles either dorsal, large, and raised on short tubes (Figs. 18.23, 18.34-18.35), or lateral and about same size as elsewhere on abdomen (Figs. 18.24, 18.32) 5 East and central United States and Canada; last pair of abdominal spiracles dorsal, similar in size to other abdominal spiracles, and borne at apex of short respiratory tubes (Figs. 10.104, 18.25, 18.29, and 18.31) NigroniaBmk^ Last pair of spiracles dorsal, large, and raised on short tubes (Figs. 18.23, 18.34-18.35) Dysmicohermes Munroe Last pair of spiracles lateral, similar in size to other abdominal spiracles and

4'.

5(4). 5'. 6(3').

sessile (Figs. 18.24 and 18.32) Spiracles on abdominal segment 8 distinct, associated with raised areas of the integument(Figs. 18.30 and 18.33); eastern and western United States and

Orohermes Evans

Canada

6'.

Neohermes Banks

Spiracles of abdominal segment 8 less conspicuous, integument not raised around spiracle (Fig. 18.28); west coast of United States and Canada

Protochauliodes Weele

Adults Sialidae

1.

Legs entirely black or brown or with tibiae slightly lighter in color than femora; head primarily yellow with some black markings; Rs distally 2-branched in both fore- and hind wings

r.

Protosialis Weele

Legs with femora reddish brown and tibiae the same color or blackish; head primarily black but with yellow or orange markings (Fig. 10.110); Rs distally 3 or 4-branched in both fore- and hind wings

Sialis Latreille

Corydalidae

1.

r.

2(1'). 2'.

Forewing with white spots in many cells (Fig. 10.109); 20 veins or more reaching wing margin posteriad of R,; M vein with 3 branches reaching wing margin (Fig. 18.36-18.37) Corydalus Latreille Forewing lacking white spots(Fig. 18.13), or white spots less widely distributed (Fig. 18.41); less than 20 veins reaching wing margin posteriad of Rj; M vein with 2 branches reaching wing margin (Figs. 18.38-18.41) 2 Posterior branch of Rs forked in both pairs of wings (Fig. 18.38) 3 Posterior branch of Rs simple in both pairs of wings (Fig. 18.39) 4

3(2).

Hind wing with posterior branch of M forked (Fig. 18.38)

3'.

Hind wing with posterior branch of M simple

4(2'). 4'.

M vein of hind wing with 3 branches reaching wing margin (Fig. 18.39) M vein of hind wing with 2 branches reaching wing margin (Fig. 18.40)

Dysmicohermes Munroe Orohermes Evans

5 6

Chapter 18 Megaloptera and Aquatic Neuroptera

575

mandible

4th layer

maxi a

antennae

'5th layer

3rd layer

ventral gill tuft lateral filament

^,'2nd layer spiracle

Figure 18.20 -♦—1st layer

Figure 18.21

Figure 18.19

dorsal proleg filament

paired claws

segment VIII lateral filament

respiratory tube

segment VIII - spiracle

spiracle

Figure 18.22

segment VIII spiracle

Figure 18.23

segment VIII

. spiracle

Figure 18.24

Figure 18.25 Figure 18.27

Figure 18.26

Figure 18.19 Section of a mutilayered egg mass of Orohermes crepusculus (Chandler) (Corydalidae). Figure 18.20 Section of a Neohermes sp. egg mass (Corydalidae). Figure 18.21 Dorsal view of larva of Corydalus sp. (Corydalidae). Figure 18.22 Dorsal view of caudal segments, respiratory tubes and spiracles of segment VIII of Chauliodes sp. larva (Corydalidae). Figure 18.23 Dorsal view of caudal segments and spiracles of segment VIM of Dysmicohermes ingens Chandler larva (Corydalidae). "s

Figure 18.24 Dorsal view of caudal segments and spiracles of segment VIII of Orohermes crepusculus (Chandler) larva (Corydalidae). Figure 18.25 Dorsal view of caudal segments and spiracles of segment VIII of NIgronia serricornis (Say) larva (Corydalidae). Figure 18.26 Dorsal view of larva of Neohermes filicornis (Banks) (Corydalidae). Figure 18.27 Dorsal view of larva of Protochauliodes spencer! Munroe (Corydalidae).

576

Chapter 18 Megaloptera and Aquatic Neuroptera

/

f//

Figure 18.30

Figure 18.28

Figure 18.29

L

fr.

.I ,

^

Figure 18.33

Figure 18.31

Figure 18.32

i. Figure 18.34

Figure 18.28 Dorsal view of left spiracle on segment VIM of Protochauliodes spenceri Munroe larva (Corydalidae). Figure 18.29 Dorsal view of left spiracle on segment VIII of Nigronia serricornis (Say) larva (Corydalidae). Figure 18.30 Dorsal view of left spiracle on segment VIII of Neohermes concolor (Davis) larva (Corydalidae). Figure 18.31 Dorsal view of left spiracle on segment VIII of Nigronia fasciatus (Walker) larva (Corydalidae). Figure 18.32 Dorsal view of left spiracle on segment VIII of Orohermes crepusculus (Ctiandler) larva (Corydalidae).

Figure 18.35

Figure 18.33 Dorsal view of left spiracle on segment VIII of Neohermes fiiicornis (Banks) larva (Corydalidae). Figure 18.34 Dorsal view of left spiracle on segment VIII of Dysmicohermes ingens Chandler larva (Corydalidae). Figure 18.35 Dorsal view of left spiracle on segment VIII of Dysmicohermes disjunctus (Walker) larva (Corydalidae).

Chapter 18 Megaloptera and Aquatic Neuroptera

Figure 18.36

3-branched

577

Figure 18.37

2-branched

2-branched

2-branched 2-branched

Figure 18.38

—— R3 R

unbranched 2-branched

* unbranched

M

Rs 3-branched

Figure 18.39

unbranched

2-branched

T

i

l

l n

unbranched

2-branched

Figure 18.40

Figure 18.36 Forewing of Corydalus sp.(Corydalidae). Figure 18.37 Adult male, Corydalus sp.(Corydalidae). Figure 18.38 Wings of Dysmicohermes disjunctus (Walker)(Corydalidae).

Figure 18.39 Wings of Neohermes sp. (Corydalidae). Figure 18.40 Wings of Chauliodes sp.(Corydalidae).

578

5(4).

5'.

6(4'). 6'.

Chapter 18 Megaloptera and Aquatic Neuroptera

Crossvein present between R3 and R4 in forewing (Fig. 18.39); antennae of male elongate, moniliform (bead-like), setigerous (bearing setae)(Fig. 18.42); apical papilla of gonapophysis lateralis absent in female (compare with Fig. 18.43) Neohermes Banks Crossvein absent between R3 and R4 in forewing; antennae filiform (thread-like): apical papilla of gonapophysis lateralis present in female (Fig. 18.43) Protochauliodes Weele Wings dark with white spots and patches (Figs. 10.108 and 18.41) Nigronia Banks Wings pale gray-brown, mottled (Fig. 18.40)

Chauliodes Latreille

KEYS TO THE GENERA OF AQUATIC NEUROPTERA

Sisyridae Larvae (after Bowles [2006])

1.

r.

Pair of dorsal setae present on abdominal segment 8(Fig. 18.48); ventral pair of medial setae on abdominal segment 8 raised on tubercles and only slightly closer together than those on segment 9(Fig. 18.50); small acute spines at bases of thoracic setae (Fig. 18.49)(not present in Climacia californica) Climacia McLachlan Pair of dorsal setae absent on abdominal segment 8; pair of ventral medial setae on abdominal segment 8 sessile and distinctly closer together than those on segment 9(Fig. 18.51); small acute spines at bases of thoracic setae absent Sisyra Burmeister

Pupae 1. Segments of labial palp similar (Fig. 18.54) 1'. Last palpal segment of labium greatly enlarged and triangular (Fig. 18.53)

Climacia McLachlan

Sisyra Burmeister

Adults

1.

Rs offorewing with one fork before pterostigmata (Fig. 18.55), forewing with brown markings (Fig. 10.111); last segment of labial palp similar in size and shape to other segments(Fig. 18.54)

r.

Rs offorewing with more than one fork before pterostigmata (Fig. 18.56), forewing uniformly brown; last segment of labial palp greatly enlarged and triangular in shape (Fig. 18.53)

Climacia McLachlan

Sisyra Burmeister

Chapter 18 Megaloptera and Aquatic Neuroptera

579

Figure 18.42

2-branched

Figure 18.41

apical papilla

gonapophysis

Figure 18.45

iateralis

Figure 18.43

Figure 18.44

,micropylar process

Figure 18.46

Figure 18.41 Forewing of Nigronia sp,(Corydaliciae). Figure 18.42 Basal part of antenna of male Neohermes sp.(Corydalidae). Figure 18.43 Lateral view of female genitalla of Protochauliodes spenceri Munroe (Corydalidae). Figure 18.44 Dorsal view of head and part of prothorax of male Corydalus sp.(eastern North America)(Corydalidae).

Figure 18.47

Figure 18.45 Dorsal view of head and prothorax of male Corydalus sp.(western North America) (Corydalidae). Figure 18.46 Lateral view of egg-burster of Climacia sp.(Sisyridae)(after Brown 1952). Figure 18.47 Egg of Climacia sp.(Sisyridae)(after Brown 1952).

580

Chapter 18

Megaloptera and Aquatic Neuroptera

styletlike mandible antenna

— seta

claw

basal

spine. inner cocoon —

- outer cocoon

segment VIII

Figure 18.49 Figure 18.48 segment VIII segment VIII

— segment IX i~segment IX

Figure 18.52

Figure 18.53

Figure 18.54 Figure 18.50

pterostigmata Sc

Figure 18.51 pterostigmata

n

Figure 18.55

Figure 18.56

Figure 18.57

Figure 18.48 Dorsal view of larva of Climacia sp. (Sisyrldae)(after Brown 1952). Figure 18.49 Right dorsal plate of pronotum of Climacia sp.(Sisyrldae)(after Parfin and Gurney 1956). Figure 18.50 Ventral view of caudal abdominal segments of Climacia areolaris (Hagen)(after Parfin and Gurney 1956). Figure 18.51 Ventral view of caudal abdominal segments of Sisyra vicaria (Walker)(after Parfin and Gurney 1956). Figure 18.52 Lateral view of pupa and cocoons of Sisyra sp. (Sisyrldae).

Figure 18.53 Labial palp of Sisyra sp.(Sisyrldae) (after Chandler 1956a). Figure 18.54 Labial palp of Climacia sp.(Sisyrldae) (after Chandler 1956a). Figure 18.55 Forewing of Climacia sp.(Sisyrldae) (after Parfin and Gurney 1956). Figure 18.56 Forewing of Sisyra sp.(Sisyrldae)(after Parfin and Gurney 1956). Figure 18.57 Female Climacia sp.(Sisyrldae)(after Brown 1952).

Chapter 18 Megaloptera and Aquatic Neuroptera

ADDITIONAL TAXONOMIC REFERENCES

Pacific Coastal Region: Evans(1972). South Carolina: Brigham et al.(1982).

General Chandler (1956a,b); Gurney and Parfin (1959); Pennak (1978); McCafferty (1981); Evans and Neunzig (1984); Peckarsky et al.(1990); New and Theischinger (1993); Penny et al. (1997); Arnett (2000); Cover and Resh (2008)

South Dakota: Johnson et al.(1997). Texas: Locklin (2008), Lowery et al.(2008).

581

Washington: Whaley et al. (2003). West Virginia: Watkins et al.(1975); Tarter (1976, 1988); Tarter et al.(1976); Tarter et al.(2010); Tarter et al. (2013).

Regional faunas Arkansas: Bowles (1989); Bowles and Sites (2015). California: Chandler (1954, 1956a,b): Whiting (1991c). Canada: Kevan (1979); Roy and Hare (1998). Colorado: Hermann and Davis(1991). Eastern United States: Tarter et al. (1976), Idaho: Biggam (1998). Illinois: Bowles and Sites (2015) Kansas: Huggins(1980); Roble (1984); Engel (2004). Kentucky: Call (1982); Tarter et al.(2006); Tarter et al.(2009). Louisiana: Locklin (2007). Maryland: Flint (2008). Minnesota: Parfin (1952). Mississippi: Poirrier and Holzenthal (1980); Lago (1981); Stark and Lago (1980, 1983). Missouri: Bowles and Sites(2015). New York: Needham and Betten (1901a). North Carolina: Cuyler (1956); Brigham et al.(1982). Oklahoma: Arnold and Drew (1987); Bowles (1989); Bowles and Sites(2015).

Taxonomic treatments at the family and generic levels Corydalidae: Munroe (1951b, 1953); Cuyler (1958); Hazard (1960); Flint(1965); Neunzig (1966); Baker and Neunzig (1968); Glorioso (1981); Evans(1984); Contreras-Ramos (1990, 1998, 2011); Neunzig and Baker (1991); Bowles and Mathis(1992); Contreras-Ramos and Harris(1998); Liu and Yang 2006; Bowles et al.(2007); Liu et al. (2015); Liu and Winterton.(2016).

Sialidae: Davis (1903); Ross (1937); Cuyler (1956); Canterbury (1978); Canterbury and Neff(1980); Neunzig and Baker (1991); Whiting (1991a, 1991b, 1994); Liu et al.(2015). Sisyridae: Parfin and Gurney (1956); Poirrier and Arceneaux (1972); Pupedis (1980); Tauber (1991); Flint(1998); Bowles(2006).

Family Genus

Corydalinae (4)

- Dobsonflies, Fishflies, Hellgrammites

Corydalidae(23)

Alderflies

Sialidae (24)-

Protosialis(2)

Sialis(22)

Species

Generally clingers— climbers

littoral)

(engulfers)

Predators—

gatherer)

Generally lotic— erosional (lentic—

a collector—

lentic— erosional

species reported to be

clingers

(sediments)

Predators— (engulfers)(one

Burrowers—

climbers—

Lotic—erosional

gatherer)

and depositional (detritus. sediments);

a collector—

lentic— erosional

Burrowers— Predators— climbers— (engulfers)(one clingers species reported to be

Habit

North

Widespread

Widespread

7.4

4

4.9

0

4

4.9 4

4

4

NW MA*

Tolerance Values

7.4 4

Trophic American Relationships Distribution SE

(sediments)

Lotic—erosional and depositional (detritus, sediments);

Habitat

*SE = Southeast, DM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasls on trophic

FIshflles

Megaloptera - Alderflies, Dobsonflies and

Order

species in parentheses)

(number of

Taxa

Table 18A Summary of ecological and distributional data for the Megaloptera (alderflies, dobsonflies, hellgrammites) and the Aquatic Neuroptera (spongillaflies). (For definition of terms see Tables 6A-6C; table prepared by K. W. Cummins, R. W. Merritt, Fl. Fl. Neunzig , and E. D. Evans)

Ecological

262, 974, 1718, 2295, 4513, 1720, 4316, 4599, 6205, 2958, 574, 642, 1172, 5899

3482,3530, 4277, 4513,4599, 4822, 5134, 5861, 5903, 6032, 3777, 4423, 6287, 6721, 2584, 3313, 4715, 4694, 4715, 5177, 3379, 6009, 3554, 5893

242, 243, 718, 894, 974, 1737, 2069, 2097, 2295, 2582, 3039,

242, 243, 718, 894, 974, 1737, 2069, 2097, 2295, 2582, 3039, 3482,3530, 4277, 4513,4599, 4822, 5134, 5861, 5903, 6032, 3777, 4423, 6287, 6721, 2584, 3313,4715,4694, 4715,5177, 3379, 6009, 3564, 5893

1720, 4316, 642, 1172, 1669, 3555, 3565

974, 1103, 2295, 4599, 1720, 4316, 4585, 1874,3556

References**

) ) ) ) ) > ) ) ))) ) ) ) ) )) ) ) ) ) ) ) ) ) )

'Jl 00

c/i

Genus

Nigronia (2)

Neohermes(6)

Dysmicohermes (2)

ChauHades(2)

Species

MEGALOPTERA-NEUROPTERA

oe

Chauliodinae (19)

Family Predators—

4513, 5034, 5891, 5904, 5667,

262, 1299, 1369, 2484, 1464,

3553

t

262, 723, 974, 1718, 2295, 3198, 4513, 5034, 635,918, 1347, 5737, 1103, 1692, 3313, 3707, 3821,4469, 5091, 5437, 3151, 3708, 4608, 4694, 6142, 637, 1106, 3586, 5896, 5900,

(in crevices and under

bark)

bark and in crevices

of woody debris)

borrowers

Predators—

Clingers— climbers—

Lotic—erosional and

depositional (coarse sediments, detritus. especially under

(engulfers)

(engulfers)

Clingers— climbers

Lotic—erosional and

Predators—

(engulfers)

spring seeps (sediments, vascular hydrophytes, leaf detritus)

(sediments. detritus)

East, Central

West, East

(Continued)

262, 840, 1073, 1077, 1279, 3199, 4077, 4314, 4653, 1347, 5891, 5898, 5904, 6062, 846, 954, 2014, 4031, 2013, 2009, 4603, 4604, 4605, 4606, 4607, 3010, t

1103, 5080, 5901, 5904, 3554

1718, 1863,4576, 5515, 639,

climbers

2.7

2.4

and depositional

0

6

1758, 2347

5.8

5.6

Ecological References**

972, 1104, 1718, 4205,4206 West, Texas

East, Central

Widespread

NW MA*

borrowers Predators—

(engulfers)

Predators—

(engulfers)

M

Clingers—

detritus, logs)

North

American

Relationships Distribution SE UM

Trophic

Tolerance Values

Lotic—erosional

Clingers— climbers—

Lentic—littoral

(swimmers)

(sediments, detritus)

(sediments.

Clingers— climbers

Lotic—eroslonal

Habit

and deposltional

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic tUnpublished data, K. W. Cummins, Kellogg Biological Station

Order

i

Corydalus(4)

Continued

(number of species in parentheses)

Taxa

Table 18A

J

00 4^

'Jt

Sisyridae (6) Spongillaflies

Family

Predators—

depositional

Widespread

Sisyra (3)

»

6.5

0

NW MA*

Ecological

973, 1043,2248, 4513,4936, 5749, 5910

973, 2295, 3821, 4936, 6451, 1043, 5749, 5910, 5489, 6445

973, 2295,4439, 4513, 4514, 4599, 4738, 4740, 1720, 3821 4936, 4839, 5749, 5910, 633, 1172, 1874,3356

972, 1718, 3664,4205,4206

972, 1737, 1719

References**

^ ) ) ) ) ) ))) ) )) ) ) ) ) ) )

widespread)

(1 eastern, 1 northern, 1

Widespread (2 Central and Eastern, 1 Western)

West

West

Climada (3)

sponges); lentic— littoral (on alpine sponges)

(piercers of clingers— Spongilla, borrowers Ephydatia, (live in or on Bryozoa etc.) sponges)

Predators—

Generally—

erosional (on climbers—

Generally lotic—

habitats)

intermittent

(including

(detritus)

(engulfers)

Clingers— climbers

Lotic—erosional

(sediments. detritus); lotic—

Protochauliodes

(6)

Predators—

(engulfers)

Lotic—erosional

M

Tolerance Values

American Trophic Relationships Distribution SE UM

Clingers—

Habit climbers

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW : Northwest, MA = Mid-Atlantic **Emphasis on trophic

Neuroptera

Order Orohermes

Continued

parentheses)

(number of species in

Taxa

Table 18A

TRICHOPTERA John C. Morse

Andrew K. Rasmussen

Clemson University, Clemson, South Carolina,

Florida Agricultural and Mechanical University, Tallahassee, Florida

Ralph W. Holzenthai

Douglas C. Carrie

University ofMinnesota, St. Paul,

Royal Ontario Museum and University of

Minnesota

Toronto, Toronto, Ontario, Canada

Desiree R. Robertson

Field Museum of Natural History, Chicago, Illinois

INTRODUCTION

The Trichoptera, or caddisflies, one of the largest

groups of aquatic insects, are closely related to the Lepidoptera. They are holometabolous and, except for a few land-dwelling larvae that are secondarily adapted to life out of water (Anderson 1967; Flint 1958; Schuster 1997), are aquatic in the immature stages. Respiration by larvae and pupae is indepen dent ofthe surface and ofatmospheric oxygen. Adults of almost all species are active, winged insects, although females of at least one North American spe cies are wingless at some time (see Ross 1944, fig. 171; Wiggins 2004, fig. 50F). Nearly 1,500 species of cad disflies are now known in North America north ofthe

Rio Grande, and these are currently assigned to 155

genera in 27 families (Rasmussen and Morse 2018). The taxonomic richness is a consequence of the broad ecological diversity ofthe order(Wiggins and Mackay 1978). Caddisflies occur in most types of freshwater habitats: spring streams and seepage areas, rivers, lakes, marshes, and temporary pools, and they have been particularly successful in subdividing resources within these habitats. General summaries of informa

tion on the biology of Trichoptera are available in several references: Betten (1934), Balduf (1939), Lepneva (1964), Malicky (1973), Ross (1944), Wig gins(1977, 1996, 2004), Mackay and Wiggins (1979), Holzenthai et al.(2015), and Table 19A. All North American families are represented in cool, lotic waters and they also have been successful

to varying degrees in exploiting freshwater habitats that are larger, warmer, and increasingly lentic (Wiggins 2004,fig. 5). Most larvae eat algae and plant materials in one form or another, especially decaying vascular plant tissue and associated microorganisms, but evidently living vascular plant tissue is less often ingested; some larvae are mainly predaceous. The food ofearly instars in all groups is largely fine organic particles.

Caddisflies have penetrated freshwater food webs particularly effectively because the construction behavior of their larvae has enabled them to subdi

vide feeding niches by building portable cases, fixed retreats, and filter nets using silk from their salivary glands. In this manner, larvae have been able to infil trate feeding niches not fully exploited by other aquatic insects that lack silk and behavior for build ing various devices, which enhance feeding efficiency, survival, and reproduction (Wiggins 2004). The 52 or so extant families of caddisflies recog

nized in the world were assigned to two suborders by Holzenthai et al. (2011), both represented in North America. The suborders Annulipalpia and Integripalpia are well established as monophyletic evolutionary lineages, including (Kjer et al. 2016) or excluding (Malm et al. 2013) a grade of four basal families that evolved early in the history of the order. The North American families in Table 19A are listed alphabeti

cally in the subordinal classification of Holzenthai etal.(2011).

585

586

Chapter 19 Trichoptera

Suborder Annulipalpia

Larvae of the Philopotamidae (Figs. 10.166, 19.32, and 19.71) live in elongate, fine-meshed nets in

Dipseudopsidae, Ecnomidae, Hydropsychidae, Philopotamidae, Polycentropodidae, Psychomyiidae,

reduced currents on the underside of rocks, where

Xiphocentronidae. These are the retreat-making caddisflies in which

they filter particles smaller than those filtered by other Trichoptera (Wallace and Malas 1976a); the special

the ecological strategy differs markedly from that of the case-makers because, instead of moving about to find food,these larvae rely on stream currents or wave action along lake shores to bring food materials to their fixed retreats. Ten families are assigned to the

ized membranous labrum serves to clear accumulated

Annulipalpia in the world fauna,and all but three,the Kambaitipsychidae, Pseudoneureclipsidae, and Stenopsychidae, occur in North America.

Larval Hydropsychidae (Figs. 10.165, 19.34, and 19.35) construct fixed retreats of organic and mineral fragments with a silken sieve net placed adjacent to the anterior entrance to filter particles from the cur rent. Mesh size of the filter nets differs: Larvae in the

Arctopsychinae, living in cold upstream sites with strong currents, spin the largest meshes and feed mainly on insect larvae; larvae in the Macronematinae, living in downstream sites with slow currents, spin the smallest mesh size and filter small particles; and those in the Diplectroninae, Hydropsychinae, and Smicrideinae, occupying sites intermediate between these two, spin sieve nets with meshes in a range ofintermediate sizes(Wallace 1975a,b; Wallace and Malas 1976b; Hauer and Stanford 1981). Larvae in Hydropsyche produce sound by rubbing the femur across ridges on the underside of the head (Jansson and Vuoristo 1979; Wiggins 2004). Evidently the sound is a defensive behavior of larvae in protecting their retreats against other hydropsychids, but whether this protective role extends generally to pred ators is not known.

Larvae in the Polycentropodidae (Figs. 10.167, 19.37,19.65-19.67)construct shelters of several types. Larvae in some genera such as Neureclipsis construct a funnel-shaped filter net of silk in slow currents and rest in the narrowed base; a single net may be 12 cm or so in length. In predaceous genera, such as Nyctiophylax, larvae make a flattened tube of silk in depres sions on rocks or logs; concealed within the tube, a larva darts out to capture prey that trip the silk threads emanating from each opening. In the family Dipseudopsidae,larvae ofthe genus Phylocenlropus fashion branching tubes of silk and sand in beds of loose sediments (Figs. 10.163, 19.33), with the ends of the tubes protruding above the sedi ment; water with food particles in suspension enters the upstream tube, passes through a filter of silk threads which retains the particles, and out again through the downstream tube (Wallace et al. 1976).

particles from the net. Although larvae ofthe Psychomyiidae(Figs. 10.168 and 19.64), Ecnomidae (Figs. 10.164, 19.70), and Xiphocentronidae (Figs. 10.169, 19.69, 19.70) live mainly in running waters, they do not filter food from the current but graze on periphyton and the fine particulate matter deposited on rocks by the water current. Constructed of fine sand and organic material over a silken lining (e.g.. Fig. 19.36), their tubular retreats are fastened to rocks and logs. Larvae of at least some psychomyiids purposefully grow gardens of periphy ton in their silken retreats(Hasselrot et al. 1996; Ings et al. 2010, 2012). Larvae of the Xiphocentronidae (Figs. 19.69, 19.428) construct similar tubes(Edwards 1961).

Suborder Integripalpia Apataniidae, Beraeidae, Brachycentridae, Calamoceratidae, Goeridae, Helicopsychidae, Lepidostomatidae, Leptoceridae, Limnephilidae, Molannidae, Odontoceridae, Phryganeidae, Rossianidae, Sericostomatidae, Thremmatidae, Uenoidae. Also,

Glossosomatidae, Hydrobiosidae, Hydroptilidae, Rhyacophilidae. Approximately 38 families are known worldwide in the Integripalpia(Holzenthal et al. 2011)and the 20 of them listed here are represented in North America. The case-making families are those whose larvae construct shelters of plant materials, rock fragments, or sometimes silk alone, which they carry with them while foraging for food(Figs. 19.1-19.31). The first 16 families listed above in Integripalpia make tubular cases. Most tube-case-making caddis larvae are detritivorous shredders, feeding on decomposing leaves primarily for the fungi and bacteria that colonize them. Others are scrapers of algal films and periphy ton, and a few are collector-gatherers of organic particles or are predators (Table 19A). Because these food resources occur in isolated patches, larvae move about in search offood,and their portable cases provide some physical and camouflaging protection against fish and other predators. Cases made of min eral fragments are beautiful examples of "masonry" with the mineral pieces fitted together with precision. Cummins(1964)observed that larvae of Pycnopsyche lepida (Hagen) on average coated ten particles with silk for every one that was actually glued in place. This fitting activity resulted in the well-constructed final

Chapter 19 Trichoptera

587

tl?J

Figure 19.3 Figure 19.4

Figure 19.5

Figure 19.1

Figure 19.6

Figure 19.2

Figure 19.9

Figure 19.10

Figure 19.7

Figure 19.1

Figure 19.8

Nectopsyche sp.(Leptoceridae) portable

case.

Figure 19.2 Figure 19.3 Figure 19.4 Figure 19.5 Figure 19.6 Figure 19.7

Figure 19.11

Triaenodes sp.(Leptoceridae) portable case. Ceraclea sp.(Leptoceridae) portable case. Oecetis sp.(Leptoceridae) portable case. Leptocerus sp.(Leptoceridae) portable case. Molanna sp.(Molannldae) portable case. Psilotreta sp.(Odontoceridae) portable case.

Figure 19.8 Beraea sp.(Beraeldae) portable case, with detail of posterior end. Figure 19.9 Helicopsyche sp.(Hellcopsychldae) portable case.

Figure 19.10 Agarodes sp.(Sericostomatldae) portable case.

Figure 19.11 case.

Gumaga sp.(Sericostomatldae) portable

588

Chapter 19 Trichoptera

Figure 19.15 Figure 19.12

Figure 19.13

Figure 19.16

Figure 19.14

Figure 19.20a

Figure 19.21

Figure 19.19 Figure 19.18 Figure 19.17

Figure 19.20b

Figure 19.12 Ptilostomis sp.(Phryganeidae) portable case, and detail of ring construction. Figure 19.13 Banksiola sp.(Phryganeidae) portable case.

Figure 19.14 Phryganea sp.(Phryganeidae) portable case, and detail of spiral construction. Figure 19.15 Fabria sp.(Phryganeidae) portable case, with detail of posterior end. Figure 19.16 Micrasema sp.(Brachycentridae) portable case.

Figure 19.20c

Figure 19.17 Brachycentrus sp.(Brachycentridae) portable case. Figure 19.18 Goeracea sp.(Goeridae) portable case, with detail of posterior end. Figure 19.19 Goera sp.(Goeridae) portable case, with detail of posterior end. Figure 19.20a-19.20c Lepidostoma spp.(Lepidostomatldae) portable cases. Figure 19.21 Theliopsyche sp.(Lepidostomatidae) portable case, with detail of posterior end.

Chapter 19 Trichoptera

product. Such cases have been marketed as jewelry (e.g., Wildscape, K. Stout). Moreover, tubular cases are an asset in respira tion because undulating movements by the larva cause a ventilating current of water to move through the tubular case, bathing the abdomen and tracheal gills. Experimental work has shown that larvae remove more oxygen from water, and thereby survive longer at low oxygen levels, when in their cases than when deprived of them (Jaag and Am buhl 1964); the rate of abdominal ventilation increases at lower oxy

gen levels (Van Dam 1938; Fox and Sidney 1953).

589

cryptic, as are those of some Calamoceratidae (Figs. 19.29-19.31, 19.95), in which Heteroplectron larvae

excavate twigs as cases (Fig. 10.178). Many larvae of the Lepidostomatidae (Figs. 10.183, 19.107), import ant components of the shredder community in cool streams, construct cases of sand grains or silk in early instars, later changing to four-sided cases of leaf and bark pieces (Figs. 19.20a-19.20c). Larvae of Thremmatidae and Uenoidae (Figs. 10.190, 19.27-19.28) live in running waters and often occur in aggregations on rocks. Uenoidae (Farula, Neothremma, and Sericostriata) inhabit cool mountain streams of western

Portable cases seem, therefore, to have released some

North America and Thremmatidae (western OUgo-

groups from respiratory dependence on stream cur rents and thus to have been an asset in the exploita tion of the resources of lentic habitats(Wiggins 1996,

phlebodes and widespread Neophylax) occur in run ning waters (Vineyard et al. 2005). Beraeidae (Figs. 10.176, 19.8, 19.111-19.112) are extremely localized in North America and have been recorded only from the water-saturated muck of spring seepage areas in the Last. Perhaps the most unusual larvae of all belong to the Helicopsychidae (Figs. 10.180, 19.9, 19.113), which construct cases coiled like the shell of a snail; these larvae graze diatoms and fine particulate matter from exposed surfaces of rocks in rivers and along wave-swept shorelines of lakes. All four extant basal families of Integripalpia occur in North America. These families (Glossosomatidae, Hydrobiosidae, Hydroptilidae, Rhyacophilidae) probably do not constitute a monophyletic group, but they do share several distinctive biological features that may have been characteristic of the ear liest species in the order. All construction activities by larvae of these four families ultimately is for pupation: at the completion of the final larval instar in the Rhyacophilidae and Hydrobiosidae; at the beginning of the final instar in the Hydroptilidae(Figs. 19.39-19.43); or at the begin ning of the first instar in the Glossosomatidae (Fig. 19.38). In the Glossosomatidae and Hydroptilidae, larvae use their precocious pupal enclosures as pro tective shelters while they forage for food. The archi tecture of the pupal enclosures in all four families is based on domes rather than tubes(Figs. 19.38, 19.39); they are constructed ofrock fragments in the Rhyaco philidae, Hydrobiosidae, and Glossosomatidae. The version used by larvae of Glossosomatidae has a transverse strap connecting the longer sides of the case beneath the middle of the larva (Figs. 10.170, 19.38) and the purse-cases of most Hydroptilidae are

2004).

The dominant tube-case-making family in North America is the Limnephilidae (Figs. 10.184, 19.4419.48, 19.51-19.53, 19.87-19.91, 19.196-19.205) with some 252 species and 41 genera in North America alone; the genera are highly diverse in case-making behavior (e.g.. Figs. 19.22-19.25), habitat, and food (Table 19A). Larvae are mainly detritivorous or omnivorous and have toothed mandibles (as in Fig. 19.49). Larvae in the Apataniidae and Goeridae have specialized mandibles with uniform scraping edges

(Fig. 19.50) and feed mainly by browsing exposed rock surfaces for diatoms and fine organic particles. Larval Phryganeidae (Figs. 10.187, 19.12-19.15, 19.76-19.80) are large, up to 40 mm in length, and usually have conspicuous yellow and black markings on the head and pronotum. Pupae in several phryganeid genera are unusual in having pupal mandibles reduced to membranous lobes(Fig. 19.551), a special ized feature correlated with the fact that they do not close the anterior end of the pupal case with a silken sieve membrane prior to metamorphosis as do other larvae (Wiggins 1960b). Leptoceridae (Figs. 19.119.5, 19.96-19.97) are biologically diverse and some are able to swim with their cases; larvae in several genera are predaceous, some in Ceraclea feed on sponges(Fig. 10.181; Resh et al. 1976). Larvae of the Odontoceridae (Figs. 10.186, 19.7, 19.108-19.110) and Sericostomatidae (Figs. 10.189, 19.10-19.11, 19.72-19.75) are primarily borrowers in loose sedi ments;some odontocerids {Psilotreta) pupate in dense clusters exposed on rocks. Larval Brachycentridae (Figs. 10.177, 19.16-19.17, 19.102-19.104) are con fined to running waters, and those of Brachycentrus are unusual among the case-making groups in filter ing food from the current with their outstretched legs. The flattened and dorsally cowled larval cases of Molannidae (Figs. 10.185, 19.6, 19.98-19.101) are

formed from two domes, mainly of silk, fastened together along the edges(Figs. 19.39-19.42). At pupa tion most larvae of these families construct a

rigid closed cocoon of tough darkened silk within the domed enclosure, leading to the name "closedcocoon-makers," which is often shortened to

590

Chapter 19 Trichoptera

Figure 19.25

Figure 19.23 Figure 19.24

Figure 19.26 Figure 19.22

f( Figure 19.27

Figure 19.31 Figure 19.28

Figure 19.29

Figure 19.22 Arctopora sp.(Limnephilldae) portable

Figure 19.27

case.

case.

Figure 19.23 Pseudostenophylax sp.(Limnephilldae) portable case. Figure 19.24 Psychoglypha sp.(Limnephilldae) portable case. Figure 19.25 Limnephilus sp.(Limnephilldae) portable case.

Figure 19.26 Rossiana sp.(Rossianidae) portable case, with detail of posterior end.

Figure 19.30

Neophylax sp.(Thremmatidae) portable

Figure 19.28 Neothremma sp.(Uenoidae) portable case.

Figure 19.29 Heteroplectron sp.(Calamoceratidae) portable case. Figure 19.30 Phylloicus sp.(Calamoceratidae) portable case, with detail of posterior end. Figure 19.31 Anisocentropus sp.(Calamoceratidae) portable case. with detail of posterior end.

Chapter 19 Trichoptera

591

Figure 19.32

Figure 19.33

Figure 19.34

Figure 19.35 \

Figure 19.37

Figure 19.36

Figure 19.32 Dolophilodes sp.(Phllopotamidae) net,

Figure 19.35 Macrostemum sp.(Hydropsychidae)

with detail of mesh.

larval retreat and capture net. Figure 19.36 Psychomyia sp. (Psychomyiidae) tubular

Figure 19.33 Phylocentropus sp.(Dipseudopsidae) buried branching tube. Figure 19.34 Hydropsyche sp.(Hydropsychidae) net and retreat.

retreats on a rock, with detail of a tube.

Figure 19.37 Neureclipsis sp.(Polycentropodidae) net.

592

Chapter 19 Trichoptera

"cocoon-makers" (Wiggins and Wichard 1989). Among all Trichoptera, these cocoons are distinctive in being osmotically semipermeable (Wichard et al. 1993). Consequently, oxygen for respiration reaches the metamorphosing pupa in these families solely by diffusion through the wall of the closed cocoon. Although all caddis larvae construct cocoons for metamorphosis, porous cocoons in the Integripalpia and Annulipalpia admit ambient water currents that carry oxygen directly to the pupa(Wiggins and Wich ard 1989; Wiggins 2004). Larvae in the families Rhyacophilidae (Figs. 10.173,19.57)and Hydrobiosidae(Figs. 10.171,19.58) forage actively and do not construct a retreat or case of any kind until just before pupation when a cell of rock fragments, usually dome-shaped, is fastened to a rock or other substrate. Within this enclosure the

larva spins a characteristic ovoid, closed cocoon of

tough, brown silk and undergoes metamorphosis inside the cocoon. Larvae are predators for the most part, although evidently some species feed on algae and vascular plant tissue; most inhabit cool running waters, some occur in transient streams. In the Rhya cophilidae, the genus Rhyacophila comprises nearly 800 species, almost entirely in the Northern Hemi sphere, and is one of the largest genera in the order; it is abundant and highly diverse in streams in western North America. The Hydrobiosidae, a family mainly of the Southern Hemisphere, extend into North America only in the Southwest. Larvae in the Glossosomatidae (Figs. 10.170, 10.196, 19.38, 19.59-19.61) construct portable cases of rock fragments resembling the domed shell of a tortoise; larvae are entirely covered by their domeshaped cases. They live in running waters and occa sionally along the wave-swept shorelines of lakes, grazing diatoms and fine particulate organic matter from the upper exposed surfaces of rocks. They often progress against the current by belaying from alter nating ends of the case: securing the upstream end with silk attached to the substrate (the "protection" belay), reversing direction in the case, and slowly swinging the downstream end upstream 180° as they graze (Monroe and Olden 2008). Oxygen-rich water

for respiration enters the case through spaces between rock pieces. In preparation for pupation, the larva removes the ventral strap and fastens the dome firmly to a rock with silk; a closed, brown, silken cocoon is

spun within the case as in the Rhyacophilidae. These larval cases are basically pupal enclosures constructed precociously at the beginning of the first instar. Larvae of the Hydroptilidae(Figs. 10.172, 19.3919.43, 19.62) are extremely small, and are free-living until the final instar when they construct purse-shaped

cases usually of two domes fastened together along two edges (Fig. 19.41), which are portable in most genera; barrel-shaped cases are constructed in a few genera (Fig. 19.43). The first four larval instars, approximately 1 mm long at maturity and completed within three weeks in some species, differ consider ably in morphology from the fifth instar, and repre sent the only example of larval hypermetamorphosis known in the Trichoptera. Larvae live in all types of permanent habitats, including springs, streams, riv ers, and lakes. Their primary food is algae, mainly the cellular contents of filamentous forms, but sessile dia

toms are ingested by species in some genera. Two of the few examples known in the Trichoptera of specific association with a food resource occur in the genera Dibusa and Palaeagapetus where final-instar larvae feed only on the freshwater red alga Lemanea or leafy liverworts of the order Jugermanniales (Fig. 10.172), respectively, and use the same plants in constructing their cases (Resh and Houp 1986; Ito et al. 2014). Although fifth-instar larvae in most hydroptilid gen era construct portable cases, those in the tribe Leucotrichiini are sedentary, fixing flattened silken domes resembling the egg capsules of leeches to rocks in running waters (Fig. 19.40). The head and thorax are extended through a small opening at either end to graze periphyton and particulate matter from the area surrounding the case. Food reserves of fifth instars cause the abdomens of hydroptilids to become dispro portionately large—depressed in the Leucotrichiini and Ptilocolepinae but compressed in most others (Wiggins 2004). Based on the specificity of case type for families of Integripalpia, Cummins et al (1965) constructed a key to the families based on case type alone.

Biology Although most species of North American caddisflies are univoltine, some require two years for development, and a few complete two generations in a year. Larvae of most species have five instars, in a few up to seven or even 14, after which they fasten the case with silk to a solid substrate, sealing off the ends. The actual pupal stage lasts from two to three weeks, although in some groups it is preceded by a prepupal phase of up to several weeks' duration

when the larva is in diapause (Wiggins 1996). When metamorphosis is complete, the pharate adult enclosed within the pupal cuticle (Figs. 19.51-19.53) leaves the pupal case and swims to the surface. Eclosion (called "hatching" by fly fishers) occurs either on the water surface of large streams or lakes, or on some emergent surface.

Chapter 19 Trichoptera

593

Figure 19.38

Figure 19.39 Figure 19.40

Figure 19.42 Figure 19.41

Figure 19.43

Figure 19.38 Glossosoma sp.(Glossosomatidae) portable case. Figure 19.39 Ochrotrichia sp.(Hydroptilidae) portable

Figure 19.41 Agraylea sp.(Hydroptilidae) portable case, right lateral and dorsal. Figure 19.42 Ithytrichia sp.(Hydroptilidae) portable

case.

case, left lateral and dorsal.

Figure 19.40 Leucotrichia sp.(Hydroptilidae) fixed retreat, dorsal and right lateral.

Figure 19.43 Neotrichia sp.(Hydroptilidae) portable case, ventral and posterior.

594

Chapter 19 Trichoptera

Adults of most species are quiescent during the day, but fly actively during the evening and hours of darkness. Some are known to feed on plant nectar (Crichton 1957). Although most adult caddisflies probably live less than one month, adult females of some Limnephilidae whose larvae inhabit temporary pools live for at least three months;their reproductive maturity is delayed by diapause until late summer and early autumn when drought conditions are wan ing (Novak and Sehnal 1963; Wiggins 1973a, 2004). Diapause intervenes to suspend development at vari ous points in the life cycles of other species. Eggs are deposited in water in most families, sometimes by females that swim (Deutsch 1985) or crawl (Elliot 1969) beneath the water surface, but above water in some groups ofthe Limnephilidae, and entirely in the absence of surface water by some species inhabiting temporary pools. A further specialization for drought occurs in some Polycentropodidae whose eggs remain in the dry basin of temporary pools and do not hatch until water is replenished the following spring (Wiggins et al. 1980). Eggs are enclosed in a matrix of spumaline (a polysaccharide complex; Hinton 1981), which in the Integripalpia is greatly expanded as water is absorbed. Otherwise, little is known about caddisfly eggs and egg masses or means for collecting and identifying them (but see, for exam ple, Hinton 1981; Wood et al. 1982; Lancaster and Glaister 2018). EXTERNAL MORPHOLOGY Larvae

Viewed dorsally (Fig. 19.48), the head capsule is subdivided into three parts by Y-shaped dorsal ecdysial lines or sutures: thefrontoclypeal apotome is sepa rated from the rounded parietals on each side by frontoclypeal sutures', posteromesally, the parietals meet dorsally along the coronal suture. Ventrally, the parietals come together along the ventral ecdysial line, but frequently the parietals are partially or completely separated mesally by the ventral apotome. Peg-like antennae (Fig. 19.48) are visible on larvae in families constructing portable cases, but are not apparent in most other groups. The eyes are groups ofstemmata. Cutting edges of the mandibles are of two basic types correlated with the method of feeding—a series of separate points or teeth(Fig. 19.49)or an entire scrap ing edge (Fig. 19.50). Silk is emitted from an opening at the tip of the labium (Fig. 19.49), which can be protracted in a long spinneret(Figs. 19.68, 19.154). The pronotum is covered by a heavily sclerotized plate subdivided by a mid-dorsal ecdysial line (Fig. 19.48);the prosternum sometimes bears small sclerites and,in somecase-makingfamilies,asemi-membranous

prosternal horn (Figs. 19.44, 19.47). The trochantin (Fig. 19.47)is a derivative of the prothoracic pleuron. The mesonotum may bear sclerotized plates (Fig. 19.48), small sclerites (Fig. 19.77), or be entirely membranous(Fig. 19.67); the metanotum (Fig. 19.48) in most families is largely membranous. Notal setae on the last two thoracic segments may be single or grouped on sclerites, but their basic arrangement into three setal areas—sa\, sal, and sal—is usually

apparent (Figs. 19.47, 19.48). In families with ambulatory larvae, the middle and hind pairs of legs are substantially longer than the first (Fig. 19.44). The abdomen (Fig. 19.44) consists of ten seg ments, usually entirely membranous except for a dorsomedian sclerite on segment IX in some families. This sclerite is not always pigmented and may be dif ficult to distinguish in certain groups. Segment I of the abdomen in families constructing portable tubecases usually bears prominent humps—one on each side and one dorsally; these humps, which serve as "spacers" facilitating uniform movement of the respi ratory current of water through the case, are retractile and may not be prominent in preserved specimens; if retracted, their presence may be detected by folds of the integument. Tracheal gills are filamentous exten sions of the body wall that contain fine tracheoles; some larvae lack gills entirely. Gills may be single (Fig. 19.44) or branched (e.g.. Fig. 19.87) and are generally arranged in dorsal, lateral, and ventral pairs of gills on each side of a segment. Lightly sclerotized ovoid rings are discernible ventrally on most abdom inal segments in larvae of the Limnephilidae (Fig. 19.44) and dorsally and laterally as well in certain limnephilid subgroups; these rings enclose chloride epithelia which are areas of the epidermis specialized for ionic transport in osmoregulation (Wichard and Komnick 1973). A lateralfringe of slender, bifid, hol low filaments extends along the mid-lateral surface of each side of most larvae in the Integripalpia (Fig. 19.44), each usually closely associated with a lateral fringe of smallforked lamellae. The function of both structures is unknown;they appear to be correlated in some way with living in portable cases, but do not occur in all Integripalpia (Kerr and Wiggins 1995). Anal prolegs terminating the abdomen each bears a pointed anal claw that sometimes has a small acces

sory hook dorsally or a comb of one or more teeth ventrally. The prolegs are short and laterally directed (Figs. 19.44, 19.45) in larvae with portable tube cases or purse cases, enabling them to hold fast to the silken lining ofthe case, but usually are longer in glossosomatids and are especially long in retreat-making and free-living larvae (Fig. 19.46). Merrill (1964) demon strated that sensory hairs at the base of the prolegs of three limnephilids and one phryganeid provide the

Chapter 19 Trichoptera

595

prosternal horn foretrochantin

-prosternal horn

prothorax

-foretrochantin

mesothorax

episternum

pleural suture |- pleu;on epimeron

metathorax

dorsal hump —■

femur

coxa

tarsus

trochanter

single

lateral fringe

^ tracheal

tarsal claw

basal seta'

Figure 19.47

gills

labrum

opening of silk gland ■ labium lateral mandible

chloride

tubercles

Figure 19.50

epithelium maxilla

labrum

Figure 19.49 dorsal sclerlte

mandible

frontoclypeal

antenna

apotome —

anal proleg dorsal

Figure 19.44

ecdysial

suture

line

coronal suture

muscle scars

parietal

pronotum

anal claw

lateral sclerlte

eye

' frontoclypeal

accessory hook mesonotum

Figure 19.45 anal claw metanotum —

Figure 19.46

Figure 19.48

Figure 19.44 Larval habitus (Limnephilldae), right lateral. Figure 19.45 Right anal proleg of a case-making larva (Limnephilldae), right lateral.

Figure 19.46 Right anal proleg of a retreat-making larva (Philopotamidae), right lateral. Figure 19.47 Larval thorax and hind leg (Limnephilldae), right lateral.

Figure 19.48

Larval head and thorax (Limnephilldae),

dorsal.

Figure 19.49 Larval mouthparts, right half, with toothed mandible (Leptoceridae), ventral. Figure 19.50 Larval right mandible with scraper edge (Uenoidae), ventral.

596

Chapter 19 Trichoptera

larva with information about case length and the need to add more building material at the leading edge of the case. When the hairs were removed, abnormally long cases resulted.

Pupae Heavily sclerotized pupal mandibles (Fig. 19.51) serve to cut an opening in the pupal case through which the insect escapes to swim to the surface for emergence (Wiggins 1960b). Stout single setae occur on various parts ofthe head, with those on the labrum hooked apically in some groups; dorsal and ventrolateral tufts of setae often occur near the bases of the

antennae (Fig. 19.51). The compacted wings conform tightly to the body, and the legs are folded ventrolaterally on the exarate pupa. Usually each of the middle tarsi bears a

dense fringe of setae (Fig. 19.52), rendering the leg more effective for swimming from the pupal case to the water surface.

Several of the abdominal segments bear paired dorsal sclerites—the hook plates (Fig. 19.52); hook plates are designated as anterior(a)or posterior(p)in accordance with their position on a particular seg ment, with at least segment V usually having both pairs. Hooks on anterior plates are directed posterad, and those on posterior plates directed anterad. In several families, segment I bears a rough spined ridge. These abdominal sclerites engage with the silken lin ing of the pupal case, enabling the insect to move within—especially important when it is ready to vacate the case for emergence. Pupal gills generally coincide with larval gills. A lateralfringe of slender filaments is variously developed along the sides in a number of families, absent in others; when present, the lateral fringes extend along the sides of several segments (Fig. 19.52), turning ventrad on segment VIII(Fig. 19.53). At the apex ofthe abdomen in many families is a pair of anal processes (Fig. 19.53); these are often elongate, but may also be short and lobate (e.g., Fig. 19.550). Adults

Structural characters of adult Trichoptera are identified in Figures 19.54—19.56. Setal warts are widely used in the key as diagnostic characters for families; usually clearly delineated by color and tex ture, the warts are somewhat dome-shaped in profile and bear macrosetae, only the basal pits of which are illustrated in the figures. Setal warts of the head in most families are reduced in various ways from the

generalized condition of Figure 19.55. The three ocelli occurring on the head in some families can be distin guished by their rounded, bead-like shape and gray or

silver color (Fig. 19.55). The number and relative lengths of segments in each maxillary palp is an important diagnostic character; segments are num bered 1-5 from base to apex(Fig. 19.54). The terminal segment (no. 5) of each maxillary palp in families of the Annulipalpia is flexible, and numerous irregular transverse striations of membrane and sclerotization

often can be seen in the cuticle, sometimes appearing as rings (Fig. 19.570, hence "Annulipalpia"). Sexual dimorphism occurs in the maxillary palpi of some families of the Trichoptera: Palpi of all females are five-segmented, but those of males are reduced in the number of segments—Phryganeidae, 4; Limnephilidae, Brachycentridae, and Helicopsychidae, 3 or less. Maxillary palpi and basal segments of the antennae may be markedly enlarged and distorted in males of the Lepidostomatidae and Sericostomatidae. An illustration of the head and first two thoracic seg ments is provided for each family as a means of sub stantiating identifications based on other characters. Tibial spurs of the legs are large, modified setae occurring usually in pairs at the apex of each tibia, and sometimes singly or paired in a preapical position (Fig. 19.54). The full complement ofspurs is expressed in an abbreviated formula, e.g., 3, 4, 4 for Figure 19.54, giving the respective total number of spurs on each of the fore-, mid-, and hind legs. Wings of caddisflies(Fig. 19.56) provide a wealth of taxonomic characters involving shape, venation, and other aspects. In general, from anterior to poste rior, the longitudinal veins include costa(C),subcosta (Sc), radius(R, with up to 6 apical branches), media (M, with up to 4 apical branches), cubitus (Cu, with up to 3 apical branches), and up to 3 anal(A) veins generally looped and fused apically in forewings; up to 4 anal veins apically independent in hind wings. These longitudinal veins are often connected by crossveins, designated by lower-case letters, usually for the longitudinal veins they connect, and some times closing special cells, such as the discoidal cell (closed by sectoral crossvein s at the fork of R2+3 and R4+5, usually with a nygma), the median cell(closed by m at the fork of Ml+2 and M3+4), the thyridial cell(closed by m-cu between M and Cul, often with a nygma), and the anal cell(closed by la-2a at the bases of lA and 2A). Other crossveins may include h (humeral crossvein between the bases of C and Sc), r (between R1 and R2), r-m (between R5 and Ml), and cu-a (between Cu2 and lA). Apical branches of the longitudinal veins may be present and numbered with Roman numerals, including Fork I(branch of R2and R3), Fork II (branch of R4 and R5, usually with a nygma). Fork III (branch of Ml and M2), Fork IV (branch of M3 and M4), and Fork V (branch of Cula and Culb). Occasionally, Sc and/or R1 may have

Chapter 19 Trichoptera

597

dorsal antennal tuft antenna

forewing

ventrolateral

antennal tuft

hind wing

mandible

seta of labrum hooked

Figure 19.51

spined ridge anterior hook plates mesotarsus

lateral fringe

IVa ^

modified for

posterior

swimming.

bookplate

abdominal

gills

lateral fringe

anal processes

Figure 19.53 Figure 19.52

pronotum

propleuron

metanotum

mesonotum

lateral ocellus

metapleuron scape,

median ocellus-

maxillary palp

coxa coxa

genital appendages(d)

labial

mesopleuron

palp

femur

VI®

trochanter

tibial spurs (preapical) tarsal

tibial spurs' (apical)

claws

spines

Figure 19.54 tarsus

Figure 19.51

Head of pupa (Limnephilidae),

frontal.

Figure 19.53 Apex of abdomen of pupa (Limnephilidae), showing lateral fringes of filaments and

Figure 19.52 Pupa (Limnephilidae), with details of median spined ridge (abdominal segment I) and hook

anal processes, ventral.

plates, habitus, dorsal.

wings omitted, left lateral.

Figure 19.54 Adult habitus (Rhyacophilidae), with

anteromesal setal wart median ocellus

flagellum pedicel

compound eye

scape

central setal area

anterior s.w.

lateral ocellus

posterior s.w.

posterolateral s.w.

prothorax pronotal s.w. median fissure mesoscutal s.w. mesothorax -

scutum

mesoscutellar s.w.

scutellum

postscutellum scutum

metathorax scutellum

postscutellum Figure 19.55

Sc

stigma I

R1

nygma nygma

arculus

Cu1b

Figure 19.56

Figure 19.55 Head and thorax of generalized adult, with wings omitted, dorsal (s.w. = setal wart).

Figure 19.56 Dolophilodes novusamericana (Phllopotamidae) right fore- and hindwings, dorsal; ac : anal cell, do = discoidal cell, mc = median cell, tc = thyridial cell, other abbreviations explained in text.

598

Chapter 19 Trichoptera

apical forks. Collectively, crossveins and the bases of apical branches often form a transverse, irregular line called the anastomosis or cord. Apical branches with their bases in the anastomosis are said to be "sessile,"

those originating well before the anastomosis "rooted," and those originating beyond it "stalked" or "pedunculate." The stigma region on the forewing anterior margin near the apices of Sc and R1 is often opaque. The apex of Cu2 is curved backward to the forewing posterior margin as an arculus, often hyaline and sometimes fused with the combined anal veins.

Segment I ofthe abdomen is reduced ventrally, an important consideration when counting segments(Fig. 19.54). A pair of pheromone glands sometimes opens on the sternum of segment V, often in raised ovoid sclerotized areas (Fig. 19.584) or in slender filaments. These are present in at least some representatives ofthe Rhyacophilidae, Glossosomatidae, Hydroptilidae, Flydropsychidae, Psychomyiidae, Polycentropodidae, Phryganeidae, Brachycentridae, Limnephilidae, Beraeidae, and Molannidae (Schmid 1980; Djernaes 2011). Males(e.g.. Fig. 19.54; Nielsen 1957)are readily distinguished from females (e.g.. Fig. 19.728; Nielsen 1980) by generally more-complex external genitalic structures terminating the abdomen.

Sampling/Collecting, Preserving, and Studying Trichoptera An excellent overview for quantitative sampling and qualitative collecting of caddisfly larvae and pupae is provided in Chapter 3, and the references provided in Table 3A. Like those immature stages, adults of caddisflies exhibit many different behaviors such that taxonomic diversity is best assessed with a variety of sampling and collecting techniques. Sam pling with emergence traps or window traps can be reasonably quantitative. Collecting with white or ultraviolet lights that are suspended before bed sheets or over pans of alcohol or soapy water or over tun neled jars of killing agent, Malaise traps that are set across flyways, aspirators that lift specimens from resting places, or aerial sweep nets that are raked through riparian vegetation can capture a wide range of day-flying or night-flying specimens. Some details for these collecting techniques were provided by Blahnik and Holzenthal (2004). Eggs and soft-bodied larvae and pupae of Trichoptera are best preserved in ethanol(EtOH) for long-term storage. Flexibility is sometimes important for manipulations of mandibles and other appendages, so that preservation in 80% ethanol is appropriate. However, because water is destructive for DNA,speci mens intended for DNA sequencing should be pre served in 95-100% ethanol (Frandsen and Thomson

599

2016). Poor specimen quality results when alcohol does not penetrate and adequately "fix" soft tissues. These specimens may become discolored, shrunken, and sub ject to decomposition. Identification of specimens of inferior quality is problematic and their use in system atic study is limited greatly. To help minimize these problems, the initial alcohol used to kill and preserve the specimens should be replaced with fresh alcohol.

Larval and pupal specimens ofthe highest quality are achieved by enhancing the initial fixation process. This can be accomplished in several ways. One method involves heating freshly captured specimens to the boiling point in either water or in alcohol. This is most easily accomplished in the field by placing the speci men in a glass vial containing 80% ethanol and then using a cigarette lighter as a heat source. Again, the specimens should be later transferred to fresh alcohol for long-term storage. An alternative, and the pre ferred method for fixing and initial preservation, is to place freshly captured specimens for two to three weeks in a special fixative solution containing forma lin, ethanol, and glacial acetic acid. Two such com mon and commercially available fixatives are Kahle's fluid and Pampel's solution. The disadvantage of boiling or using these special fixatives is that specimen DNA is damaged and unusable for sequencing. Addi tionally,formalin presents a health hazard and special disposal procedures are required. Adult Trichoptera also can be preserved in etha nol. Doing so preserves some flexibility and the char acters of the mesonotum. On the other hand, careful

preservation ofspecimens dry on pins maintains body color better and keeps most of the hair, including those hairs that contribute to color patterns of wings. Also, DNA sequencing remains an option longer for pinned specimens than for alcohol-preserved speci mens (Frandsen and Thomson 2016). Because of these different advantages, if a series of specimens for a species is captured, it is best to preserve some adults in alcohol and some specimens on pins (Blahnik and Holzenthal 2004). Adults ofsome families and many genera are iden

tified by characters of wing venation. Because hair often obscures wing venation, especially crossveins, it may be necessary to remove the hair and mount wings on microscope slides or between cover slips. This is accomplished most effectively by removing wings from one side of the specimen (customarily the right side), stroking them gently with a pair of soft-bristle watercolor brushes under water with a few drops of ethanol to clean all the hair from both sides of each

wing, then allowing the wings to dry under a cover slip pressed and sealed with glue or tape on a micro scope slide or between two cover slips (Blahnik and Holzenthal 2004).

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Chapter 19 Trichoptera

KEY TO THE FAMiLIES OF TRICHOPTERA LARVAE

1.

Larvae construct portable cases of sand grains or small rock fragments, coiled to resemble snail shells (Figs. 19.9, 19.178, 19.460a, 19.460b); anal claw comb-shaped (Figs. 19.113, 19.177); widespread HELICOPSYCHIDAE .... Helicopsyche (p. 618)

1'.

Larvae construct cylindrical portable cases that do not resemble coiled snail

shells, or larva does not construct portable case; anal claw with stout apical hook (Figs. 19.45, 19.46)

2(1'). 2' 3(2).

3'.

4(3').

4'.

5(2').

5'.

2 Metanotum entirely covered by pair of sclerites (Figs. 19.62, 19.63) 3 Metanotum entirely membranous (Fig. 19.71), or largely so but with several pairs of smaller sclerites (Fig. 19.90) 5 Abdomen with ventrolateral rows of branched gills, and with prominent tuft of long setae at base of anal proleg (Fig. 19.63). Larvae construct fixed retreats of detritus and rock fragments (Figs. 19.34, 19.35); widespread HYDROPSYCHIDAE (p. 622) Abdomen without either branched gills or tuft of setae at base of anal proleg (Fig. 19.62) 4 Anal claw large and at least as long as its sclerotized basal segment, anal proleg projecting freely from abdomen, (Fig. 19.70); body length typically more than 5 mm. Larvae construct fixed tubular retreats of sand (e.g.. Fig. 19.36); Texas ECNOMIDAE .... Austi-otinodes(p. 616) Anal claw relatively small, anal proleg usually not projecting freely from abdomen (Figs. 19.62, 19.234), sometimes projecting freely (Figs. 19.233, 19.235); body length typically less than 5 mm. Larvae construct purse-shaped portable cases of silk (Figs. 19.42, 19.240, 19.243, 19.244, 19.246, 19.504, 19.505, 19.507) sometimes with sand (Fig. 19.39) or plant material (Figs. 19.41, 19.237-19.239, 19.245, 19.504, 19.506); flat silken domes fastened to rocks (Figs. 19.40, 19.241); or cylindrical cases covered with tiny sand grains

(Figs. 19.43, 19.242); widespread HYDROPTILIDAE (p. 625) Antennae markedly elongate and prominent, at least 6 times as long as wide (Fig. 19.96), and/or sclerotized plates on mesonotum lightly pigmented except for pair of dark curved lines on posterior half(Fig. 19.97). Larvae construct portable cases of various materials (Figs. 19.1-19.5, 19.265-19.269, 19.497-19.501, 19.503); widespread LEPTOCERIDAE(p. 630) Antennae of normal length, not more than 3 times as long as wide (Fig.19.107), or not apparent(Fig. 19.67); mesonotum never with pair of dark curved lines as above

6(5').

6'.

7(6). 7'. 8(7).

6

Mesonotum largely or entirely membranous (Fig. 19.71), or with small sclerites covering not more than half of notum (Fig. 19.77); pronotum never with anterolateral lobe (Fig. 19.76) 7 Mesonotum largely covered by sclerotized plates, variously subdivided (Figs. 19.102-19.104, 19.88), and typically pigmented, although sometimes lightly (Fig. 19.112); pronotum sometimes with prominent anterolateral lobe (Figs. 19.111, 19.112) 15 Abdominal tergum IX with sclerite, sometimes pale and inconspicuous or obscured by posterior margin of abdominal tergum VIII (Figs. 19.59, 19.60) 8 Abdominal tergum IX entirely membranous(Fig. 19.73) 11 Metanotal sa3 typically consisting of cluster of setae arising from small rounded or ovoid sclerite (Figs. 19.77, 19.372-19.374); prosternal horn present (Figs. 19.79 and 19.368). Larvae construct tubular portable cases, typically of plant materials (Figs. 19.12-19.15, 19.376-19.380, 19.453, 19.454, 19.474, 19.495, 19.496); widespread PHRYGANEIDAE (p. 646)

Chapter 19 Trichoptera

601

chelate

Figure 19.58

foreleg

accessory

ry

hook

Figure 19.62

secondary lateral claw

Figure 19.59

Figure 19.57

Figure 19.60 Figure 19.61

ry

Figure 19.57 Rhyacophila sp.(Rhyacophilidae) larval habitus, dorsal.

ry

Figure 19.58 Atopsyche sp.(Hydrobiosidae) right larval foreleg, right lateral. Figure 19.59 Glossosoma sp. (Glossosomatidae) larval habitus, with detail of segments IX and X and right anal proleg, right lateral.

Figure 19.60 Glossosoma sp.(Glossosomatidae) larval segments IX and X and anal prolegs, caudal. Figure 19.61 Agapetus sp.(Glossosomatidae) larval head and thorax, dorsal.

Figure 19.62 Agraylea sp.(Hydroptilidae) larval habitus, right lateral, with enlargement of chloride epithelium, dorsal.

trochantin

coxa

I Figure 19.64

Figure 19.63

branched

gitis trochantin

tuft of setae

Figure 19.66

T

Figure 19.65

Figure 19.67

Figure 19.63 Hydropsyche sp.(Hydropsychidae) larval habitus, right lateral.

Figure 19.64 Psychomyia sp.(Psychomyildae) larval habitus, with details of right foretrochantin and right anal claw, right lateral.

Figure 19.65 Nyctiophylax sp.(Polycentropodidae) larval habitus, with detail of right anal claw, right lateral. 602

Figure 19.66 Neureclipsis sp.(Polycentropodidae) larval habitus, with details of right foretrochantin and right anal claw, right lateral. Figure 19.67 Neureclipsis sp.(Polycentropodidae) larval head and pro- and mesothoraces, dorsal.

Chapter 19 Trichoptera

603

8'.

Metanotal sa3 consisting of single seta with or without sclerite (Fig. 19.61); prosternal horn absent(Fig. 19.59). Larvae either without portable tubular cases, or with tortoise-like domed cases of rock fragments(Fig. 19.38)

9(8').

Anal prolegs each with basal half broadly joined with segment IX, its anal claw with at least one dorsal accessory hook (Figs. 19.59, 19.60). Larvae construct tortoise-like domed portable cases of rock fragments (Figs. 19.38, 19.167, 19.168, 19.459a, 19.459b); widespread GLOSSOSOMATIDAE(p. 616) Anal prolegs each mostly free from segment IX, its anal claw without dorsal accessory hook, although secondary lateral claw may be present (Fig. 19.57). Larvae free-living, without cases or fixed retreats until pupation 10

9'.

9

10(9').

Tibia, tarsus, and claw of each foreleg shortened, articulated against extended lobe of femur to form chelate appendage (Fig. 19.58); Southwest HYDROBIOSIDAE .... Atopsyche (p. 622)

10'.

Foreleg normal, not modified as chelate appendage (Fig. 19.57); widespread

11(7').

RHYACOPHILIDAE(p. 656) Labrum membranous and T-shaped (Fig. 19.71), often withdrawn from view in preserved specimens. Larvae construct fixed sac-shaped nets of silk (Fig. 19.32); widespread PHILOPOTAMIDAE(p. 646)

11'.

Labrum sclerotized, rounded, articulated in normal way and never

withdrawn (Fig. 19.67) 12 Mesopleuron extended anteriorly on each side as lobate process (Fig. 19.69); tibiae and tarsi of all legs fused together (Fig. 19.69). Larvae construct fixed, meandering, tubes of sand on rocks; Arizona, Texas XIPHOCENTRONIDAE(p. 666) 12'. Mesopleura unmodified, tibiae and tarsi separate on all legs (Fig. 19.65) 13 13(12'). Foretrochantins each extended as broadened, hatchet-shaped lobe best seen from ventrolateral direction (Fig. 19.64). Larvae construct fixed tubular 12(11').

retreats of sand and debris on rocks and logs(Fig. 19.36);

13'.

widespread PSYCHOMYIIDAE(p. 656) Foretrochantins each with apex acute, not expanded into broadened lobe (Fig.19.66) 14

14(13'). Tarsi of all legs markedly flattened, tibiae shorter than tarsi (Fig. 19.68). Larvae construct tubes of sand and silk, buried in usually soft sediments

(Fig. 19.33); East

DIPSEUDOPSIDAE .... Phylocentropus (p. 616)

14'.

Tarsi of all legs more or less cylindrical and not flattened, tibiae as long as or longer than tarsi (Fig. 19.65). Larvae construct exposed funnel-shaped or tubular filter nets of silk (Fig. 19.37) or flattened retreats; widespread POLYCENTROPODIDAE(p. 652)

15(6').

Abdominal segment I lacking both dorsal and lateral humps (Fig. 19.102), metanotal ral absent(Fig. 19.103) or represented only by single seta without sclerite. Larvae construct portable cases of silk and usually sand and/or transversely arranged plant materials that are either cylindrical (Figs. 19.16, 19.134-19.136, 19.467, 19.469, 19.491) or 4-sided (Figs. 19.17, 19.132, 19.133, 19.468, 19.470, 19.490, 19.492); widespread BRACHYCENTRIDAE(p. 613) Abdominal segment I always with lateral humps although not always prominent, and usually with median dorsal hump (Figs. 19.76, 19.86); metanotal sa\ always present, typically represented by sclerite bearing several setae (Fig. 19.81) but with at least a single seta. Larvae construct portable cases of widely differing form and materials 16

15'.

T-shaped labrum

mesopleuron

Figure 19.69

Figure 19.68 Figure 19.70 Figure 19.71

antenna

trochantin

lateral sclerite

Figure 19.75

cluster of setae

Figure 19.73

Figure 19.72

Figure 19.74

Figure 19.68 Phylocentropus sp.(Dipseudopsidae) larval habitus, with details of right fore- and middle tarsi and claws, right lateral. Figure 19.69 Xiphocentron sp.(Xiphocentronidae) larval prothorax and mesopleuron, right lateral. Figure 19.70 Austrotinodes sp.(Ecnomldae) larval habitus, right lateral. Figure 19.71 Chimarra sp.(Philopotamidae) larval head and thorax, with detail of labrum, dorsal. 604

Figure 19.72 Agarodes sp.(Sericostomatidae) larval habitus, with detail of right anal claw, right lateral. Figure 19.73 Agarodes sp.(Sericostomatidae) larval segments IX and X and anal prolegs, dorsal. Figure 19.74 Agarodes sp.(Sericostomatidae) larval head and thorax, dorsal.

Figure 19.75 Fattigia sp. (Sericostomatidae) larval right foretrochantin, right lateral.

prosternal horn

Figure 19.79

Figure 19.77

Figure 19.80

Figure 19.83

Figure 19.76

Figure 19.78

Figure 19.82

Figure 19.81

Figure 19.84

Figure 19.76 Oligostomis sp. (Phryganeldae) larval habitus, right lateral. Figure 19.77 Oligostomis sp.(Phryganeldae) larval

Figure 19.85

Figure 19.86

Figure 19.82 Moseiyana sp.(Apatanlidae) larval left mandible, ventral.

Figure 19.83 Allomyla sp.(Apatanlidae) larval left

head and thorax, dorsal.

mandible, ventral.

Figure 19.78 Agrypnia sp.(Phryganeldae) larval head

Figure 19.84 Apatania sp.(Apatanlidae) larval head

and thorax, dorsal.

and thorax, dorsal.

Figure 19.79 Agrypnia sp.(Phryganeldae) larval prothorax, ventral. Figure 19.80 Yphria sp.(Phryganeldae) larval head

Figure 19.85 Neophylax sp.(Thremmatidae) larval

and thorax, dorsal.

Figure 19.81

head and thorax, dorsal.

Figure 19.86 Neophylax sp.(Thremmatidae) larval habitus, right lateral.

Moseiyana sp.(Apatanlidae) larval head

and thorax, dorsal. 605

606

Chapter 19 Trichoptera

antenna

prosternal horn

Figure 19.91 Figure 19.89

Figure 19.87

Figure 19.90

Figure 19.88

Figure 19.92

Figure 19.94

Figure 19.93

Figure 19.87 Anabolia sp.(Limnephilidae) larval habitus, right iaterai. Figure 19.88 Anabolia sp.(Limnephiiidae) larvai head and thorax, dorsai.

Figure 19.89 Platycentropus sp.(Limnephiiidae) iarvai head and prosternai horn, right iaterai. Figure 19.90 Umnephilus sp.(Limnephiiidae) larval head and thorax, dorsal.

Figure 19.91

Homophylax sp.(Limnephiiidae) larval

labrum, dorsal.

Figure 19.92 Goereilla sp.(Rossianidae) larval habitus, with detail of anal claw, right iaterai. Figure 19.93 Goereilla sp.(Rossianidae) larval head and thorax, dorsai.

Figure 19.94 Rossiana sp.(Rossianidae) larval head and thorax, anterior right lateral.

Chapter 19 Trichoptera

16(15').

^

Mesopleura extended anteriorly on each side as prominent, acutely or bluntly pointed process (Figs. 19.105, 19.106). Larvae construct cases of rock

fragments (Figs. 19.18-19.19, 19.173-19.176, 19.482-19.486); East, Midwest, Northwest

^

607

GOERIDAE(p. 618)

16'.

Mesopleura not extended anteriorly as pointed processes (e.g., Figs. 19.88, 19.95)

17(16').

Labrum with transverse row of approximately 16 setae across central area (Fig. 19.95). Larvae construct cases of wood and other plant materials variously arranged (Figs. 19.29-19.31, 19.151, 19.152, 19.461-19.463); East, Southeast, Southwest, West Coast CALAMOCERATIDAE(p. 616)

17

17'.

Labrum not as above, typically with only 6 setae across central area (Fig.19.91)

18(17').

Antennae situated on each side near anterior margin of eye (Fig. 19.107); abdominal segment I without median dorsal hump (Fig. 19.107). Larvae construct cases of sand (Figs. 19.21, 19.493) or plant materials

18

(Figs. 19.20b, 19.20c), with 4-sided cases of panels of plant materials common (Figs. 19.20a, 19.466); widespread LEPIDOSTOMATIDAE(p. 630) 18'. Antennae not near anterior margins of eyes, situated approximately as close to anterior margin of head capsule as to eyes(Fig. 19.89) or closer (Fig. 19.74); abdominal segment I almost always with median dorsal hump (Fig.19.72) 19 19(18'). Antennae situated approximately midway between anterior margin of head capsule and eyes (Fig. 19.89); prosternal horn typically present(Fig. 19.89) although sometimes short; chloride epithelia typically present on at least some abdominal segments(Fig. 19.44) 20 19'. Antennae situated at or near anterior margin of head capsule (Fig. 19.74); prosternal horn and chloride epithelia absent(Fig. 19.72) 24 20(19). Mesonotum notched anteromedially (Figs. 19.85, 19.421, 19.422, 19.425-19.427) 21 20'.

Mesonotum not notched anteromedially (Figs. 19.88, 19.90)

21(20).

Larvae very slender, without gills; pronotum broadest anteriorly in dorsal view and anteromedial notch of mesonotum deep (Figs. 19.425-19.427); constructing slender cases of silk (Fig. 19.456) or sand and silk (Figs. 19.28, 19.487, 19.488); West

22

UENOIDAE(p. 660)

21'.

Larvae not slender, with gills; pronotum broadest about middle in dorsal view and anteromedial notch of mesonotum shallow (Figs. 19.85, 19.421, 19.422); constructing stouter cases of rock fragments, often with small stones arranged linearly along each side (Figs. 19.27, 19.494); widespread THREMMATIDAE(p. 660)

22(20').

Mandibles each typically with uniform scraper blade (Fig. 19.83) or, if mandible toothed (Fig. 19.82), then more than 25 setae present on membranous area of metanotum between sa\ sclerites (Fig. 19.81); metanotal ^al sclerites absent in some genera. Larvae construct cases mainly of mineral materials (Figs. 19.126, 19.429, 19.430, 19.481, 19.502); Appalachian Mtns., North, West APATANIIDAE (p. 611)

22'.

Mandibles almost always toothed (Fig. 19.49); setae typically absent from metanotum between sa\ sclerites (Fig. 19.90) or, if present, then fewer than 25 setae

23(22').

23

Mesonotum with one large sclerite on each side of midline (Fig. 19.88); chloride epithelia typically present on at least some abdominal segments(Fig. 19.87). Larvae construct cases of plant (Figs. 19.22, 19.25, 19.337-19.339, 19.341, 19.342, 19.432-19.434, 19.440, 19.441b, 19.445-19.447, 19.449, 19.450, 19.480,

19.509-19.511) or mineral materials (Figs. 19.23, 19.335, 19.336, 19.340, 19.343, 19.438, 19.441a, 19.442a, 19.448, 19.451, 19.452a, 19.508), or a combination of the two (Figs. 19.24, 19.431, 19.435-19.437, 19.439, 19.442c-l 9.444, 19.452b), occasionally with mollusk shells (Fig. 19.442b); widespread LIMNEPFIILIDAE (p. 633)

curved line

Figure 19.96

Figure 19.97

Figure 19.95

Figure 19.99 Figure 19.101

Figure 19.98

Figure 19.100

Figure 19.95 Anisocentropus sp.(Calamoceratidae)

Figure 19.99 Molanna sp.(Molannldae) larval head

larval head and thorax, with detail of labrum, dorsal.

and thorax, dorsal.

Figure 19.96 Triaenodes sp.(Leptocerldae) larval Figure 19.97 Ceraclea sp.(Leptocerldae) larval head

Figure 19.100 Molannodes sp.(Molannldae) larval habitus, with detail of right hind tarsus, right lateral. Figure 19.101 Molannodes sp.(Molannldae) larval

and thorax, with detail of left antenna, dorsal.

head and thorax, dorsal.

head and thorax, dorsal.

Figure 19.98 Molanna sp.(Molannldae) larval habitus, with detail of right hind tarsal claw, right lateral. 608

f

Figure 19.104

Figure 19.103

Figure 19.102

mesopleuron

^

\

MirMV.;?:

r -V;L;^

Figure 19.105

Figure 19.106

Figure 19.107

Figure 19.102 Micrasema sp.(Brachycentridae) larval

Figure 19.106 Goera sp.(Goerldae) larval head and

habitus, right lateral.

thorax, dorsal.

Figure 19.103 Micrasema sp.(Brachycentridae) larval

Figure 19.107 Lepidostoma sp.(Lepidostomatidae)

head and thorax, dorsal.

larval habitus, with details of apex of hind right coxa and eye and antenna, right lateral.

Figure 19.104 Brachycentrus sp.(Brachycentridae) larval head and thorax, dorsal.

Figure 19.105 Goeracea sp.(Goerldae) larval head and thorax, dorsal. 609

610

Chapter 19 Trichoptera

trochantin

Figure 19.109

lateral sclerite

Figure 19.111

I

•• • ,•

Figure 19.108

Figure 19.113 Figure 19.112

Figure 19.108 Psilotreta sp.(Odontoceridae) larval habitus, with details of right foretrochantin and anal proleg, right lateral. Figure 19.109 Psilotreta sp.(Odontoceridae) larval

Figure 19.111

Beraea sp.(Beraeidae) larval habitus,

with detaiis of right serrate lamellae and anal proleg, right lateral. Figure 19.112 Beraea sp. (Beraeidae) iarval head and

head and thorax, dorsal.

thorax, dorsal.

Figure 19.110 Psilotreta sp.(Odontoceridae) larval

Figure 19.113 Helicopsyche sp.(Helicopsychidae) larval right anal claw, right lateral.

segment IX and anal prolegs, dorsal.

Chapter 19 Trichoptera

23'.

611

Mesonotum with 2(Fig. 19.94) or 3(Fig. 19.93) smaller sclerites on each side of midline; chloride epithelia absent(Fig. 19.92). Larvae construct cases almost entirely of rock fragments(Figs. 19.26, 19.411, 19.455, 19.464); Northwest

ROSSIANIDAE (p. 656)

24(19'). Tarsal claw of each hind leg modified to form short stub or slender filament (Figs. 19.98, 19.100). Larvae construct cases of sand grains with flanges on sides and overhanging anterior opening (Figs. 19.6, 19.457, 19.458); Central, East, far Northwest MOLANNIDAE (p. 642) 24'. Tarsal claws of hind legs not different in structure from those of other legs (Fig. 19.72) 25 25(24'). Pronotum with transverse carina or ridge extended into rounded anterolateral lobe on each side (Figs. 19.111, 19.112). Larvae construct cases of sand grains (Figs. 19.8, 19.489); East(highly localized) BERAEIDAE .... Beraea (p. 613) 25'.

Pronotum without transverse carina, and anterolateral corners not lobate,

although pointed in some genera (Fig. 19.109) 26(25'). Dorsum of each anal proleg with cluster of approximately 30 or more setae

26'.

26

mesal of lateral sclerite (Fig. 19. 73), lateral sclerite relatively small in dorsal view (Fig. 19.73); foretrochantins relatively large and hook-shaped apically (Fig. 19.75). Larvae construct cases mainly of fine rock fragments (Figs. 19.10, 19.11, 19.412, 19.475-19.477); widespread SERICOSTOMATIDAE(p. 660) Dorsum of each anal proleg with no more than 3-5 setae mesal of lateral scleidte, although with short spines in some genera, lateral sclerite relatively large (Fig. 19.110); foretrochantins smaller than above and not hooked apically (Fig. 19.108). Larvae construct cases of rock fragments (Figs. 19.7, 19.348, 19.349, 19.465, 19.471-19.473, 19.478, 19.479); widespread ODONTOCERIDAE(p. 642)

KEYS TO THE GENERA OF TRICHOPTERA LARVAE

Through the efforts of various workers, especially H.H. Ross, O.S. Flint, and G.B. Wiggins, larvae of almost all of the North American caddisfly genera have been associated and described; in fact, only 4 of the 155 North American genera recognized in this chapter are not yet known in the larval stage. The keys to larvae of North American genera are based on the work of Wiggins(1996). Even more so than for families, accurate determinations for larvae of genera are most likely obtained when last instar specimens are examined. North American genera for which larvae have not yet been described include 1 genus of Hydropsychidae (Oropsyche), 2 of Limnephilidae (Chilosligmodes and Leptophylax), and 1 of Philopotamidae (Sisko). Apataniidae 1. Metanotal iulsclerites present, about same size as sal sclerites (Fig. 19.114) 1'.

Metanotal 5fllsclerites absent, ^alsetae forming transverse row (Fig. 19.115) or patch

2(1).

(Fig. 19.116) Basal seta of each tarsal claw short, much shorter than its claw (Fig. 19.119);

2

4

mandibles each with several teeth (Figs. 19.82, 19.123); many metanotal setae arising from membranes(Fig. 19.117); case of fine sand grains, strongly tapered and curved, covered with shiny silk (Fig. 19.429); Oregon,

Washington 2'.

Moselyana comosa Denning'

Basal seta of each tarsal claw long, extending to or almost to tip of claw (Fig. 19.120); mandibles each with apical edge entire, without teeth

(Fig. 19.124); most metanotal setae confined to sclerites (Fig. 19.114) 'This genus is represented in North America by only one species(see also Table 19A).

3

612

Chapter 19 Trichoptera

Figure 19.114

, \\\ UW/'////// \\

Figure 19.117

/

sa1

//

Figure 19.115

Figure 19.116

sciente

Figure 19.120 seta seta

Figure 19.119

Figure 19.118 teeth carina

Figure 19.123

Figure 19.124

stout primary seta

\

Figure 19.121

Figure 19.126

Figure 19.122

Figure 19.125

Figure 19.114 Manophylax annulatus Wiggins (Apataniidae) metanotum, dorsai. Figure 19.115 Apatania arizona Wiggins (Apataniidae) meso- and metanota, dorsai.

Figure 19.116 Pedomoecus sierra Ross (Apataniidae) meso- and metanota, dorsai.

Figure 19.117 Moselyana comosa Denning (Apataniidae) metanotum, dorsal. Figure 19.118 Manophylax annulatus Wiggins (Apataniidae) abdominal sternum I, ventral. Figure 19.119 Moselyana comosa Denning (Apataniidae) left metathoracic tarsal claw, posterior (left lateral). Figure 19.120 Manophylax annulatus Wiggins (Apataniidae) left metathoracic tarsai claw, posterior (left lateral).

© R.W. Holzenthal 2{)(I6

Figure 19.121 Allomyla scotti(Wiggins)(Apataniidae) head, right lateral. Figure 19.122 Manophylax annulatus Wiggins (Apataniidae) head, right lateral. Figure 19.123 Moselyana comosa Denning (Apataniidae) left mandible, ventral. Figure 19.124 Manophylax annulatus Wiggins (Apataniidae) left mandible, ventral. Figure 19.125 Pedomoecus sierra Ross (Apataniidae) head, dorsal.

Figure 19.126 Pedomoecus sierra Ross (Apataniidae) larval case.

Chapter 19 Trichoptera

3(2').

3'.

Abdominal sternum I with anteromedian sclerite (Fig. 19.118), with or without central unsclerotized area; head unmodified and uniformly convex (Fig. 19.122); case of rock fragments, somewhat depressed, tapered and slightly curved, with plant pieces dorsolaterally; Alaska, Appalachian Mtns., Idaho(madicolous or dry rock mountain habitats)

Manophylax

Abdominal sternum I without anteromedian sclerite; dorsum of head flattened,

frequently with prominent carina (Fig. 19.121); case of coarse rocks, tapered, curved, with larger pebbles laterally (Figs. 10.175, 19.430); West(mountains) 4(1').

613

Allomyia

Metanotal sal setae arranged as transverse row (Fig. 19.115), mesonotum with 2 large, undivided plates (Fig. 19.115); dorsum of head with most primary setae unmodified; case of rocks, strongly tapered, anterior opening usually oblique

with dorsal edge extending beyond ventral edge for final instar (Fig. 19.502); Appalachian Mtns., North, Southwest 4'.

Apatania

Metanotal ^al setae in transversely elliptical patch (Fig. 19.116); mesonotum with 2 median and 2 lateral plates (Fig. 19.116); dorsum of head with some primary setae unusually large, stout(Fig. 19.125); case of rocks, smooth, strongly tapered, slightly curved (Figs. 19.126, 19.481); Northwest Pedomoecus sierra

Beraeidae Pronotum with transverse carina extended as rounded anterolateral lobes

(Figs. 19.127, 19.128); mesonotum consisting of single undivided plate; metanotum lacking sclerites and with single transverse elliptical patch of ial setae (Fig. 19.112); lateral lamellae of abdomen serrate; lateral sclerite of each anal proleg conical with large apical seta (Fig. 19.111); case of fine sand, curved and tapered, smooth (Figs. 10.176, 19.8, 19.489); East Brachycentridae 1. Meso- and metathoracic legs long, their femora about as long as head capsule, their tibiae each produced distally into prominent process from which stout spur arises (Fig. 19.130); case usually square in cross section, composed of small pieces of plant materials fastened transversely (Figs. 10.177, 19.17, 19.132, 19.468, 19.470), although case sometimes cylindrical and largely of silken secretion or with a mixture of sand and plant material(Fig. 19.467), or occasionally with small rock projections (Fig. 19.136); widespread 1'. Meso- and metathoracic legs shorter, their femora much shorter than head capsule (Fig. 19.131), each tibia not produced distally into prominent process, although

2(1').

2'.

spur arising from about same point on unmodified tibia (Fig. 19.131) Ventral apotome of head longer than wide, narrowed somewhat posteriorly; rudimentary prosternal horn present on anterior part of prosternum (Fig. 19.129); case 4-sided and tapered, composed of short pieces of plant material placed crosswise with loose ends often protruding (Figs. 19.133, 19.490, 19.492) Ventral apotome of head usually wider than long (Fig. 19.137), sometimes squarish (Fig. 19.138); prosternal horn absent; case cylindrical, tapered, straight or curved, composed of lengths of plant material wound around the circumference (Figs. 19.16, 19.134) or of silk or silk and rock material (Fig. 19.135)

'This genus is represented in North America by only one species (see also Table 19A).

Beraea

Brachycentrus

2

3

4

carna

Figure 19.

Figure 19.128

ventral

apotome

F-

'r tYS. i I /\ horn

Figure 19.129 0

Figure 19.127 S.-^\

fe

r:

rw

Figure 19.136

Figure 19.132

Figure 19.133

Figure 19.134

Figure 19.135

R.W. Holzenthal 2006

Figure 19.127 Beraea gorteba Ross (Beraeidae) head

Figure 19.132 Brachycentrus sp.(Brachycentridae)

and thorax, dorsal.

larval case.

Figure 19.128 Beraea gorteba Ross (Beraeidae) pronotum, right lateral. Figure 19.129 Adicrophleps hitchcocki Flint (Brachycentridae) head and prosternum, ventral. Figure 19.130 Brachycentrus sp. (Brachycentridae) right metathoracic leg, posterior (right lateral). Figure 19.131 Micrasema wataga Ross (Brachycentridae) right metathoracic leg, posterior (right lateral).

Figure 19.133 Adicrophleps hitchcocki Flint (Brachycentridae) larval case. Figure 19.134 Micrasema wataga Ross (Brachycentridae) larval case. Figure 19.135 Micrasema rusticum (Hagen) (Brachycentridae) larval case. Figure 19.136 Brachycentrus echo (Ross) (Brachycentridae) larval case.

614

Chapter 19 Trichoptera

ventral

pronotal

pronotal

groove

groove

615

apotome

Figure 19.138

Figure 19.137

Figure 19.139

Figure 19.140

sal

Figure 19.143

variable suture

Figure 19.141

Figure 19.144

Figure 19.142 Figure 19.145 © R.W. Holzenthal 2006

Figure 19.137 Amiocentrus aspilus (Ross) (Brachycentridae) head, ventral. Figure 19.138 Micrasema wataga Ross (Brachycentridae) head, ventral. Figure 19.139 Micrasema wataga Ross (Brachycentridae) pronotum, right lateral. Figure 19.140 Amiocentrus aspiius (Ross) (Brachycentridae) pronotum, right iaterai. Figure 19.141 Eobrachycentres geiidae Wiggins (Brachycentridae) mesonotum, dorsal.

Figure 19.142 Adicrophleps hitchcocki Flint (Brachycentridae) mesonotum, dorsal. Figure 19.143 Micrasema wataga Ross (Brachycentridae) mesonotum, dorsai. Figure 19.144 Amiocentrus aspilus (Ross) (Brachycentridae) mesonotum, dorsal. Figure 19.145 Micrasema wataga Ross (Brachycentridae) abdominal segment X, caudoventral.

616

3(2).

Chapter 19 Trichoptera

Each half of mesonotum usually entire, lateral quarter partially delineated by variable suture, posterior margin raised and colored dark brown (Fig. 19.141);

Northwest 3'.

Eobrachycentms gelidaeV^'xggms^

Each half of mesonotum divided into 3 separate sclerites, posterior margin not conspicuously raised or colored (Fig. 19.142);

Northeast 4(2').

Adicrophleps hitchcocki Flint'

Transverse pronotal groove curving anteriorly, with ends of groove usually meeting anterior margin (Fig. 19.139), each end sometimes forming rounded lateral lobe; brown, sclerotized band on either side of anus (Fig. 19.145); mesonotal sa\ with multiple setae (Fig. 19.143) or solitary seta on each side; widespread

4'.

Micrasema

Pronotal groove not curving anteriorly and never reaching anterior margin (Fig. 19.140); no brown sclerotized bands near anus; mesonotal sal with

solitary seta on each side (Fig. 19.144); Northwest

Amiocentms aspilus (Ross)'

Calamoceratidae

1.

T.

Anterolateral corners of pronotum produced into prominent lobes (Figs. 19.146, 19.147); gills with 2 or 3 branches (Fig. 19.149)

2

Anterolateral corners of pronotum somewhat extended (Fig. 19.148), but much less than above; gill filaments single (Fig. 19.150); cases of hollowed twigs (Figs. 10.178, 19.29, 19.151, 19.462a, 19.462b); East, West Coast

2(1).

2'.

Heteroplectron

Metathoracic legs about as long as mesothoracic legs; anterolateral corners of pronotum pointed (Fig. 19.147,); case of pieces of bark and leaves (Figs. 19.152, 19.463); Southwest

Phylloicus

Metathoracic legs about twice as long as mesothoracic legs; anterolateral corners of pronotum rounded (Fig. 19.146); case of 2 leaf pieces, dorsal piece overhanging ventral one (Figs. 19.31, 19.461);

Southeast

Anisocentropus pyraloides(Walker)'

Dipseudopsidae Tarsi of all legs compressed, tibiae shorter than tarsi (Fig. 19.153); spinneret half as long as head capsule (Fig. 19.154); retreats branching tubes of silk covered with sand and buried in fine sand, except tips of branches exposed (Figs. 10.163, 19.33); East

Phylocentropus

Ecnomidae

Flead depressed and nearly as long as thorax (Figs. 10.164, 19.70, 19.155); foretrochantins each longer than its coxa and half as long as head, depressed (Figs. 19.70, 19.155); retreat fixed tube of rock fragments held together with silk; Texas

Austrotinodes texensis Bowles'

Glossosomatidae^

1.

Mesonotum with 2 or 3 sclerites (Figs. 19.156, 19.157, sometimes hard to see); head with ventromesal margins of genae not thickened, posterior median ventral ecdysial line about 1.5 times as long as each anterior divergent branch (Figs. 19.159, 19.160); anal opening without dark, sclerotized line on each side

2

'This genus is represented in North America by only one species (see also Table 19A). ^ The cases of Glossosomatidae genera are all dome-like dorsally, composed of stones (Figs. 10.196, 19.38, 19.168), often with large flat stones laterally (Fig. 10.170, 19.167), each with a transverse band of finer sand ventrally.

Chapter 19 Trichoptera

617

lobe'

Figure 19.148

Figure 19.147

Figure 19.146

')•! /v IS^

Wm ,

Figure 19.188

OCQO" -.r

II

>

f/

Cfl'l'/ Vr, )/ 0 -■/ ®

'"I'l l

V I //1 1

' >

{?-> ^

Figure 19.190

Figure 19.189

Figure 19.192 furrow

Figure 19.193

Figure 19.191 ) R.W. Holzenthal 2006

Figure 19.180 Arctopsyche irrorata Banks (Hydropsychldae) head, ventral. Figure 19.181 Parapsyche cardis Ross (Hydropsychldae) head, ventral. Figure 19.182 Diplectrona modesta Banks (Hydropsychidae) head, ventral. Figure 19.183 Hydropsyche betteni Ross (Hydropsychidae) head, ventral. Figure 19.184 Smicridea fasciatella MacLachlan (Hydropsychldae) head, ventral. Figure 19.185 Potamyia flava (Hagen) (Hydropsychidae) head, ventral. Figure 19.186 Macrostemum Carolina (Banks) (Hydropsychidae) gill of abdominal segment IV.

Figure 19.187 Hydropsyche betteni Ross (Hydropsychidae); gill of abdominal segment IV.

Figure 19.188

Cheumatopsyche sp. (Hydropsychidae)

left foretrochantin, left lateral.

Figure 19.189 Potamyia fiava (Hagen) (Hydropsychidae) left foretrochantin, left lateral. Figure 19.190 Dipiectrona modesta Banks (Hydropsychidae) pronotum, dorsal. Figure 19.191 Homoplectra sp. (Hydropsychidae) pronotum, dorsal. Figure 19.192 Parapsyche cardis Ross (Hydropsychidae) abdominal segment IV, left lateral. Figure 19.193 Arctopsyche irrorata Banks

(Hydropsychidae) abdominal segment IV, left lateral.

622

Chapter 19 Trichoptera

Hydrobiosidae Forelegs chelate, each with shortened tibia, tarsus, and claw close against concave extension offemur (Figs. 19.58, 19.179); free-living (Fig. 10.171); Southwest

Atopsyche

Hydropsychidae^'^ 1. 1'.

2(1').

2'.

Genae of head capsule completely separated by single ventral apotome (Figs. 19.180, 19.181) Genae touching ventrally, separating ventral apotome into anterior and posterior parts (Figs. 19.182, 19.183, 19.185) or posterior part inconspicuous(Fig. 19.184) Posterior ventral apotome much longer than broad, at least half as long as median ecdysial line where genae touch (Fig. 19.182); frontoclypeal apotome broad behind eyes

3'.

Abdominal gills each with up to 40 filaments arising fairly uniformly along central stalk (Fig. 19.186); foretrochantins never forked (Figs. 19.204, 19.205) Abdominal gills each with about 10 filaments arising mostly near apex of central stalk (Fig. 19.187); foretrochantins usually forked (Fig. 19.188), sometimes not(Fig. 19.189)

4(1).

4'.

5(2).

5'. 6(3').

Most abdominal segments dorsally with tuft of long setae and/or scale hairs on each sal and sa3 position (Fig, 19.192); ventral apotome of head usually nearly rectangular (Fig. 19.181); widespread Most abdominal segments with single long seta in each sal and sal position, frequently with 1 or 2 shorter setae, but without tuft (Fig. 19.193); ventral apotome narrowed posteriorly (Figs. 10.165, 19.180); widespread

7(6').

5

3

9

6

Parapsyche

Arctopsyche

Pronotum with transverse furrow separating narrower posterior l/3rd from

broader anterior 2/3rds(Fig. 19.191); East, West Pronotum without transverse furrow, but constricted only slightly at posterior border (Fig. 19.190); widespread Abdominal sternum VIII with single median sclerite (Fig. 19.194); submentum rounded or sinuous apically, but not notched (Fig. 19.184); Southwest

6'.

2

Posterior ventral apotome no longer than broad, much less than half as long as median ecdysial line (Figs. 19.183, 19.185) or inconspicuous(Fig. 19.184); frontoclypeal apotome not expanded behind eyes, V-shaped (Fig. 19.206) or U-shaped (Figs. 19.198, 19.199, 19.207)

3(2').

4

Abdominal sternum VIII with pair of sclerites (Figs. 19.195-19.197); submentum notched apically (Fig. 19.183)

Homoplectra Diplectrona

Smicridea

7

Prosternum with pair of large sclerites in intersegmental fold posterior to prosternal plate (Fig. 19.200); frontoclypeus entire (Fig. 19.198);

widespread T.

Prosternum with pair of usually small sclerites posterior to prosternal plate (Fig. 19.201); if sclerites large, frontoclypeus with shallow mesal excision (Fig. 19.199)

Hydropsyche

8

'The larva of Oropsyche (North Carolina) is unknown, but probably keys to couplet 5. The only representative of the genus in North America is O. howellae Ross.

The retreats of most genera (except Macrostemum) are covered with sand or plant material, aligned with the current, and each provided with a capture net offset to one side of the anterior end; the capture net is often supported by pieces of debris and silken guy-lines (Fig. 19.34), with its rectangular mesh size corresponding with the optimal current speed for the species.

Chapter 19 Trichoptera

t

623

//

Figure 19.194

Figure 19.195

Figure 19.198 excision

notch

Figure 19.199

Figure 19.197

Figure 19.196

e R.W, Holzeiithal 2006

Figure 19.194 Smicridea fasciatella McLachlan (Hydropsychidae) abdominal sternum VIII, ventral. Figure 19.195 Hydropsyche betteni Ross (Hydropsychidae) abdominal sternum VIII, ventral. Figure 19.196 Cheumatopsyche sp.(Hydropsychidae)

Figure 19.197 Potamyia flava (Hagen) (Hydropsychidae) abdominai sterna VIII and IX, ventral. Figure 19.198 Hydropsyche betteni Ross (Hydropsychidae) head, dorsai. Figure 19.199 Cheumatopsyche sp.(Hydropsychidae)

abdominai sterna Vlil and IX, ventral.

head, dorsal.

624

Chapter 19 Trichoptera

sclerite

Figure 19.200

Figure sclerite

setal fringe

Figure 19.204 flange

Figure 19.202

Figure 19.203

foretrochantin

Figure 19.205

Figure 19.206

Figure 19.207

Figure 19.200 Hydropsyche betteni Ross (Hydropsychidae) prosternum, ventral.

Figure 19.201 Cheumatopsyche sp.(Hydropsychidae) prosternum, ventral. Figure 19.202 Potamyia flava (Hagen) (Hydropsychidae); right mandible, dorsal. Figure 19.203 Cheumatopsyche sp.(Hydropsychidae) right mandible, dorsal. Figure 19.204 Macrostemum Carolina (Banks) (Hydropsychidae) left prothoracic leg, posterior (left lateral).

©R.W. Holzenthal 2006

Figure 19.205 Leptonema sp.(Hydropsychidae) left prothoracic leg, posterior (ieft lateral). Figure 19.206 Macrostemum Carolina (Banks) (Hydropsychidae) head, dorsal. Figure 19.207 Leptonema sp.(Hydropsychidae) head, dorsal.

Chapter 19 Trichoptera

8(7).

Anterior ventral apotome of head with prominent anteromedian projection (Fig. 19.185); posterior margin of each sclerite on abdominal sternum IX entire (Fig. 19.197); lateral border of each mandible flanged (Fig. 19.202); foretrochantins forked or not(Figs. 19.188, 19.189); Central,

8'.

Anterior ventral apotome without anteromedian projection (similar to Fig. 19.183; posterior margin of each sclerite on abdominal sternum IX notched (Fig. 19.196); mandibles not flanged (Fig. 19.203); foretrochantins

East

Potamyiaflava (Hagen)'

forked (Fig. 19.188); widespread 9(3).

9'.

625

Cheumatopsyche

Tibia and tarsus of each prothoracic leg with dense, dorsal setal fringe (Fig. 19.204); dorsum of head flattened and margined with sharp carina (Fig. 19.206); retreat an open-ended chamber of fine sand and silk, with its silken capture net of elongated mesh spun across chamber and with larva in tubular diverticulatum beside main chamber (Fig. 19.35); Central, East

Macrostemum

Tibia and tarsus of each prothoracic leg lacking dense, dorsal setal fringe (Fig. 19.205); dorsum of head convex and without carina (Fig. 19.207);

Texas

Leptonema albovirens(Walker)'

Hydroptilidae (final larval instar only)

•

1. r. 2(1).

Abdomen dorsoventrally depressed (Figs. 19.208-19.211) Abdomen laterally compressed (Figs. 19.233-236) Abdominal segments V and VI usually abruptly broader than others in dorsal aspect(Figs. 19.208, 19.209); case depressed, oval, made of

2 5

silk with small circular opening near each end, fastened to rock

^

(Figs. 19.40, 19.241) 2'. 3(2).

3

Abdominal segments V and VI never abruptly broader than others in dorsal aspect(Figs. 19.210, 19.211) Abdominal tergites II-VII with pair of small circular punctures near midline (Fig. 19.208 inset); basal seta of tarsal claw on each leg large,

claw appearing bifid (Fig. 19.212); West 3'.

4(2').

4'.

5'.

Zumatrichia notosa(Rossy

Abdominal tergites II-VII solid, without punctures(Fig. 19.209 inset); basal seta of tarsal claw on each leg much shorter and thinner than claw (Fig. 19.213); widespread

Leucotrichia

Abdominal segments I-VIII each with truncate, fleshy tubercle on each side (Fig. 19.211 inset); pro-, meso-, and metanotal plates each divided on meson by ecdysial suture (Fig. 19.211); case flat, elliptical valves covered with pieces of liverwort (Figs. 10.172, 19.237, 19.506); Northeast, Northwest Abdominal segments without lateral tubercles (Fig. 19.210); pronotal plate divided on meson by ecdysial suture, but meso- and metanotal plates not divided (Fig. 19.210); larva free-living without case;

Southwest 5(1').

4

Palaeagapetus

Alisotrichia arizonica (Blickle and Denning)'

Tarsal claws stout and abruptly curved, each with thick, blunt basal seta (Fig. 19.214) Tarsal claws slender, gradually curved, each with thin, pointed basal seta (Figs. 19.216-19.218), or seta apparently absent (Fig. 19.215)

'This genus is represented in North America by only one species(see also Table 19A).

6 7

626

Chapter 19 Trichoptera

ecdysial suture

ecdysial

tubercle

suture

VIII

Figure 19.211

IX

Figure 19.208

Figure 19.209

Figure 19.210

Figure 19.212

basal seta

Figure 19.213

basal seta

Figure 19.214

Figure 19.215 Figure 19.217

basal seta

dorsal ring

sclerlte

r-

Figure 19.218

-A Figure 19.219

Figure 19.220

Figure 19.208 Zumatrichia notosa (Ross) (Hydroptilidae) larva, dorsal; inset, sclerlte of abdominal tergum III.

Figure 19.209 Leucotrichia sp. (Hydroptilidae) larva, dorsal; Inset, sclerlte of abdominal tergum III. Figure 19.210 Alisotrichia sp.(Hydroptilidae) larva, dorsal.

Figure 19.211 Palaeagapetus celsus (Ross) (Hydroptilidae) larva, dorsal; Inset, left fleshy tubercle of abdominal segment II. Figure 19.212 Zumatrichia notosa (Ross) (Hydroptilidae) right mesothoracic tarsal claw, posterior (right lateral).

Figure 19.213 Leucotrichia sp. (Hydroptilidae) right mesothoracic tarsal claw, posterior (right lateral).

© R.W. Holzenthal 2006

Figure 19.214 Dibusa angata Ross (Hydroptilidae) right mesothoracic tarsal claw, posterior (right lateral). Figure 19.215 Metrichia nigritta (Banks) (Hydroptilidae) right mesothoracic tarsal claw, posterior (right lateral). Figure 19.216 Oxyethira sp.(Hydroptilidae) right metathoracic leg, posterior (right lateral). Figure 19.217 hiydroptila sp.(Hydroptilidae) right metathoracic leg, posterior (right lateral). Figure 19.218 Ithytrichia sp.(Hydroptilidae) right metathoracic leg, posterior (right lateral). Figure 19.219 Dibusa angata Ross (Hydroptilidae) abdominal segment III, dorsal. Figure 19.220 Stactobieila delira (Ross) (Hydroptilidae) abdominal segment III, dorsal.

Chapter 19 Trichoptera

6(5).

Dorsal abdominal setae stout, each arising from small sclerite, dorsal rings distinct (Fig. 19.219); case of 2 long, parallel-sided valves of red

6'.

Dorsal abdominal setae thin and without basal sclerites, dorsal rings indistinct (Fig. 19.220); 2 elliptical valves of silk case with few or no inclusions (Fig. 19.239); widespread

algae (Figs. 19.238, 19.504); East

7(5'). 7'.

8(7). 8'. 9(8). 9'. 10(9).

Dibusa angata Ross'

Each tarsal claw with basal seta (Figs. 19.216-19.218) Tarsal claws of middle and hind legs each apparently lacking basal seta (Fig. 19.215); case bivalved, made of silk with algal filaments included concentrically (similar to Fig. 19.245); Central, Southwest Protibiae each with prominent ventral lobe bearing short, stout setae (Figs. 19.221, 19.222)

Stactobiella 8

Metrichia 9

Protibiae lacking prominent lobes, their setae normal (Fig. 19.223), if present Thoracic legs approximately same length (Fig. 19.224)

14 10

Middle and hind legs much longer than forelegs (Fig. 19.225) 3 filamentous gills arising from posterior end of abdomen, 1 from dorsomedian position on segment IX, other 2 at lateral sclerites of anal prolegs, often difficult to distinguish from setae (Fig. 19.226)

13

10'.

Filamentous gills absent from posterior end of abdomen

11(IO').

Meso- and metanota usually with projecting anterolateral lobes(Fig. 19.224); abdominal segments usually each with dorsomedian sclerite and transverse ventral sulcus; case portable, laterally compressed, usually consisting of 2 silk valves covered with sand or occasionally filamentous algal strands (similar to Figs. 19.239, 19.245), but sometimes dorsoventrally depressed, with dorsal valve carried like tortoise shell and ventral sheet flat (Fig. 19.39); widespread

11'.

Meso- and metanota without projecting anterolateral lobes (similar in this character to Fig. 19.225); abdominal segments lacking dorsomedian sclerites and transverse ventral sulci; case portable, laterally compressed, usually consisting of 2 valves constructed of filamentous algal strands

12(10).

Base of each mesotarsal claw smoothly contoured with its apicoventral margin

(similar to Fig. 19.238); California

^

627

12 11

Ochrotrichia

Nothotrichia shasta Harris and Armitage'

(Fig- 19.229); case portable, compressed, consisting of 2 silken valves usually 12'.

covered with sand grains or sometimes diatoms (Fig. 10.194); widespread Base of each mesotarsal claw quadrate and angular (especially claws of middle and hind legs), not smoothly contoured with its apicoventral margin (Fig. 19.230); case portable, compressed, consisting of 2 silken

Hydroptila

valves with little additional material added to exterior surface;

13(9').

13'.

Arkansas Paucicalcaria ozarkensis Mathis and Bowles' Antennae long and slender, longer than diameter of cluster of stemmata (Fig. 19.231); ventral lobe of each protibia parallel-sided (Fig. 19.221); case portable, compressed, entirely of silk, shaped like flask, open posteriorly (Figs. 19.240, 19.507); widespread Oxyethiva Antennae shorter than diameter of cluster of stemmata (Fig. 19.232); ventral lobe of each protibia triangular (Fig. 19.222); case portable, compressed, incorporating filamentous algae in concentric circles (Figs. 19.41, 19.245); widespread except not deep Southeast

'This genus is represented in North America by only one species (see also Table 19A).

Agraylea

628

Chapter 19 Trichoptera

Figure 19.221

Figure 19.224

Figure 19.225 tibia

Figure 19.222 Figure 19.227

Figure 19.228 Figure 19.223

Figure 19.226 antenna

Figure 19.229 /

Figure 19.230 Figure 19.231

Figure 19.232

anal

proleg

anal proleg

Figure 19.233

Figure 19.235

Figure 19.234

Figure 19.236 © R.W. Holzenthal 2006

Figure 19.221 Oxyethira sp. (Hydroptilidae) right prothoracic leg, posterior (right lateral). Figure 19.222 Agraylea multipunctata Curtis (Hydroptilidae) right prothoracic leg, posterior (right lateral). Figure 19.223 Mayatrichia sp.(Hydroptilidae) right prothoracic leg, posterior (right lateral). Figure 19.224 Ochrotrichia sp. (Hydroptilidae) head and thorax, right lateral. Figure 19.225 Oxyethira sp. (Hydroptilidae) head and thorax, right laterai. Figure 19.226 Hydroptila sp.(Hydroptilidae) abdominal segments IX and X, left lateral. Figure 19.227 Mayatrichia sp.(Hydroptilidae) right femur, tibia, and tarsus, posterior (right laterai). Figure 19.228 Neotrichia sp.(Hydroptilidae) right femur, tibia, and tarsus, posterior (right lateral).

Figure 19.229 Hydroptila sp. (Hydroptilidae) right tarsus and tarsal ciaw, posterior (right lateral). Figure 19.230 Paucicalcaria ozarkensis Mathis and Bowles (Hydroptilidae) right tarsus and tarsal claw, posterior (right iateral). Figure 19.231 Oxyethira sp.(Hydroptilidae) head, right lateral. Figure 19.232 Agraylea multipunctata Curtis (Hydroptilidae) head, right lateral.

Figure 19.233 Neotrichia sp. (Hydroptilidae) abdomen, left lateral.

Figure 19.234 Orthotrichia sp. (Hydroptilidae) abdomen, left lateral.

Figure 19.235 Mayatrichia ayama Mosely (Hydroptilidae) abdomen, left lateral. Figure 19.236 Ithytrichia sp.(Hydroptilidae) abdomen, left lateral.

Chapter 19 Trichoptera

629

1^1 Figure 19.237

Figure 19.242

Figure 19.238

Figure 19.239

Figure 19.243

Figure 19.240

Figure 19.244

Figure 19.241

Figure 19.245

Figure 19.246

© R.W, Holzenthal 2006

Figure 19.237 Palaeagapetus celsus (Ross) (Hydroptilldae) larval case, dorsal. Figure 19.238 Dibusa angata Ross (Hydroptilldae) larval case, lateral.

Figure 19.239 Stactobiella delira (Ross) (Hydroptilldae) larval case, lateral. Figure 19.240 Oxyethira sp. (Hydroptilldae) larval Figure 19.241

Figure 19.243 Mayatrichia ayama Mosely (Hydroptilldae) larval case, lateral. Figure 19.244 Ithytrichia sp.(Hydroptilldae) larval case, lateral.

Figure 19.245 Agraylea sp. (Hydroptilldae) larval case, lateral.

case, lateral. case, dorsal.

Figure 19.242 Neotrichia sp.(Hydroptilldae) larval case, ventral.

Leucotrichia sp.(Hydroptilldae) larval

Figure 19.246 Orthotrichia sp.(Hydroptilldae) larval case, dorsal.

630

14(8').

14'. 15(14).

15'.

16(14').

16'.

Chapter 19 Trichoptera

Anal prolegs long and cylindrical, projecting prominently beyond general body outline (Figs. 19.233, 19.235) Anal prolegs short, conforming to general body outline, not projecting

prominently (Fig. 19.234) Mesotibiae each with pair of short, stout ventral setae located apically or nearly apically (Fig. 19.227); case portable, made of silk or including soft mineral material, tapered posteriorly, cylindrical but usually with longitudinal or transverse and longitudinal ridges (Fig. 19.243); widespread Mesotibiae each with pair of ventral setae located about l/3rd distance from apex (Fig. 19.228); case portable, made with fine sand grains, cylindrical (Figs. 19.43, 19.242); widespread Most abdominal segments with prominent, pointed, dorsal and ventral projections (Fig. 19.236); flat silk case open posteriorly, reduced to small circular opening anteriorly (Figs. 19.42, 19.244, 19.505); widespread Abdominal segments without dorsal and ventral projections (Fig. 19.234); silk case with longitudinal ridges (Fig. 19.246); widespread

Lepidostomatidae 1. Ventral apotome of head as long as, or longer than, median ecdysial line (Fig. 19.247); case usually 4-sided, of quadrate pieces of leaves or bark (Figs. 19.20a, 19.466), but pieces may be arranged irregularly (Fig. 19.20c), transversely (Fig. 10.183), or spirally (Fig. 19.20b), or case may be of sand grains; widespread 1'.

Ventral apotome of head shorter than median ecdysial line (Fig. 19.248); case of sand grains (Figs. 19.21, 19.493); East

15

16

Mayatrichia

Neotrichia

Ithytrichia

Orthotrichia

Lepidostoma Theliopsyche

Leptoceridae 1.

Tarsal claw of each mesothoracic leg hooked and stout; tarsus curved (Fig. 19.249); slender case of transparent silk (Figs. 19.5, 19.267, 19.500);

r.

Tarsal claw of each metathoracic leg slightly curved and slender; tarsus straight (Fig. 19.250) Sclerotized, concave plate with marginal spines on each side of anal opening and extending onto ventral lobe (Fig. 19.251); cylindrical case of stones (Figs. 19.266, 19.498); Central, East Sclerotized, spiny plates absent, although patches of spines or setae may be present(Fig. 19.254) Maxillary palpi extending far beyond labrum; mandibles long and blade-like, with sharp apical tooth separated from remainder of teeth (Fig. 19.252 and inset); cases of various types and materials (Figs. 19.4, 19.501); widespread Maxillary palpi extending little, if any, beyond labrum; mandibles short, wide, with teeth grouped close to apex around central concavity (Fig. 19.253 and inset) Mesonotum with pair of dark, curved bars on weakly sclerotized plates (Figs. 19.97, 19.270); abdomen broad basally, tapering posteriorly, with gills usually in clusters of 2 or more (Fig. 19.257); cases of various shapes and materials (Fig. 19.3), sometimes including spicules and pieces of freshwater sponges (Figs. 10.181, 19.499); widespread

Central, East

2(1').

2'.

3(2').

3'.

4(3').

'This genus is represented in North America by only one species(see also Table 19A),

Leptocems americanus(Banks)' 2

Setodes 3

Oecetis

4

Ceraclea

ventral

apotome

Figure 19.247

Figure 19.250 Figure 19.251 Figure 19.248

maxillary palp submesal

lateral

spines

spines

"I'll

Figure 19.252

Figure 19.254

Figure 19.253

spines

Figure 19.255

Figure 19.256

1 Figure 19.257 Figure 19.259

© R.W. Holzenthal 2006

Figure 19.258

Figure 19.247 Lepidostoma sp.(Lepidostomatidae) head, ventral. Figure 19.248 head, ventral.

Theliopsyche sp.(Lepidostomatidae)

Figure 19.249 Leptocerus americanus (Banks) (Leptoceridae) left mesothoracic leg, posterior, left lateral. Figure 19.250 Oecetis sp.(Leptoceridae) left mesothoracic leg, posterior, left lateral. Figure 19.251 Setodes incertus (Walker) (Leptoceridae) abdominal segment X, caudoventral. Figure 19.252 Oecetis sp.(Leptoceridae) head, dorsal; Inset, left mandible.

Figure 19.254 Triaenodes tarda Milne (Leptoceridae); abdominal segments IX-X, ventral. Figure 19.255 Triaenodes sp.(Leptoceridae) mandibles, ventral.

Figure 19.256 Mystacides sp.(Leptoceridae) mandibles, ventral.

Figure 19.257 Ceraclea sp.(Leptoceridae) abdomen, dorsal.

Figure 19.258 Triaenodes tarda Milne (Leptoceridae) abdomen, dorsal.

Figure 19.259 Nectopsyche sp.(Leptoceridae) abdominal segments IX and X, ventral.

Figure 19.253 Ceraclea maculata (Banks) (Leptoceridae) head, dorsal; Inset, left mandible.

631

ventral

apotomev

%

m

Figure 19.260 Figure 19.261

Figure 19.262

\

constriction

Figure 19.266

Figure 19.263

Figure 19.267

Figure 19.264

Figure 19.265

Figure 19.268

Figure 19.269

R.W. Holzenthal 2018

Figure 19.270

Figure 19.260 Nectopsyche sp.(Leptoceridae) head, ventral.

Figure 19.261

Triaenodes tarda Milne (Leptoceridae)

head, ventral.

Figure 19.262 Nectopsyche sp.(Leptoceridae) left metathoracic leg, posterior (left lateral). Figure 19.263 Triaenodes tarda Milne (Leptoceridae) left metathoracic leg, posterior (left lateral). Figure 19.264 Mystacides sp.(Leptoceridae) left metathoracic leg, posterior (left lateral). Figure 19.265 Triaenodes tarda Milne (Leptoceridae) larval case.

632

Figure 19.271

Figure 19.266 Setodes incertus (Walker) (Leptoceridae) larval case. Figure 19.267 Leptocerus americanus (Banks) (Leptoceridae) lan/al case. Figure 19.268 Mystacides sp.(Leptoceridae) larval case.

Figure 19.269

Nectopsyche sp.(Leptoceridae) larval

case.

Figure 19.270 Ceraciea sp.(Leptoceridae) thorax, dorsal.

Figure 19.271 dorsal.

Nectopsyche sp.(Leptoceridae) thorax,

Chapter 19 Trichoptera

4'. 5(4').

5'.

6(5').

6'.

Mesonotum without pair of dark bars(Fig. 19.271); abdominal segments I-VII more slender, nearly parallel-sided, with gills single (Fig. 19.258) or absent Ventral apotome of head triangular (Fig. 19.260); tibia of each hind leg usually without apparent constriction (Fig. 19.262); pair of ventral, submesal bands of uniformly small spines beside anal opening (Fig. 19.259) or spines absent in this position, but no lateral patches of longer spines; slender case of plant fragments, fine sand (Fig. 10.182), and/or diatoms with usually 1 twig or conifer needle extending length of case and beyond 1 or both ends(Figs. 19.1, 19.269, 19.497); widespread Ventral apotome of head of mature larva rectangular (Fig. 19.261), if triangular, case a spiral of plant pieces (e.g.. Fig. 19.265); tibia of each hind leg with translucent constriction, apparently dividing it into 2 subequal parts (Figs. 19.263, 19.264); patch of longer spines lateral of each band of shorter submesal spines(Fig. 19.254) Mandibles strongly asymmetrical (Fig. 19.255); metathoracic legs each usually with close-set fringe of long hairs (Fig. 19.263); slender case a spiral of plant pieces (Figs. 19.2, 19.265, 19.503); widespread

633

5

Nectopsyche

6

Triaenodes

Mandibles only slightly asymmetrical(Fig. 19.256); metathoracic legs with only few, scattered, long hairs (Fig. 19.264); irregular case of plant and mineral materials, with twigs or conifer needles extending beyond ends

(Fig. 19.268); widespread

Mystacides

Limnephilidae^ 1.

Anterior margin of pronotum densely fringed with long hairs; dorsum of head flat and with 2 bands of dense scale hairs (Fig. 19.272); case tapered, depressed, made of transverse bits of wood and bark (Fig. 19.480);

r.

Anterior margin of pronotum and dorsum of head without dense hairs (Fig. 19.290) or hairs not of type or arrangement described above (Figs. 19.288, 19.289) Most abdominal gills single (Fig. 19.273) Most dorsal and ventral gills multiple (Fig. 19.274), lateral gills sometimes single (Fig. 19.332) Metanotal sal sclerites large, distance between sal sclerites no more than twice maximum dimension of single sal sclerite and .sal sclerites not fused (Fig. 19.276); case slender, straight, scarcely tapered tube, made of coarse rocks often with long plant material attached (Fig. 19.431);

West

2(1'). 2'. 3(2).

^

Cryptochia

West

18

Ecclisomyia

3'.

Metanotal sal sclerites small, distance between sal sclerites more than twice

4(3').

maximum dimension of single sal sclerite (Fig. 19.277), sal sclerites sometimes fused (Figs. 19.280) Each mesonotal plate wider than long, shorter mesally than laterally (Fig. 19.278); abdominal segment VIII with transverse posterodorsal line of slender, closely spaced setae (Fig. 19.275); case of short pieces of Sphagnum moss laid transversely (Fig. 19.341);

North

2 3

4

Phanocelia canadensis(Banks)'

'This genus is represented in North America by only one species(see also Table 19A). 'Larvae are unknown for Chilostigmodes(Alaska to Labrador), and Leptophylax (North Central states). These genera are represented in North America by only C. areolatus(Walker) and L. gracilis Banks, respectively.

634

4'.

5(4').

Chapter 19 Trichoptera

Mesonotal plates of varying width, but length nearly same mesally and laterally (Fig. 19.279); abdominal segment VIII with or without posterodorsal line of setae, but less dense than above, if present(Fig. 19.306) 1 or 2 sclerites adjacent to base of each lateral hump of abdominal segment I (Figs. 19.281-19.286; sclerites often only lightly pigmented and difficult to see, but distinguishable by relatively shinier surfaces)

5'.

No sclerites adjacent to lateral humps of abdominal segment I

6(5).

Large single sclerite at base of each lateral hump of abdominal segment I enclosing posterior half of hump and extending posterodorsad as irregular lobe (Fig. 19.282); case of leaves or bark formed into flattened tube with seams along narrow lateral flanges (Fig. 19.432); North,

6'.

1 or 2 small sclerites without irregular lobes at base of each lateral hump of abdominal segment I (Figs. 19.283, 19.284)

7'. 8(7).

Chyranda centralis(Banks)'

1 long sclerite at posterior edge of base of each lateral hump of abdominal segment I (Figs. 19.281, 19.284) 2 or more sclerites at base of each lateral hump on abdominal segment I (Figs. 19.283, 19.285) Sclerite at base of each lateral hump of abdominal segment I only half as tall as basal width of hump (Fig. 19.284); case smooth, thin-walled, nearly straight, little-tapered tube, made of irregularly arranged bark pieces (Fig. 19.433) or occasional flat rocks, rarely 3-sided; West

8'.

Sclerite at base of each lateral hump of abdominal segment I nearly as tall as basal width of hump (Fig. 19.281)

9(8').

Metanotal xul sclerites fused (Fig. 19.280); abdominal sternum II with chloride epithelium (in which case abdominal segment IX with only single seta on each side of dorsal sclerite, similar to Fig. 19.287, Hydatophylax argus, eastern North America) or without chloride epithelium (in which case abdominal segment IX with tuft of 3-6 setae, Fig. 19.287 inset, H. hesperus, western North America); case of wood or leaves in irregular outline

9'.

Metanotal 5al sclerites not fused although often contiguous (Fig. 19.277), abdominal sternum II without chloride epithelium and abdominal segment IX with only single seta on each side of dorsal sclerite (Fig. 19.287); case of twigs, gravel, or leaves, variously shaped (Fig. 19.435), occasionally 3-sided; Central, East, North

8

10

Homophylax

Hydatophylax

2 small ring sclerites posterodorsally at base of each lateral hump of abdominal segment I (Fig. 19.285); case tubular, slightly curved, tapered, made mostly of rocks with some small pieces of wood incorporated (Fig. 19.436)

10'. 11(10).

2 or more sclerites dissimilar in shape at base of each lateral hump (Fig. 19.283) Sclerotized parts more reddish brown; small secondary setae numerous on head; metanotum with setae scattered between sclerites bearing primary setal areas; abdominal segment VII usually lacking posterodorsal gills;

IF.

Sclerotized parts darker brown (tending to become reddish brown in

California

7

9

(Fig. 19.434); widespread except not Southwest

10(7').

6

13

West

7(6').

5

Pycnopsyche

11 12

Desmona

preserved material); secondary setae absent on head; metanotal setae between primary setal area sclerites reduced or absent; abdominal

segment VII usually with posterodorsal gills; West 'This genus is represented in North America by only one species(see also Table 19A).

Monophylax mono(Denning)'

Chapter 19 Trichoptera

12(10').

12'. 13(5').

1 or 2 small rounded posterodorsal sclerites and 1 long dorsal sclerite at base of each lateral hump of abdominal segment I (Fig. 19.283) or dorsal area at base of hump with several small discrete sclerites, 1 surrounding each seta; case rough, straight, untapered tube, made of rocks and wood fragments (Fig. 19.24), sometimes with trailing pieces or cases of other caddisflies attached (Fig. 19.437); North, West 1 irregular posterior sclerite and 1 irregular dorsal sclerotized area surrounding bases of 2-3 setae (Fig. 19.286) Anterior margin of pronotum with flat scale hairs, dorsum of head flat (Fig. 19.288); case slightly curved, coarse, made of minerals (Fig. 19.438); Northwest

635

Psychoglypha 15

Philocasca

13'.

Anterior margin of pronotum without flat scale hairs, setae normal; dorsum of head usually convex (Fig. 19.290, flat only in western Pseudostenophylax edwardsi. Fig. 19.292) 14(13'). Mesonotal ^al and sal distinct, separated by gaps free of setae (Figs. 19.293, 19.294) 14'. Mesonotal 5al and sal connected by continuous longitudinal band of setae on each side of meson (Fig. 19.291) 15(12',14). Pronotum covered with fine spines (Fig. 19.293 inset); case of plant and rock fragments (Fig. 19.439), sometimes with snail opercula;

Northwest(Arctic)

(Figs. 19.522a, 19.51 lb); Northwest

18(2'). 18'. 19(18).

16

C/osfoec« rf/s/Hwcta(Banks)'

Chilostigma itascae Wiggins'

Head and pronotum strongly inflated and with pebbled texture (Fig. 19.289); case smooth, tapered, curved, made of small rocks

(Fig. 19.340); Northwest 17'.

17

Mesonotal sclerites each wider than long (Fig. 19.295); case straight, made irregularly of small pieces of leaves and bark;

Minnesota 17(14').

15

Grensia praeterita (Walker)'

15'. Pronotum smooth and shiny, without fine spines (Fig. 19.294) 16(15'). Mesonotal sclerites each about as long as wide (Fig. 19.294); case of leaf pieces with wide flanges at each side of depressed tube 16'.

14

Ecclisocosmoecus scylla (Milne)'

Head and pronotum not unusually inflated (Fig. 19.290), although dorsum of head flat in western species (Fig. 19.292), sclerotized areas not pebbled; case smooth, tapered, curved, made of rocks (Figs. 10.184, 19.23), Central, East, Northwest Most gills with 2 or 3 branches, none with more than 4(Fig. 19.274) At least some gills with more than 4 branches Dorsum of head with 2 bands of contrasting color extending from

Pseudostenophylax 19 40

coronal suture to bases of mandibles(Fig. 19.296) and/or narrowed

19'.

20(19).

20'.

posterior portion of frontoclypeal apotome with 3 light areas: 1 along each side and 1 at posterior extremity (Fig. 19.297) Dorsum of head lacking bands or other well-defined, contrasting areas, usually uniform in color or with prominent light or dark spots only at points of muscle attachment(Fig. 19.311) Dorsum of head with prominent dark bands on light background, lateral bands extending from coronal suture to base of each mandible, median band on frontoclypeus (Fig. 19.296) Dorsum of head lacking dark lateral bands, but narrowed posterior part of frontoclypeus with 3 light areas: 1 along each side and 1 at posterior extremity (Fig. 19.297)

'This genus is represented in North America by only one species (see also Table 19A).

20

26

21

23

636

Chapter 19 Trichoptera

w:.A h

mi

Figure 19.273

Figure 19.274 Figure 19.278

Figure 19.279 Figure 19.272

Figure 19.276

Figure 19.280

Figure 19.277

Figure 19.275

dorsal hump

sclerite

lateral

hump sclerite sclerite

'// Figure 19.283 Figure 19.281

Figure 19.285

Figure 19.282

Figure 19.272 Cryptochia pilosa (Banks) (Umnephilidae) head and pronotum, dorsal. Figure 19.273 Pycnopsyche sp.(LImnephllidae) abdominal segment IV, left lateral. Figure 19.274 Limnephilus sp. (LImnephllidae) abdominal segment IV, left lateral. Figure 19.275 Phanocelia canadensis (Banks) (Umnephilidae) abdominal segments VIII and IX, dorsal. Figure 19.276 Ecctisomyia sp. (LImnephllidae) metanotum, dorsal.

Figure 19.277 Pycnopsyche sp.(LImnephllidae) metanotum, dorsal.

Figure 19.278 Phanocelia canadensis (Banks) (LImnephllidae) mesonotum, dorsal. Figure 19.279 Clostoeca disjuncta (Banks) (LImnephllidae) mesonotum, dorsal.

Figure 19.284

Figure 19.286

Figure 19.280 Hydatophylax sp.(LImnephllidae) metanotum, dorsal.

Figure 19.281 Pycnopsyche sp.(LImnephllidae) abdominal segment I, left lateral. Figure 19.282 Chyranda centralis (Banks) (LImnephllidae) abdominal segment I, left lateral. Figure 19.283 Psychogiypha sp.(LImnephllidae) abdominal segment I, left lateral. Figure 19.284 Homophylax sp.(LImnephllidae) abdominal segment I, left lateral. Figure 19.285 Desmona bethuia Denning (LImnephllidae) abdominal segment I, left lateral. Figure 19.286 Clostoeca disjuncta (Banks) (LImnephllidae) abdominal segment I, left lateral.

Chapter 19 Trichoptera

637

multiple setae

K si lit

Figure 19.289

Figure 19.288

Figure 19.287

Figure 19.290 I

I V

iV ////

Figure 19.291

fine spines.

Figure 19.292

Figure 19.293

Figure 19.295

Figure 19.294

) R.W. Holzenthal 2006

Figure 19.287 Pycnops/che sp.(Limnephilidae)

Figure 19.291

abdominal segments IX and X, left lateral; inset =

(Limnephiiidae) mesonotum, dorsal.

Ecclisocosmoecus scylla (Milne)

Hydatophylax hesperus (Banks)(Limnephiiidae), abdominal tergum IX lateral setae, left lateral. Figure 19.288 Philocasca rivularis W\gg\ns (Limnephiiidae) head and pronotum, dorsal. Figure 19.289 Ecc//socosmoecus scy/Za (Milne) (Limnephiiidae) head and pronotum, dorsal. Figure 19.290 Pseudostenophylax sparsus (Banks) (Limnephiiidae) head and pronotum, dorsal.

Figure 19.292 Pseudostenophylax edwardsi(Banks) (Limnephiiidae) head, left dorsolateral oblique. Figure 19.293 Grensia praeterita (Walker) (Limnephiiidae) thorax, dorsal; inset, microspines enlarged. Figure 19.294 Clostoeca disjuncta (Banks) (Limnephiiidae) thorax, dorsal. Figure 19.295 Chilostigma itascae Wiggins (Limnephiiidae) meso- and metanota, dorsal.

638

Chapter 19 Trichoptera

21(20).

Dark dorsal bands on head fused at junction of coronal and frontoclypeal sutures to form U-shaped marking, pronotum with narrow dark bands along anterior border and across dorsum (Fig. 19.298); case of leaf pieces arranged

21'.

Dark dorsal bands on head extended posterad beyond junction of coronal and frontoclypeal sutures to form V-shaped marking (Fig. 19.296), pronotal markings variable Chloride epithelia present dorsally (as well as dorsolaterally and ventrally) on several abdominal segments(Fig. 19.299); rough, tubular case of wood or leaf fragments, sometimes 3-sided, changed to fine gravel before pupation;

transversely or longitudinally (Fig. 19.440); North

22(21').

Nemotaulim hostilis(Hagen)'

22

Northwest Halesochila taylori(Banks)' Chloride epithelia absent dorsally (present ventrally and sometimes dorsolaterally) 25

22'. 23(20'). Abdominal sternum I usually with more than 100 setae overall, with setal areas merged (Fig. 19.300); small spines on head and pronotum (Fig. 19.297); short, stout setae on lateral sclerite of each anal proleg (Fig. 19.305); case cylindrical, made usually of irregular pieces of twigs and bark (Fig. 19.441b), sometimes small rocks(Fig. 19.441a); West 23'.

Abdominal sternum I with fewer than 100 setae overall, with setal

areas discrete; usually without spines on head and pronotum; usually without short, stout setae on lateral sclerite of anal proleg 24(23').

24'.

Chloride epithelia present dorsally, laterally, and ventrally on most abdominal segments (similar to Fig. 19.299); case of plant and rock materials (Fig. 19.343); North Chloride epithelia absent dorsally, may be present dorsolaterally but always present ventrally on most abdominal segments

25

Philarctiis bergrothi McLachlan'

26(19'). Femur of each hind leg with 2 major setae on ventral edge—these setae sometimes unequal in length (Fig. 19.303) 26'. Femur of each hind leg with more than 2 major setae on ventral

27'.

edge (Fig. 19.304) Pronotum, especially anterior margin (Fig. 19.307), and lateral sclerite of each anal proleg (Fig. 19.305) both with short, stout setae

28'. 29(27'). 29'.

27 36

28

Pronotum usually, lateral sclerite of anal proleg always, without short, stout setae

28(27).

Limnephilus (in part)

Mesothoracic femora each with 1 major seta 1/4 distance from base (remote from midpoint of femur) and 1 major seta near midpoint of femur (Fig. 19.302); case of fine sand, sedge seeds, or snail shells;

Northwest

27(26).

24

Asynarchus (in part)

25(22',24'). Mesothoracic femora each with 2 major setae situated near midpoint offemur (Fig. 19.301); cases of wide range of shapes and materials (Figs. 19.25, 19.442a-l9.442c); widespread except not deep Southeast (i.e., not east of Texas or south of Arkansas and South Carolina) 25'.

Clistoronia

Tibiae and tarsi of all legs each with dark, contrasting band (Fig. 19.303); case cylindrical, made of sand, twigs, and bark (Fig. 19.443); Central, North Tibiae and tarsi lacking dark bands; case smooth, cylindrical, mostly of small stones with wood fragments (Fig. 19.444); East Chloride epithelia present dorsally on at least some abdominal segments (as in Fig. 19.299) Chloride epithelia absent dorsally on abdominal segments

'This genus is represented in North America by only one species (see also Table 19A).

29

Glyphopsyche Frenesia 30

33

Chapter 19 Trichoptera

639

— dark band

Figure 19.297

Figure 19.296 chloride

f\

r\

f-\

epithelia

\ )

[ !

M

Figure 19.298

'iC

igure 19.299

i i/.\V' V

Figure 19.300 major seta

Figure 19.303 stout seta

Figure 19.301

major seta

Figure 19.302

stout seta

Figure 19.304

Figure 19.305

Figure 19.307

Figure 19.306

® R.W, Holzenthal 2006

Figure 19.296 Halesochila taylori(Banks) (Limnephilldae) head, dorsal. Figure 19.297 Clistoronia magnifica (Banks)

(Limnephilidae) head, dorsal; inset, microspines enlarged. Figure 19.298 Nemotaulius hostilis (Hagen) (Limnephilidae) head and pronotum, dorsal. Figure 19.299 Halesochila taylori(Banks)(Limnephilidae) abdomen, left lateral; insets, dorsal chloride epithelia. Figure 19.300 Clistoronia magnifica (Banks) (Limnephilidae) abdominal segment I, ventral. Figure 19.301 Limnephilus sp.(Limnephilidae) left mesothoracic femur, posterior (left lateral). Figure 19.302 Philarctus bergrothi MacLachlan (Limnephilidae) left mesothoracic femur, posterior (left lateral).

Figure 19.303 Glyphopsyche irrorata (Fabricius) (Limnephilidae) left metathoracic leg, posterior (left lateral). Figure 19.304 Dicosmoecus sp.(Limnephilidae) left metathoracic leg, posterior (left lateral). Figure 19.305 Clistoronia magnifica (Banks) (Limnephilidae) left lateral sclerite and anal proleg, left lateral.

Figure 19.306 Desmona bethuia Denning (Limnephilidae) abdominal segments VIII and IX, dorsal. Figure 19.307 Frenesia missa Milne (Limnephilidae) pronotum, dorsal.

640

Chapter 19 Trichoptera

30(29).

Metanotal sal with few setae, usually 2, and no scierite (Fig. 19.308); case smooth, cylindrical, made of long sedge or grass pieces; North 30'. Metanotal sal with more than 2 setae and with scierite (Fig. 19.311) 31(30'). Dorsum of head with numerous large spots often coalescing in places into diffuse blotches, or small discrete spots, especially on frontoclypeal

Arctopora 31

apotome (Fig. 19.311)

31'.

32

Dorsum of head with varied markings, but not spots; case of plant and

rock materials (Fig. 19.343); North

Asynarchus(in part)

32(31).

Anterolateral corners of pronotum each with small patch of spines (Figs. 19.87, 19.88, 19.311); case cylindrical, made of plant materials (Fig. 19.338), sometimes 3-sided; widespread 32'. Anterolateral corners of pronotum usually lacking patches of spines 33(29'). Prosternal horn extending beyond head capsule to mentum of labium (Figs. 19.89, 19.309); case cylindrical, made of transverse, narrow, projecting pieces of plant material (Fig. 19.445); widespread 33'. Prosternal horn extending only to distal edge of head capsule (Fig. 19.310) 34(33'). Chloride epithelia absent laterally, but present ventrally on abdominal sterna II-VII; mesonotal sal with single seta (Fig. 19.314); abdominal sternum 1 with single pair of sal setae (Fig. 19.316); case of pieces of sedge leaves arranged lengthwise and irregularly (Fig. 19.342); Northwest Terr.,

Yukon Terr.(Arctic tundra) 34'.

Anabolia 35

Platycentropus 34

Sphagnophylax meiops Wiggins and Winchester^

Chloride epithelia occasionally present laterally, always present ventrally on abdominal sterna II-VII; mesonotal sal usually with more than 1 seta (Fig. 19.313); abdominal sternum I usually with 2 or more sal setae (Fig. 19.315)

35

35(32',34') Dorsum of head light brownish yellow with numerous discrete, small,

dark spots (Fig. 19.312); case cylindrical, made of longitudinally arranged sedge or similar leaves (Fig. 19.337); North Grammotaulius 35'. Dorsum of head with varied markings, usually darker than above; cases of wide range of shapes and materials (Figs. 19.25, 19.442a-19.442c); widespread except not deep Southeast (i.e., not east of Texas or south of Arkansas and South Carolina) Limnephilus (in part) 36(26'). Tibiae with several pairs of stout, spur-like setae (Fig. 19.317) 37 36'. Tibiae each with only 1 pair of stout, spur-like setae at apex (Fig. 19.318) 38 37(36). Abdominal tergum I with transverse row of setae posterior to median dorsal hump (Fig. 19.319); scale hairs on dorsum of head (Fig. 19.325); abdominal sternum II with 2 chloride epithelia (Fig. 19.323); case irregularly outlined, made of small pebbles arranged in slightly curved and

flattened cylinder (Fig. 19.335); Northwest 37'.

38(36').

Allocosmoecus partitas Banks'

Abdominal tergum I usually lacking setae posterior to median hump (Fig. 19.320); dorsum of head without scale hairs; abdominal sternum II with single chloride epithelium (Fig. 19.324), 3 epithelia (smaller epithelia laterally), or without epithelia; case of final instar of fine gravel, slightly depressed (Figs. 19.336, 19.508), case of younger larvae with plant materials; West Metanotal .sal sclerites usually fused (Fig. 19.321), occasionally separated by small gap; case of hollow twig with ring of bark pieces anteriorly (Fig. 19.446a) or case entirely of wood fragments(Fig. 19.446b);

West 'This genus is represented in North America by only one species(see also Table 19A).

Dicosmoecus

Amphicosmoecus canax (Ross)'

Chapter 19 Trichoptera

641

Figure 19.308 muscle scars

spines

Figure 19.312

Figure 19.309 prosternal horn

Figure 19.313

Figure 19.310 Figure 19.311

Figure 19.314

fe "■■■■

/

Figure 19,315 Figure 19.316

Figure 19.308

Arctopora sp. (Limnephilidae)

Figure 19.313

© R.W. Holzenlhal 2006

Grammotaulius sp. (Limnephilidae)

metanotum, dorsal.

mesonotum, dorsal.

Figure 19.309 Platycentropus radiatus (Say) (Limnephilidae) head and prosternal horn, left lateral. Figure 19.310 Limnephilus sp. (Limnephilidae) head and prosternal horn, left lateral. Figure 19.311 Anabolla bimaculata (Walker) (Limnephilidae) head and thorax, dorsal. Figure 19.312 Grammotaulius sp. (Limnephilidae)

Figure 19.314 Sphagnophyiax meiops Wiggins and Winchester (Limnephilidae) mesonotum, dorsal. Figure 19.315 Grammotaulius sp. (Limnephilidae)

head, dorsal.

abdominal sternum I, ventral.

Figure 19.316 Sphagnophyiax meiops Wiggins and Winchester (Limnephilidae) abdominal sternum I, ventral.

642

Chapter 19 Trichoptera

38'. Metanotal sclerites clearly separate (Fig. 19.322) 39(38'). Pronotum anteriorly with erect submarginal setae widely spaced, especially mesally, and with some short, stout, marginal setae among fine marginal hairs (Fig. 19.328); abdominal tergum VII with a single long seta on either side of midline posterodorsally, sometimes each side with 1 or 2 much smaller setae (Fig. 19.330); lateral abdominal gills usually lacking from segment V,sometimes from IV or terminating with anterior position of segment IV; case of final instar made of stout pieces of wood (Fig. 19.447) or of fine rock fragments (Fig. 19.448), cases of earlier instars made of

39

pliable plant materials; Northwest

Eocosmoecus

39'.

Pronotum anteriorly with submarginal setae more closely spaced, especially submesally, and all of similar length and thickness, without short, stout setae among fine marginal hairs (Fig. 19.329); abdominal tergum VIl with 1-5 long setae on either side of midline posterodorsally (Fig. 19.331); lateral abdominal gills terminating with anterior position of segment V; case of pieces of wood and bark (Fig. 19.449); North, West 40(18'). Femora of meso- and metathoracic legs each with approximately 5 major setae along ventral edge (Fig. 19.326); case curved, scarcely tapered, made of wood, bark, twigs, and leaves (Figs. 19.339, 19.509, 19.510)

Onocosmoecus

or of sand; Central, East

40'. 41(40').

Ironoquia

Femora of meso- and usually metathoracic legs each with 2 major setae along ventral edge (Fig. 19.327)

41

Metanotum with all setae confined to primary sclerites (Fig. 19.333); case cylindrical, made of longitudinally arranged lengths of sedge leaves or of fragments of bark and leaves (Fig. 19.450); North, West

Lenarchus

41'.

Metanotum with at least few setae between primary sclerites (Fig. 19.334)

42(41').

Surface of head without short, fine, acuminate spines; West

42'.

Surface of head with short, fine, acuminate spines

43(42').

Pronotal surface minutely pebbled, without short spines;

Psychoronia 43

Southwest 43'.

42

Crenophylax sperryi(Banks)'

Pronotal surface with short spines, appearing as golden pubescence on surface held at oblique angle in transmitted light; North, West

Hesperophylax

Molannidae^

1.

Tarsal claw of each metathoracic leg curved, broad, setose, much shorter than tarsus (Fig. 19.344); Central, East

1'.

Tarsal claw of each metathoracic leg forming slender filament as

long as tarsus (Fig. 19.345); far Northwest

Molanna

Molannodes tinctus (Zetterstedt)'

Odontoceridae

1.

Prothoracic femora each about as broad as its tibia, prothoracic tibia about 4 times as long as its tarsus, single apical spur of prothoracic tibia broad, clasp-like (Fig. 19.346); case curved and tapered, made of rock fragments (Figs. 19.348, 19.479); Southern Appalachian

Mountains r.

Pseudogoera singularis Carpenter'

Prothoracic femora each distinctly broader than its tibia, prothoracic

tibia as long as its tarsus, both apical spurs of prothoracic tibia slender (Fig. 19.347)

2

'This genus is represented in North America by only one species(see also Table 19A). ' Larvae of Molannidae build depressed cases ofsand, each with lateral flanges and a hood over the anterior opening, completely hiding the larva beneath (Figs. 10.185, 19,6, 19.457, 19.458).

Chapter 19 Trichoptera

643

\\\\\ i" ;/ iM Spur-like setae

Figure 19.319 spur-like setae

setal row

I'l l I' li i ii,'

Figure 19.321

Figure 19.317 Figure 19.318 Figure 19.320 'J

Figure 19.322

Figure 19.323 chloride epithelium major

Figure 19.325 Figure 19.326

Figure 19.327

Figure 19.324

major seta

Figure 19.330 single-branched gill

WW/

\iljl

Figure 19.328 Figure 19.332

Figure 19.331

Figure 19.329 Figure 19.333

Figure 19.317 Dicosmoecus sp.(Limnephilidae) left metatlioracic tibia and tarsus, posterior (left lateral). Figure 19.318 Onocosmoecus unicolor (Banks) (Limnephilidae) left metathoracic tibia and tarsus,

posterior (left lateral). Figure 19.319 Allocosmoecus partitus Banks (Limnephilidae) abdominal tergum I, dorsal. Figure 19.320 Dicosmoecus sp.(Limnephilidae) abdominal tergum I, dorsal. Figure 19.321 Amphicosmoecus canax (Ross) (Limnephilidae) metanotum, dorsal. Figure 19.322 Onocosmoecus unicolor (Banks) (Limnephilidae) metanotum, dorsal. Figure 19.323 Allocosmoecus partitus Banks (Limnephilidae) abdominal sternum II, ventral. Figure 19.324 Dicosmoecus sp.(Limnephilidae)

Figure 19.334

Figure 19.326 Ironoqula sp.(Limnephilidae) left metathoracic leg, posterior (left lateral). Figure 19.327 Hesperophylax sp.(Limnephilidae) left metathoracic femur, posterior (left lateral). Figure 19.328 Eocosmoecus frontalls (Banks) (Limnephilidae) pronotum, dorsal. Figure 19.329 Onocosmoecus unicolor (Banks) (Limnephilidae) pronotum, dorsal. Figure 19.330 Eocosmoecus schmldl(Wiggins) (Limnephilidae) abdominal segment VII, dorsal. Figure 19.331 Onocosmoecus unicolor (Banks) (Limnephilidae) abdominal segment VII, dorsal. Figure 19.332 Psychoronia costalls (Banks) (Limnephilidae) abdominal segment II, left lateral. Figure 19.333 Lenarchus sp.(Limnephilidae) metanotum, dorsal.

abdominal sternum II, ventral.

Figure 19.334 Hesperophylax sp.(Limnephilidae)

Figure 19.325 Allocosmoecus partitus Banks (Limnephilidae) scale hairs on dorsum of head.

metanotum, dorsal.

644

Chapter 19 Trichoptera

Figure 19.335

Figure 19.336 Figure 19.337

Figure 19.341

Figure 19.339 Figure 19.338

Figure 19.342

Figure 19.343

Figure 19.340 © R.W. Holzenthal 2006

Figure 19.335 Allocosmoecus partitas Banks (Limnephilidae) larval case. Figure 19.336 Dicosmoecus sp.(Limnephilidae) larval case.

Figure 19.337

Grammotaulius sp.(Limnephilidae)

larval case.

Figure 19.338 Anabolia bimaculata (Walker) (Limnephilidae) larval case. Figure 19.339 Ironoquia sp.(Limnephilidae) larval case.

Figure 19.340 Ecclisocosmoecus scylla (Milne) (Limnephilidae) larval case. Figure 19.341 Phanocelia canadensis (Banks) (Limnephilidae) larval case. Figure 19.342 Sphagnophyiax meiops Wiggins and Winchester (Limnephilidae) larval case. Figure 19.343 Asynarchus montanus (Banks) (Limnephilidae) larval case.

Chapter 19 Trichoptera

645

Figure 19.346

femur

filament

tarsal

Figure 19.347 Figure 19.345

Figure 19.344

Figure 19.349 Figure 19.348

Figure 19.350 spines

%;)\

'If I

Figure 19.351

Figure 19.354

Figure 19.352

© R.W. Holzenthal 2006

Figure 19.353

Figure 19.355

Figure 19.344 Molanna tryphena Betten (Molannidae) right metathoracic tarsus, posterior (right iaterai). Figure 19.345 Molannodes tinctus (Zetterstedt) (Moiannidae) right metathoracic tarsus, posterior (right lateral).

Figure 19.346 Pseudogoera singularis Carpenter (Odontoceridae) left prothoracic leg, posterior (left lateral). Figure 19.347 Psilotreta sp. (Odontoceridae) left prothoracic leg, posterior (left lateral). Figure 19.348 Pseudogoera singularis Carpenter (Odontoceridae) larval case. Figure 19.349 Psilotreta sp.(Odontoceridae) larval case.

Figure 19.350 Psilotreta sp.(Odontoceridae) pronotum, left lateral. Figure 19.351 Namamyia plutonis Banks (Odontoceridae) pronotum, left lateral. Figure 19.352 Parthlna vierra Denning (Odontoceridae) left anal proleg, left lateral. Figure 19.353 Psilotreta sp.(Odontoceridae) left anal proleg, left lateral. Figure 19.354 Marilia sp.(Odontoceridae) thorax, dorsal.

Figure 19.355 Nerophilus callfornicus (Hagen) (Odontoceridae) thorax, dorsal.

646

Chapter 19 Trichoptera

2(1'). 2'.

Anterolateral corner of pronotum produced, sharply pointed (Fig. 19.350) Anterolateral corner of pronotum not produced, rounded (Fig. 19.351);

3

case curved, slightly tapered, made of sand (Figs. 10.186, 19.471-19.473)

4

3(2).

Ventral apotome of head long, completely separating genae (Fig. 19.356); claw of each anal proleg stout, both claw and lateral sclerite with straight spines as well as setae (Fig. 19.352); case of fine sand grains with

3'.

Ventral apotome of head short, separating genae only in anterior l/3rd (Fig. 19.357); claw of each anal proleg more slender, claw and lateral sclerite with setae only

silken exterior (Fig. 19.478); West

4(2').

Parthina

(Fig. 19.353); case slightly curved and tapered, made of coarse and fine rock fragments, very sturdy (Figs. 19.7, 19.349, 19.465); East Mesonotal plates each subdivided into 3 sclerites; metanotal sa\ sclerites large, subrectangular, contiguous mesally (Fig. 19.354); Central, Southwest, Vermont

4'.

5(4').

Marilia

Mesonotal plates undivided; metanotal sa\ sclerites small, oval, separated by distance equal to, or greater than, greatest width of 1 of them (Fig. 19.355) Abdominal sternum I with 2 clusters of gill filaments, 2 pairs of setae

(Fig. 19.358); California, Oregon 5'.

Psilotreta

5

Nerophilus californicus(Hagen)'

Abdominal sternum I without gills, with many setae (Fig. 19.359); California, Oregon Namamyia /j/Hton/s Banks'

Philopotamidae^-^ 1.

r.

Anterior margin of frontoclypeus with prominent notch asymmetrically right of midline (Fig. 19.360); foretrochantin small, scarcely projecting (Fig. 19.364) and prothoracic coxae with long, slender, subapical, seta-bearing process(Fig. 19.364); head with seta no. 18 at level of posterior point of ventral apotome (Fig. 19.365); widespread Anterior margin of frontoclypeus variable, prominently notched

Chimarra

(similar to Fig. 19.360), slightly sinuous (Fig. 19.361), or completely

2(1'). 2'.

symmetrical(Fig. 19.362); foretrochantin small as above or elongate, fingerlike (Fig. 19.363); prothoracic coxae without long subapical processes (Fig. 19.363); head with seta no. 18 approximately halfway between posterior edge of ventral apotome and occipital foramen (Fig. 19.366) Anterior margin of frontoclypeus slightly (Fig. 19.361) to markedly (as in Fig. 19.360) asymmetrical; foretrochantin projecting, finger-like (Fig. 19.363) Anterior margin of frontoclypeus evenly convex, symmetrical(Fig. 19.362); foretrochantin small, scarcely projecting (as in Fig. 19.364); larva as in Fig. 10.166; widespread

3(2). 3'.

2

3

Wormaldia

Anterior margin of frontoclypeus slightly asymmetrical(Fig. 19.361); widespread Dolophilodes Anterior margin of frontoclypeus markedly asymmetrical (similar to Fig. 19.360); Southeast(mountains) Fumonta major(Banks)'

Phryganeidae 1. Genae of head almost completely separated by ventral apotome (Fig. 19.368); case entirely of plant fragments r. Genae of head mostly contiguous ventrally, separated anteriorly by tiny ventral apotome (Fig. 19.367); case of plant and mineral fragments, mineral fragments mostly anterior and ventral (Figs. 19.376, 19.474), pupal case

entirely of mica-like fragments; California, Oregon

2

Yphria californica (Banks)'

'This genus is represented in North America by only one species(see also Table 19A). ' The larva is unknown for Sisko(West), represented in North America by S. sisko (Ross)and Sisko oregona (Denning). ' The finger-like retreats of Philopotamidae larvae are each fastened to the underside of a rock, often in groups of retreats. The net has the finest mesh of any known caddisfly retreat and is distended by the flow of water, collapsing when removed from water. Nets range in size from 25 to 60 mm long and from 2.5 to 5 mm wide.

Chapter 19 Trichoptera

Mi 'V

647

)/'V

ventral

apotome

Figure 19.359

Figure 19.358

Figure 19.356

Figure 19.357

Figure 19.360

Figure 19.362

Figure 19.361 foretrochantin

no. 18

foretrochantin

Figure 19.363

Figure 19.365 coxa! process

) R.W.(lolxenthal 2006

Figure 19.364 Figure 19.366

Figure 19.356 Parthina vierra Denning (Odontoceridae) head, ventral. Figure 19.357 Psilotreta sp.(Odontoceridae) head, ventral.

Figure 19.358 Nerophilus californicus (Hagen) (Odontoceridae) abdominal sternum I, ventral. Figure 19.359 Namamyia plutonis Banks (Odontoceridae) abdominal sternum I, ventral. Figure 19.360 Chimarra sp.(Philopotamidae) head,

head, dorsal.

Figure 19.363 Dolophilodes sp.(Philopotamidae) right foretrochantin and prothoracic coxa, right lateral. Figure 19.364 Chimarra sp.(Philopotamidae) right foretrochantin and prothoracic coxa, right lateral. Figure 19.365 Chimarra sp.(Philopotamidae) head, ventral.

Figure 19.366 head, ventral.

dorsal.

Figure 19.361

Figure 19.362 Wormaldia sp.(Philopotamidae) head, dorsal.

Dolophilodes sp.(Philopotamidae)

Dolophilodes sp.(Philopotamidae)

648

Chapter 19 Trichoptera

ventral

apotome ventral

antenna

apotome

I

Figure 19.367

stemmata stemellum

I

Figure 19.369

Figure 19.370

Figure 19.368 Figure 19.371

Figure 19.372

Figure 19.373

Figure 19.374 C) R.W. Holzenthal 2006

Figure 19.375

Figure 19.367 Yphria californica (Banks) (Phryganeidae) head, ventral. Figure 19.368 Agrypnia vestita (Walker) (Phryganeidae) head and prothorax, ventral. Figure 19.369 Beothukus complicatus (Banks) (Phryganeidae) right anterolaterai corner of head, dorsai.

Figure 19.370 Oligostomis ocelligera (Walker) (Phryganeidae) right anterolaterai corner of head, dorsai.

Figure 19.371

Fabria inornata (Banks)(Phryganeidae)

pronotum, dorsal.

Figure 19.372 Ptilostomis sp.(Phryganeidae) thorax, dorsai.

Figure 19.373 Hagenella canadensis (Banks) (Phryganeidae) head and thorax, dorsal. Figure 19.374 Banksiola dossuaria (Say) (Phryganeidae) head and thorax, dorsal. Figure 19.375 Oligostomis oceiligera (Walker) (Phryganeidae) mesonotum, dorsal.

Chapter 19 Trichoptera

2(1).

2'.

649

Mesonotal ral sclerites several times larger than sa3 sclerites, each with sal seta near its anterior edge (Fig. 19.375); case of ring construction (Fig. 19.380)

3

Mesonotal ^nl sclerites absent(Fig. 19.372) or much smaller than sa3 sclerites (Fig. 19.373) and each with its 5ul seta centrally located; case generally of ring (Figs. 19.12, 19.380) or spiral (Figs. 19.377-19.379) construction, slightly curved or straight, respectively

4

3(2).

Antennae each as long as width of pigmented area of clustered stemmata (Fig. 19.369); case straight, with plant materials arranged in discrete rings

3'.

Antennae much shorter than width of pigmented area of clustered stemmata (Fig. 19.370); case slightly curved, with plant materials arranged in discrete rings or bands (Fig. 19.454); East except not deep Southeast

4(2').

Head and pronotum uniformly light brown except for darker muscle scars on head (Fig. 19.373); case of ring construction (Fig. 19.496); Northcentral, Northeast Hagenella canadensis(Banks) Head and pronotum with distinct, dark bands (Fig. 19.374) 5

or bands(Fig. 19.453); Northcentral, Northeast

4'.

5(4'). 5'. 6(5).

6'.

7(5'). 7'.

8(7).

8'.

Beothukus complicatus(Banks)'

Ventral combs of prothoracic coxae conspicuous, their individual teeth evident at magnification of 50X (Fig. 19.383) Ventral combs of prothoracic coxae small, each comb appearing as tiny raised points at 50X magnification (Fig. 19.382) Prothoracic sternellum usually present(Fig. 19.368); ventral combs of mesothoracic coxae with basal axes both transverse and parallel to long axis of coxa (Fig. 19.384); case usually of spiral construction (Figs. 10.187, 19.495); widespread

Oligostomis

6 7

Agrypnia

Prothoracic sternellum absent; ventral combs of mesothoracic coxae with basal axes only transverse to long axis of coxa (Fig. 19.381); case of spiral

construction (Figs. 19.14, 19.377); widespread Meso- and metanota with pair oflongitudinal, irregular, dark bands (Fig. 19.374); case of spiral construction

Phryganea 8

Meso- and metanota nearly uniform in color (Fig. 19.372); case

variously constructed 9 Abdominal segments VI and VII with anterodorsal gills, segment VII without posteroventral gills (Fig. 19.385); case often with pieces of plant material trailing posteriorly (Figs. 19.13, 19.378); widespread Banksiola Abdominal segments VI and VII without anterodorsal gills, segment VII with posteroventral gills (Fig. 19.386); case as in Figure 19.377, without trailing ends; Alaska,

Yukon Territory

Oligotricha lapponica (Hagen)'

9(7').

Pronotum with dark line along anterior margin, without dark, central, transverse markings (Fig. 19.371); case of spiral construction but with trailing ends of plant fragments giving bushy appearance (Figs. 19.15, 19.379); North Fabria inornata (Banks)'

9'.

Pronotum without dark line along anterior margin, with dark transverse

markings near center of each sclerite (Fig. 19.372); case of ring construction without trailing ends (Figs. 19.12, 19.380); widespread except not Southwest

Ptilostomis

'This genus is represented in North Atnerica by only one species(see also Table 19A). ' The larva keyed here has not been positively associated, but it is similar to those of European species of Hagenella and is "almost certainly H. canadensis"(Wiggins 1998),

650

Chapter 19 Trichoptera

Figure 19.376

Figure 19.377 Figure 19.378 Figure 19.379

Figure 19.380

coxa comb

Figure 19.381

Figure 19.382

Figure 19.383

Figure 19.384

anterodorsal

_ posteroventral

" gill Figure 19.386

Figure 19.385

©R.W. Holzenthal 2018

Figure 19.376 Yphria californica (Banks) (Phryganeidae) larval case. Figure 19.377 Phryganea sp.(Phryganeidae)

Figure 19.382 Banksiola dossuaria (Say) (Phryganeidae) left prothoracic coxa, ventral. Figure 19.383 Phryganea sp.(Phryganeidae) left prothoracic coxa, ventral.

larval case.

Figure 19.378 Banksiola dossuaria (Say) (Phryganeidae) larval case. Figure 19.379 Fabria inornata (Banks)(Phryganeidae) larval case.

Figure 19.384 Agrypnia vestita (Walker) (Phryganeidae) left mesothoracic coxa, ventral. Figure 19.385 Banksiola dossuaria (Say) (Phryganeidae) abdominal segments VI and VII, left

Figure 19.380 Ptilostomis sp.(Phryganeidae) larval

lateral.

case.

Figure 19.386 Oiigotricha iapponica (Hagen) (Phryganeidae) abdominal segments VI and VII, left

Figure 19.381

Phryganea sp.(Phryganeidae) left

mesothoracic coxa, ventral.

lateral.

Chapter 19 Trichoptera

651

Figure 19,391 accessory

Figure 19.387

Figure 19.388

spine spines

Figure 19.389

Figure 19.390

tarsus

Figure 19.392

bristle

Figure 19.395

Figure 19.396

Figure 19.394

sa1

Figure 19.393 sa2

Figure 19.397

© R.W. Holzenthal 2006

Figure 19.398

Figure 19.387 Polyplectropus sp.(Polycentropodidae)

Figure 19.394 Nyctiophylax sp.(Polycentropodidae)

left anal claw, left lateral.

pronotum, left dorsolateral oblique. Figure 19.395 Polycentropus sp. (Polycentropodidae) left prothoracic tibia, tarsus, and tarsal claw, posterior

Figure 19.388 Nyctiophylax sp.(Polycentropodidae) left anal claw, left lateral.

Figure 19.389 Plectrocnemia sp.(Polycentropodidae) left anal claw, left lateral.

Figure 19.390 Holocentropus sp.(Polycentropodidae) left anal claw, left lateral.

Figure 19.391 Neureclipsis sp.(Polycentropodidae) left anal proleg and claw, left lateral; Inset, anal claw enlarged. Figure 19.392 Cernotina spicata Ross (Polycentropodidae) left anal proleg and claw, left lateral; Inset, dorsal region of base of claw. Figure 19.393 Cyrnellus fraternus (Banks) (Polycentropodidae) left anal proleg and claw, left lateral; Inset, dorsal region of base of claw.

(left lateral). Figure 19.396

Cernotina spicata Ross (Polycentropodidae) left prothoracic tibia, tarsus, and tarsal claw, posterior (left lateral). Figure 19.397 Polycentropus sp.(Polycentropodidae) thorax, dorsal.

Figure 19.398 Cyrneiius fraternus (Banks) (Polycentropodidae) thorax, dorsal.

652

Chapter 19 Trichoptera

Polycentropodidae 1. Anal claws each with 6 or fewer conspicuous teeth along ventral, concave margin (Figs. 19.387, 19.388) r.

2

Anal claws without ventral teeth (Figs. 19.389, 19.390, 19.392, 19.393) or each with 10 or more tiny spines(Fig. 19.391) along ventral, concave margin

3

2(1).

Teeth on each anal claw much shorter than apical hook, dorsal accessory spine present (Figs. 19.65, 19.388); pronotum with short, stout bristle near each lateral margin (Fig. 19.394); silken retreat forming rectangular tent over depression in wood or rock, open at two ends; widespread except not Southwest

2'.

Teeth on each anal claw almost as long as apical hook, dorsal accessory spine absent(Fig. 19.387); pronotum without short, lateral bristles; silken retreat similar to that of Nyctiophylax-, Southwest

3(1').

3'.

4(3').

4'.

Basal segment of each anal proleg about as long as distal segment and with only 2 or 3 apicoventral setae (Fig. 19.391); anal claw with many tiny spines along ventral, concave margin (Fig. 19.391 inset); larva as in Fig. 10.167; silken retreat trumpet-shaped, recurved, up to 12 cm long, slender basally and broad apically, with 3-4 cm opening facing current(Fig. 19.37); widespread except not Southwest Basal segment of each anal proleg obviously longer than distal segment in mature specimens and with many setae scattered over most of its ventral and dorsal surfaces (Figs. 19.392, 19.393); anal claw without tiny ventral spines (Figs.19.392, 19.393) Dorsal region between anal claw and sclerite of distal segment of each anal proleg with 2 dark bands contiguous mesally (Fig. 19.392 inset); meso- and metanotal sal setae short, not more than l/3rd as long as longest sal setae (Fig. 19.397) Dorsal region between anal claw and sclerite of distal segment of each anal proleg with 2 dark bands completely separated mesally (Fig. 19.393 inset); meso- and metanotal ^al setae about as long as longest sal setae (Fig. 19.398); silken retreat similar to that of Nyctiophylax, but more nearly circular; Central,

East

Polyplectropus

Neureclipsis

4

5

Cyrnellusfraternus(Banks)'

5(4).

Prothoracic tarsi broad and only 1/2 as long as prothoracic tibiae (Fig. 19.395); silken retreat bag-like structure expanded by current;

5'.

Prothoracic tarsi narrow and at least 2/3rds as long as prothoracic tibiae (Fig. 19.396)

widespread

6(5').

Nyctiophylax

Polycentwpus'° 6

Anal claws obtusely curved (Fig. 19.389); capture net loosely constructed, flat, spider-like, with funnel-shaped silken retreat in middle, surrounded by

maze of silken threads; widespread except not Southwest 6'.

Anal claws curved approximately 90° (Fig. 19.392)

7(6').

Anal claws each with 2 or 3 dorsal accessory spines(Fig. 19.390, spines tiny and difficult to see with dissecting microscope, sometimes broken); capture net bowlor funnel-shaped with tubular retreat perpendicular to it in middle and supported

by maze of silken threads in surrounding vegetation; Central, North

Plectrocnemia^^ 7

Holocentropus^°

'This genus is represented in North America by only one species(see also Table 19A), Considerable caution must be exercised when distinguishing among Cernotina, Holocentropus, Plectrocnemia, and Pulycentropus because the larvae of so few species are known (Wiggins 1996).

Chapter 19 Trichoptera

653

Figure 19.401 Figure 19.400

Figure 19.399

submental

Figure 19.402

sderite

protuberance

submental sclerite

Figure 19.404

Figure 19.403

ventral

Figure 19.405

apotome

Figure 19.406

© R.W. Holzenthal 2006

Figure 19.407

Figure 19.399 Psychomyia flavida Hagen (Psychomyiidae) left anal claw, left lateral. Figure 19.400 Tinodes sp.(Psychomyiidae) left anal claw, left lateral.

Figure 19.401 Lype diversa (Banks)(Psychomyiidae) mandibles, dorsal.

Figure 19.402 Tinodes sp.(Psychomyiidae) mandibles, dorsal.

Figure 19.403 Psychomyia flavida Hagen (Psychomyiidae) head, ventral.

Figure 19.404 Tinodes sp.(Psychomyiidae) head, ventral.

Figure 19.405 Tinodes sp.(Psychomyiidae) ventral apotome and submental sclerites, ventral. Figure 19.406 Lype diversa (Banks)(Psychomyiidae) ventral apotome and submental sclerites, ventral. Figure 19.407 Himaiopsyche phryganea (Ross) (Rhyacophilidae) thorax and abdominal segments I and II (distal portions of thoracic legs omitted), left lateral.

654

Chapter 19 Trichoptera

mesepisternum

0 oO o

Figure 19.409

S'OO

0 QQ O

" o 0 Oo

booO

O o

Figure 19.410

Figure 19.408

Figure 19.411

Figure 19.412 Figure 19.414

Figure 19.417

Figure 19.413

Figure 19.415

Figure 19.416

carna

Figure 19.418

© R.W. Holzcnthal 2006

Figure 19.419

Figure 19.420

Figure 19.408 Rosslana montana Denning (Rossianidae) head and thorax, dorsal. Figure 19.409 Goereilla baumanni Denning (Rossianidae) mesothorax, left lateral. Figure 19.410 Rossiana montana Denning (Rossianidae) mesothorax, left lateral. Figure 19.411 Rossiana montana Denning (Rossianidae) larval case. Figure 19.412 Gumaga nigricola (McLachlan) (Sericostomatidae) larval case. Figure 19.413 Agarodes libalis Ross and Scott (Sericostomatidae) pronotum, left lateral. Figure 19.414 Gumaga nigricola (McLachlan) (Sericostomatidae) pronotum, left lateral.

Figure 19.415 Fattigia pele (Ross)(Sericostomatidae) posterior edge of abdominal segment iX, dorsal. Figure 19.416 Agarodes sp.(Sericostomatidae) posterior edge of abdominal segment IX, dorsal. Figure 19.417 Agarodes libalis Ross and Scott (Sericostomatidae) metanotum, dorsal. Figure 19.418 Gumaga nigricola (MacLachlan) (Sericostomatidae) metanotum, dorsal. Figure 19.419 Fattigia pele (Ross)(Sericostomatidae) head, left lateral.

Figure 19.420 Agarodes libalis Ross and Scott (Sericostomatidae) head, left lateral.

Chapter 19 Trichoptera

655

\ J] j '1 ,emargination TT

emargination

Figure 19.423

T"

Jc>. Figure 19.424

Figure 19.422

Figure 19.421

lobate

foretrochantin

Figure 19.426

Figure 19.425

darkened

Figure 19.427

posterolateral

darkened posterolateral

Figure 19.428

corner

© R.W, Holzcnthai 2006

thorax, dorsal.

Figure 19.426 Sericostriata surdickae Wiggins, Weaver, and Unzicker (Uenoidae) pro- and mesonota,

Figure 19.422 Neophylax sp.(Thremmatidae) thorax,

dorsal.

dorsal.

Figure 19.427 Neothremma alicia Dodds and Hisaw (Uenoidae) pro- and mesonota, dorsal. Figure 19.428 Xiphocentron sp.(Xiphocentronidae)

Figure 19.421

Oligophlebodes sp.(Thremmatidae)

Figure 19.423 Neothremma sp.(Uenoidae) abdominal segment II, left lateral. Figure 19.424 Farula sp.(Uenoidae) abdominal segment II, left lateral. Figure 19.425 Farula jewetti Denning (Uenoidae) pro- and mesonota, dorsal.

head and thorax, left lateral.

656

7'.

Chapter 19 Trichoptera

Anal claws each with only 1 dorsal accessory spine (Fig. 19.392); capture net silken, tent-like, covering depression in wood or rock, with flared opening

at each end; Central, East

Cernotina^°

Psychomyiidae 1. Anal claw with 3 or 4 conspicuous teeth along ventral, concave margin (Fig. 19.399)

2

1'.

Anal claw without teeth on ventral, concave margin (Fig. 19.400)

3

2(1).

Paired submental sclerites on ventral surface of labium each longer than broad (Fig. 19.403); larva as in Fig. 10.168; retreat on rock, silken meandering tube several centimeters long and covered with sand (Fig. 19.36); widespread Paired submental sclerites on ventral surface of labium broader than long (Fig. 19.404); retreat on rock, similar to that of Psychomyia, up to 22 mm

2'.

long and 2-3 mm wide; Arkansas, Missouri(Ozarks) 3(1').

Paduniella nearctica Flint'

Dorsolateral edge of each mandible without protuberance, but with small dorsal condyle close to base, lateral setae about l/3rd of distance from base (Fig. 19.401); submental sclerites each about l/3rd as long as wide (Fig. 19.406); silken retreat with slightly arched roof, camouflaged with detrital inclusions,

in groove of submerged wood; Central, East 3'.

Psychomyia

Lype diversa (Banks)'

Dorsolateral edge of each mandible with rounded protuberance, dorsal condyle sometimes prominent, lateral setae about midway from base (Fig. 19.402); submental sclerites each about half as long as wide (Fig. 19.405); silken retreat flattened, elongate, covered with sand, attached usually to rock; West

Tinodes

Rhyacophilidae^^ 1.

Dense tuft of stout gills on each side of meso- and metathorax and abdominal segments I-VIII (Figs. 10.174, 19.407); final instar

1'.

Tufts of gills absent or not as dense or not on as many abdominal segments as above (Figs. 10.173, 19.57); final instar larva less than 25 mm long; widespread

larva up to 32 mm long; Pacific states

Himalopsychephryganea (Ross)'

Rhyacophila

Rossianidae

1.

Dorsum of head concave, with posterolateral flanges (Fig. 19.408); pronotum with pair of concavities (Fig. 19.408); mesepisterna each granulate but lacking spiny lobe (Fig. 19.410); case curved, scarcely tapered, made of coarse rock fragments (Figs. 10.188, 19.26, 19.411, 19.464);

Northwest r.

Rossiana montana Denning'

Dorsum of head rounded,lacking flanges; pronotum rounded; mesepisterna each with short, rounded, spiny prominence (Fig. 19.409); case curved, tapered, relatively smooth, made of sand with few small

pieces of detritus (Fig. 19.455); Northwest

Goereilla baumanni Denning'

'This genus is represented in North America by only one species(see also Table 19A). Considerable caution must be exercised when distinguishing among Cernotina, Holocentropus, Plectrocnemia, and Polycentropus because the larvae of so few species are known (Wiggins 1996).

"Larvae are free-living, constructing neither cases nor retreats, until pupation.

a Figure 19.429

Figure 19.430 Figure 19.433

Figure 19.432 Figure 19.431

Figure 19.438

Figure 19.438

Figure 19.434

Figure 19.437

Figure 19.435 )James C.(Skip) Hodges, Jr. 2018

Figure 19.429 Moselyana comosa Denn'mg (Apataniidae) larval case, left lateral. Figure 19.430 A//omy;a scoff/(Wiggins)(Apataniidae)

Figure 19.434 Hydatophylax sp.(Limnephilidae) larval case, ventral. Figure 19.435 Pycnopsyche sp.(Limnephilidae) larval

larval case, left lateral.

case, left lateral

Figure 19.431 Ecclisomyia sp.(Limnephilidae) larval

Figure 19.436 Desmona bethula Denning (Limneptiilidae) larval case, left lateral. Figure 19.437 Psychoglypha mazamae Denning

case, left lateral.

Figure 19.432 Chyranda centralis (Banks) (Limnephilidae) larval case, ventral. Figure 19.433 Homophylax andax Ross (Limnephilidae) larval case, left lateral.

(Limnephilidae) larval case, left lateral. Figure 19.438 Philocasca demita Ross (Limnephilidae) larval case, left lateral. 657

Figure 19.439

Figure 19.440

Figure 19.441a Figure 19.441b Figure 19.442a

Figure 19.442b

Figure 19.443 Figure 19.444 Figure 19.442c © James C.(Skip) Hodges,Jr. 2018

Figure 19.439 Grensia praeterita (Walker) (Limnephilldae) larval case, left lateral. Figure 19.440 Nemotaulius hostilis (Hagen) (Limnephilldae) larval case, ventral. Figure 19.441a Glistoronia magnifica (Banks) (Limnephiiidae) larval case, left lateral. Rgure 19.441b Glistoronia magnifica (Banks) (Limnephiiidae) larval case, left lateral. Figure 19.442a Limnepfiiius externus Hagen (Limnephiiidae) larval case, left lateral. 658

Figure 19.442b Limnephiius externus Hagen (Limnephilldae) larval case, left lateral.

Figure 19.442c Limnephiius externus Hagen (Limnephiiidae) larval case, ventral.

Figure 19.443 Giyphopsyche irrorata (Fabricius) (Limnephiiidae) larval case, left lateral.

Figure 19.444 Frenesia missa (Milne)(Limnephiiidae) larval case, left lateral.

Figure 19.445 Piatycentropus radiatus (Say) (Limnephiiidae) larval case, left lateral.

Figure 19.446a

Figure 19.446b

Fjgu^e 19.447

Figure 19.449 Figure 19.450

4

il

Figure 19.451

Figure 19.452a

Figure 19.452b Figure 19.453

Figure 19.454 © James C.(Skip) Hodges,Jr. 2018

Figure 19.446a Amphicosmoecus canax(Ross) (Limnephilidae) larval case, left lateral. Figure 19.446b Amphicosmoecus canax(Ross) (Limnephilidae) iarvai case, left lateral. Figure 19.447 Eocosmoecus frontalis (Banks) (Limnephilidae) larval case, left lateral. Figure 19.448 Eocosmoecus schmidi(Wiggins) (Limnephilidae) larval case, left ventrolateral. Figure 19.449 Onocosmoecus unicolor (Banks) (Limnephilidae) larval case, left lateral. Figure 19.450 Lenarchus vastus (Hagen) (Limnephilidae) larval case, left lateral. Figure 19.451 Psychoronia costalis (Banks) (Limnephilidae) iarvai case, left lateral.

Figure 19.452a Hesperophylax sp.(Limnephilidae) larval case, left lateral.

Figure 19.452b Hesperophylax sp.(Limnephilidae) larval case, left lateral.

Figure 19.453 Beothukus complicatus (Banks) (Phryganeidae) larval case, left lateral. Figure 19.454 Oligostomis pardalis (Walker) (Phryganeidae) larval case, left lateral. Figure 19.455 Goereilla baumanni Denning (Rossianidae) larval case, left lateral. Figure 19.456 Sericostriata surdickae Wiggins, Weaver, and Unzicker (Uenoidae) iarvai case, left lateral.

659

660

Chapter 19 Trichoptera

Sericostomatidae

1.

Anterolateral corners of pronotum each acute, projecting (Fig. 19.413); metanotal sal with many setae on transverse sclerites (Fig. 19.417); case curved, tapered, relatively smooth, made of fine sand (Figs. 10.189, 19.10, 19.475, 19.476)

1'.

Anterolateral corners of pronotum each rounded, not projecting (Fig. 19.414); each metanotal sal with single seta and no sclerite (Fig. 19.418); case of small sand grains, frequently long, slender (Figs. 19.11, 19.412, 19.477); West(as far east as western Kansas)

2(1).

Abdominal tergum IX with about 40 setae (Fig. 19.415); head flat dorsally with lateral carinae prominent(Fig. 19.419); Southern

Appalachian Mountains (higher elevations) 2'.

Abdominal tergum IX with about 15 setae (Fig. 19.416); head rounded dorsally with lateral carinae not as prominent(Fig. 19.420); East, Central (as far west as Minnesota and eastern Texas, middle and lower elevations)

2

Gumaga

Fattigia pele (Ross)'

Agarodes

Thremmatidae

1.

r.

Pronotum with prominent, lateral, longitudinal ridges; anterior edge of mesonotum with only shallow and broadly triangular emargination in middle (Fig. 19.421); case smooth, strongly tapered and curved, composed of rocks without larger lateral pebbles; West Pronotum without prominent longitudinal ridges; anterior edge of mesonotum with deeper emargination in middle truncate or more sharply incised at its lateral edges(Fig. 19.422); case of coarse rocks, several larger pebbles laterally (Figs. 19.27, 19.494); widespread except not Southwest

Oligophlebodes

Neophylax

Uenoidae

1.

1'.

2(1').

T.

Mesonotal sclerites each with anterior margin straight except for lobed anteromesal excision (Fig. 19.426); pronotum with anterior margin straight and anterolateral corner angulate (Fig. 19.426); case of dark, tough silk, tapered and slightly curved, dorsal surface usually with parallel longitudinal ridges formed of silk, often with slight spiral (Figs. 10.190, 19.456); Idaho,

Montana Sericostriata surdickae Wiggins, Weaver, and Unzicker' Mesonotal sclerites each with anterior margin rounded except for common, unlobed anteromesal notch between them (Figs. 19.425, 19.427); pronotum with anterior margin and anterolateral edge curved (Figs. 19.425, 19.427) Darkened posterolateral corner of each mesonotal sclerite extending anterad on lateral margin approximately to middle of sclerite (Fig. 19.427); filaments of abdominal lateral fringe arising along half or more of most segments (Fig. 19.423); case of sand grains with thin, silken lining over interior and exterior surfaces (Figs. 19.28, 19.488); West Darkened posterolateral corner of each mesonotal sclerite extending along posterior l/3rd of lateral margin (Fig. 19.425); filaments of abdominal lateral fringe scattered and arising along less than half of each segment, but with prominent and discrete tuft of filaments on anterior edge of segment II (Fig. 19.424); case as above, but more slender (Fig. 19.487); Pacific states

'This genus is represented in North America by only one species(see also Table 19A).

2

Neothremma

Farula

—^

Figure 19.4S9a'

Figure 19.457

Figure 19.458

Figure 19.460a

Figure 19.460b

%Figure 19.459b

Figure 19.461 Figure 19.462b

Figure 19.463

Figure 19.462a

© James C.(Skip) Hodges,Jr. 2018

Figure 19.457 Molanna blenda Sibley (Molannidae) larval case, ventral.

Figure 19.458 Molannodes tinctus (Zetterstedt) (Molannidae) larval case, ventral. Figure 19.459a Glossosoma sp.(Glossosomatidae) larval case, dorsal.

Figure 19.459b

Glossosoma sp.(Glossosomatidae)

larval case, ventral.

Figure 19.460a Helicopsyche borealis (Hagen) (Hellcopsychidae) larval case, dorsal.

Figure 19.460b Helicopsyche borealis (Hagen) (Hellcopsychidae) larval case, ventral. Figure 19.461 Anisocentropus pyraloides (Walker) (Calamoceratidae) larval case, ventral. Figure 19.462a Heteroplectron amerlcanum (Walker) (Calamoceratidae) larval case, slightly left ventrolateral. Figure 19.462b Heteroplectron amerlcanum (Walker) (Calamoceratidae) larval case, ventral. Figure 19.463 Phyllolcus aeneus (Hagen) (Calamoceratidae) larval case, ventral. 661

Figure 19.464

Figure 19.471

19,405

Figure 19.472

Figure 19.466 Figure 19.467 Figure 19.468

Figure 19.469 pjgure 19.470

Figure 19.476

Figure 19.473 Figure 19.475

Figure 19.477

Figure 19.474

© James C.(Skip) Hodges,Jr. 2018

Figure 19.464 Rossiana montana Denning (Rossianidae) larval case, left lateral. Figure 19.465 Psilotreta amera Ross (Odontoceridae) Figure 19.466 Lepidostoma flinti Wallace and

Figure 19.471 Namamyia plutonis Banks (Odontoceridae) larval case, left lateral. Figure 19.472 Nerophilus californicus (Hagen) (Odontoceridae) larval case, left lateral. Figure 19.473 Marilia nobsca Milne (Odontoceridae)

Sherberger (Lepldostomatidae) larval case, sligfitly left

larval case, left lateral.

ventrolateral.

Figure 19.474 Yphria californica (Banks) (Phryganeidae) larval case, left lateral. Figure 19.475 Agarodes griseus Banks (Sericostomatidae) larval case, left lateral. Figure 19.476 Fattigia pete (Ross)(Sericostomatidae)

larval case, left lateral.

Figure 19.467 Brachycentrus americanus (Banks) (Brachycentrldae) larval case, slightly left ventrolateral. Figure 19.468 Brachycentrus chelatus Ross (Brachycentrldae) larval case, slightly left ventrolateral. Figure 19.469 Brachycentrus etowahensis Wallace (Brachycentrldae) larval case, slightly left ventrolateral. Figure 19.470 Brachycentrus numerosus (Say) (Brachycentrldae) larval case, slightly left ventrolateral. 662

larval case, left lateral.

Figure 19.477 Gumaga nigricula (McLachlan) (Sericostomatidae) larval case, left lateral.

Figure 19.482

Figure 19.481

Figure 19.478 Figure 19.479

Figure 19.483

Figure 19.480

Figure 19.484

Figure 19.485 Figure 19.488

■'9-487

.,g ^gg

© James C. (Skip) Hodges, Jr. 2018

Figure 19.478 Parthina vierra Denning (Odontoceridae) larval case, left lateral. Figure 19.479 Pseudogoera singularis Carpenter (Odontoceridae) larval case, left lateral. Figure 19.480 Cryptochia pilosa (Banks) (LImnepfillldae) larval case, slightly left ventrolateral. Figure 19.481 Pedomoecus sierra Ross (Apatanlldae) larval case, left lateral.

Figure 19.482

Goeracea genota (Ross) (Goerldae)

larval case, ventral.

Figure 19.483

Figure 19.484

Goerita betteni Ross (Goerldae), larval

case, left lateral.

Figure 19.485

Goerita fiinti Parker (Goerldae) larval

case, left lateral.

Figure 19.486

Lepania cascada Ross (Goerldae)

larval case, left lateral.

Figure 19.487

Faruia sp. (Uenoldae) larval case, left

lateral.

Figure 19.488

Neothremma sp. (Uenoldae) larval

case, left lateral.

Goera calcarata Banks (Goerldae)

larval case, ventral. 663

Figure 19.489

Figure 19.490

Figure 19.491

Figure 19.492

Figure 19.498

Figure 19.495

Figure 19.493

Figure 19.499 Figure 19.600

Figure 19.494

Figure 19.501

Figure 19.496 Figure 19.497 © James C.(Skip) Hodges,Jr. 2018

Figure 19.489 Beraea gorteba Ross (Beraeidae) larval case, left lateral.

Figure 19.490 Eobrachycentrus gelidae Wiggins (Brachycentridae) larval case, slightly left ventrolateral. Figure 19.491 Amiocentrus aspilus (Ross) (Brachycentridae) larval case, left lateral. Figure 19.492 Adicrophleps hitchcocki Flint (Brachycentridae) larval case, slightly left ventrolateral. Figure 19.493 Theliopsyche sp.(Lepidostomatidae) larval case, left lateral.

Figure 19.494 Neophylax ornatus Banks (Thremmatidae) larval case, ventral. Figure 19.495 Agrypnia improba (Hagen) (Phryganeidae) larval case, left lateral. 664

Figure 19.496 Hagenella canadensis (Banks) (Phryganeidae) larval case, left lateral. Figure 19.497 Nectopsyche tavara (Ross) (Leptoceridae) larval case, left lateral. Figure 19.498 Setodes dixiensis Holzenthal (Leptoceridae) larval case, left lateral. Figure 19.499 Ceraclea sp. (Leptoceridae) larval case, left lateral.

Figure 19.500 Leptocerus americanus (Banks) (Leptoceridae) larval case, left lateral. Figure 19.501 Oecetis cinerascens (Hagen) (Leptoceridae) larval case, slightly left ventrolateral.

Figure 19.502

^

Figure 19.504

Figure 19.505

Figure 19.507 Figure 19.506

Figure 19.503

Frgure 19.611a

Figure 19.511b

Figure 19.509 Figure 19.508

Figure 19.510

© James C.(Skip) Hodges,Jr. 2018

Figure 19.502 Apatania sp.(Apatanildae) larval case, slightly left ventrolateral. Figure 19.503 Triaenodes marginatus Sibley (Leptoceridae) larval case, left lateral. Figure 19.504 Dibusa angata Ross (Hydroptllidae)

Figure 19.508 Dicosmoecus atripes (Hagen) (Limnephilidae) larval case, left lateral. Figure 19.509 Ironoquia sp.(Llmnephilidae) larval

larval case, left lateral.

case, left lateral.

Figure 19.505 Ithytrichia clavata Morton (Hydroptllidae) larval case, left lateral. Figure 19.506 Palaeagapetus celsus (Ross) (Hydroptllidae) larval case, dorsal. Figure 19.507 Oxyethira sp.(Hydroptllidae) larval

Figure 19.511a Clostoeca disjuncta (Banks) case, ventral. (Limnephilidae) larval I Figure 19.511b Clostoeca disjuncta (Banks) case, ventral. (Limnephilidae) larval I

case, left lateral.

Figure 19.510 Ironoquia sp.(Limnephilidae) larval

case, left lateral.

665

666

Chapter 19 Trichoptera

Xiphocentronidae^^ Foretrochantin small, partly membranous, separated from pleuron by suture (Fig. 19.428); mesopleura each with lobate process extending anterodorsad (Fig. 19.428); larva as in Fig. 10.169; retreat tube of fine sand grains on rocks,

sometimes several centimeters long

Cnodocentron yavapai Moulton and Stewart'(Arizona) or Xiphocentron messapus Schmid'(Texas)

KEYS TO THE FAMILIES OF TRICHOPTERA PUPAE

Because pupae remain unknown in many genera, some specimens may not coincide with diagnostic characters employed in this key to families. Diagnostic characters to pupae of some genera were given by Ross(1944), but are not yet available for most genera. Most pupae may be identified to genus and species, however, by reference to last instar larval sclerites retained in the pupal case (except Leptoceridae, which rid their pupal cases of these sclerites) and to structures of the pharate adult within the pupal cuticle. 1. Abdomen terminated in 1 simple lobe (Fig. 19.512), which may bear pair of setal tufts (Fig. 19.515)(ignore ventral membranous lobes containing developing genitalia) 2 T. 2(1). 2'. 3(2).

3'.

Abdomen terminated in pair oflobes (Figs. 19.527, 19.550) or slender sclerotized processes (Fig. 19.53) 6 Abdominal segments III, IV,and V each with 2 pairs of hook plates (Fig. 19.512) 3 Abdominal segment III with no more than 1 pair of anterior hook plates; segments lY and V each with 1 or 2 pairs of hook plates (Figs. 19.515, 19.518) 4 Mandible with apical and subapical teeth (Fig. 19.513); body length typically greater than 5 mm; widespread RHYACOPHILIDAE

Mandible with only apical teeth or points (Fig. 19.516); body length typically less than 5 mm; widespread

4(2'). 4'.

5(4').

5'.

6(T). 6'.

7(6).

HYDROPTILIDAE

Abdominal segment IV with 1 pair of hook plates; segment V with 2 pairs of hook plates (Fig. 19.523); widespread PHILOPOTAMIDAE Abdominal segment IV with 2 pairs of hook plates and V with 1 or 2 pairs of hook plates (Figs. 19.514, 19.515, 19.518, 19.519) 5 Mandibles serrate mesally or not, each with 1 (Fig. 19.520) or 2 (Fig. 19.521) prominent mesal, subapical teeth conspicuously larger than serrations; hook plates Vp present or absent(Fig. 19.519), but if present then also with hook plates VIIIp and sometimes IXp (Fig. 19.518); widespread GLOSSOSOMATIDAE Mandibles finely serrate subapicomesally, with larger, irregular serrations sub-basomesally, without conspicuous mesal teeth (Fig. 19.522); hook plates Vp present, but hook plates VIIIp and IXp absent(Fig. 19.514); Southwest HYDROBIOSIDAE,Atopsyche Mandibles with both apical teeth and subapical teeth (Fig. 19.533) 7 Mandibles each with only single apical point or tooth (Fig. 19.524), in a few groups with serrations along mesal edge; or mandibles sometimes shorter than labrum and semi-membranous (Fig. 19.551) 8 Abdominal segments III and IV each with two pairs of hook plates; segment V with only one pair of hook plates (Fig. 19.534); widespread HYDROPSYCHIDAE

'This genus is represented in North America by only one species(see also Table 19A). The larva of Cnodocentron (Caenocentron) yavapai Moulton and Stewart(Moulton and Stewart 1997a) is indistinguishable from that of Xiphocentron messapus Schmid (Edwards 1961; Wiggins 1977, 1996).

Chapter 19 Trichoptera

667

\ ,;>IVP r,vp VI

subapical teeth Figure 19.513 Figure 19.512 Figure 19.514

IN m

Figure 19.516 iva mm

Va «

V

IVp

m

Figure 19.517

Figure 19.515

Figure 19.512 Rhyacophila sp.(Rhyacophilidae) pupal abdomen, with details of hook plates, dorsal. Figure 19.513 Rhyacophila sp.(Rhyacophilidae) pupal head, frontal. Figure 19.514 Atopsyche sp. (Hydrobiosidae) pupal abdomen, with details of hook plates, dorsal.

Figure 19.515 Glossosoma sp.(Glossosomatidae) pupal abdomen, with detaiis of hook piates, dorsal. Figure 19.516 Agrayiea sp. (Hydroptilidae) pupal head, frontal.

Figure 19.517 Agrayiea sp. (Hydroptilidae) pupal abdomen, with detaiis of hook piates, dorsal.

668

Chapter 19 Trichoptera

fVW;-T'*5-*

rVlX IVa

m? ^v„ Figure 19.520 Vlllp

Figure 19.518

-sjlla

Figure 19.521

/5^IVa .^IVp

W' /l^VIa

N Villa

TJ

Figure 19.519

Figure 19.518 Anagapetus debilis (Glossosomatidae) pupal abdomen, with details of hook plates, dorsal (after Genco and Morse, 2017). Figure 19.519 Protoptila maculata (Glossosomatidae) pupal abdomen, with details of hook plates, dorsal (after Genco and Morse, 2017).

Figure 19.522

Figure 19.520 Protoptila maculata (Glossosomatidae) pupal head, frontal (after Genco and Morse, 2017). Figure 19.521 Anagapetus debilis (Glossosomatidae), pupal head, frontal (after Genco and Morse, 2017). Figure 19.522 Atopsyche sp.(Hydrobiosidae), pupal mandibles, dorsal (after Rueda Martin, 2006).

Chapter 19 Trichoptera

669

Figure 19.524

Figure 19.523 Figure 19.525

Figure 19.526

Figure 19.527

Figure 19.523 Dolophilodes sp.(Philopotamidae) pupal abdomen, with details of hook plates, dorsal. Figure 19.524 Polycentropus sp.(Polycentropodldae) pupal head, frontal. Figure 19.525 Polycentropus sp.(Polycentropodldae) pupal abdomen, with details of hook plates and anal processes, dorsal.

Figure 19.528

Figure 19.526 Psychomyia sp.(Psychomyiidae) pupal head, frontal.

Figure 19.527 Psychomyia sp.(Psychomyiidae) pupal abdomen, with details of hook plates and anal processes, dorsal.

Figure 19.528 Xiphocentron sp.(Xiphocentronidae) pupal abdomen, with details of hook plates, dorsal.

e

Q Figure 19.531a

Figure 19.531b

Figure 19.529

Figure 19.530

Figure 19.533

III

IV

i.

__Vp

IVp

v,f(^ VII

Figure 19.534 Figure 19.535

Figure 19.536

Figure 19.529 Phylocentropus sp.(Dipseudopsldae) pupal abdomen, with details of hook plates, dorsal. Figure 19.530 Austrotinodes sp.(Ecnomidae) pupal abdominal hook plates, dorsal (after O.S. Flint, 1973). Figure 19.531 a, 19.531 b Manophylax butleri (Apataniidae), male and female pupal anal processes,

Figure 19.534 Hydropsyche sp.(Hydropsychidae) pupal abdomen, with details of hook plates and anal

respectively, dorsal (after Schuster 1997). Figure 19.532 Austrotinodes sp.(Ecnomidae) pupal right mandible, dorsal (after O.S. Flint 1973).

Figure 19.536 Helicopsyche sp.(Helicopsychidae) pupal abdomen, with details of hook plates and anal processes, dorsal.

Figure 19.533 Hydropsyche sp.(Hydropsychidae) pupal head, frontal. 670

processes, dorsal.

Figure 19.535 Fattigia sp.(Sericostomatidae) pupal abdomen, with details of hook plates and anal processes, dorsal.

Chapter 19 Trichoptera

671

IV

Va

^4 CrIts'VI .CSe^-

'fiSPvii

Figure 19.539

Figure 19.538

Figure 19.537

frontal

projection

m

Figure 19.540

VII

I)#

Figure 19.543

Figure 19.541

Figure 19.537 Lepidostoma sp. (Lepidostomatidae) pupal abdomen, with details of hook plates and anal processes, dorsal. Figure 19.538 Goereilla sp.(Rosslanldae) pupal abdomen, with details of hook plates, dorsal. Figure 19.539 Goereilla sp.(Rosslanldae) pupal anal processes, dorsal and right lateral. Figure 19.540 Goereilla sp.(Rosslanldae) pupal head, right lateral.

Figure 19.542

Figure 19.541 Heteroplectron sp.(Calamoceratidae) pupal abdomen, with details of hook plates and anal processes, dorsal.

Figure 19.542 Neothremma sp.(Uenoidae) pupal abdomen, with details of hook plates and anal processes, dorsal.

Figure 19.543 Neothremma sp.(Uenoidae) pupal abdominal segment VIII, showing lateral fringe, ventral.

672

7'. 8(6',?').

Chapter 19 Trichoptera

Abdominal segments III and IV each with one pair of hook plates; segment V with two pairs of hook plates (Fig. 19.550) Abdomen terminated by one or more pairs of apically rounded semi-membranous lobes (Figs. 19.525, 19.529), or by short, broad, truncate and more sclerotized lobes that are typically concave dorsally or caudally (Fig. 19.550)

8'.

Abdomen terminated by one pair of sclerotized processes, typically slender and elongate (Fig. 19.53), but sometimes triangular (Fig. 19.537) or conical

9(8).

Abdominal segment I with median dorsal lobe or ridge (Fig. 19.550); anal lobe short, broad, more or less truncate and typically concave dorsally (Fig. 19.550) or caudally; widespread

(Fig. 19.556)

9'. 10(9').

8

9

14

PHRYGANEIDAE

Abdominal segment I without median dorsal ridge, anal lobe simple and rounded, bearing tufts of setae (Fig. 19.525)

10

Abdominal segment II without hook plates (Fig. 19.529)

11

10'. 11(10).

Abdominal segment II with hook plates (Fig. 19.527) 12 Terminal abdominal segment with short lobe on each side of longer anal lobe (Fig. 19.529); East DIPSEUDOPSIDAE,Phylocentropus 11'. Terminal abdominal segment with only single pair of apical lobes (Fig. 19.525); widespread POLYCENTROPODIDAE 12(10'). Mandibles each with apex attenuated as sclerotized filament(Fig. 19.526) 13 12'. Mandibles not attenuated apically (Fig. 19.532); Texas ECNOMIDAE,Austrotinodes texensis Bowles' 13(12). Abdominal segment VIII with hook plates; segment V with posterior hook plates(Vp)much wider than long (Fig. 19.527); widespread PSYCHOMYIIDAE 13'. Abdominal segment VIII without hook plates; segment V with posterior hook plates(Vp)longer than wide (Fig. 19.528); Arizona, Texas

14(8').

14'.

XIPHOCENTRONIDAE

Antennae markedly longer than body and coiled around end of abdomen, held in place by pair of setose dorsal lobes(Fig. 19.544); widespread LEPTOCERIDAE Antennae little, if any, longer than body and not coiled around end of abdomen 15

15(14'). Labrum with major setae hooked apically (Fig. 19.51)

Labrum with setae typically straight apically and not hooked (Fig. 19.545), although slightly inflected in some groups(Fig. 19.555) 16(15'). Abdomen with dense lateral fringe of hair-like filaments on several segments

27

15'.

(e.g.. Figs. 19.52, 19.53)

16'. 17(16').

17'.

18(17').

Abdomen without lateral fringe (Fig. 19.549) or with only few filaments near male inferior appendage sheaths Abdomen with anterior hook plates in the form of a single hook; apices of anal processes tapered and curved dorsad, bifid apically (Fig. 19.549); East(highly localized)

16 19 17

BERAEIDAE,Beraea

Abdomen with anterior hook plates each bearing 2 or more hooks; anal processes not tapered and straight, blunt apically (Fig. 19.536) or hooked laterad and acute apically (Figs. 19.531a, 19.531b); widespread 18 Anal processes short and straight, each with several mesal setae and 2 long apical setae (Fig. 19.536); widespread HELICOPSYCHIDAE,Helicopsyche

'This genus is represented in North America by only one species(see also Table 19A).

Chapter 19 Trichoptera

673

antenna

Figure 19.545

Figure 19.544

apical Figure 19.546

bristles

VaK VP apex attenuated

Figure 19.547

Figure 19.1

Figure 19.549

Figure 19.544 Oecetis sp.(Leptoceridae) pupal abdominal segments VIII, IX, X, and anal processes, showing coiled antennae, left lateral. Figure 19.545 Molanna sp.(Molannidae) pupal head,

Figure 19.547 Psiiotreta sp. (Odontoceridae) pupal head, with detail of attenuate apex of right mandible, frontal.

frontal.

Figure 19.548 Psiiotreta sp.(Odontoceridae) pupal abdomen, with details of hook plates and anal

Figure 19.546 Molanna sp.(Molannidae) pupal abdomen, with details of hook plates and anal processes, dorsal.

Figure 19.549 Beraea sp.(Beraeidae) pupal abdomen, with details of hook plates and anal processes, dorsal.

processes, dorsal.

674

Chapter 19 Trichoptera

Figure 19.551

Figure 19.553 Figure 19.550 Figure 19.552

IV

Va ttil

Figure 19.555

Figure 19.554 Figure 19.556

Figure 19.550 Banksiola sp.(Phryganeidae) pupal abdomen, with details of hook plates and anal processes, dorsal. Figure 19.551 Ptilostomis sp.(Phryganeidae) pupal head, frontal.

Figure 19.552 Apatania sp. (Apataniidae) pupal head, frontal.

Figure 19.553 Apatania sp.(Apataniidae) pupal abdomen, with details of hook plates and anal processes, dorsal.

Figure 19.554 Brachycentrus sp.(Brachycentridae) pupal abdomen, with details of hook plates and anal processes, dorsal. Figure 19.555 Brachycentrus sp.(Brachycentridae) pupal head, frontal. Figure 19.556 Goeracea sp.(Goerldae) pupal abdomen, with details of hook plates and anal processes, dorsal.

Chapter 19 Trichoptera

18'.

Anal processes long and hooked laterad apically, each with 3 long subapical setae (Figs. 19.531a, 19.531b); Kentucky, Tennessee, West Virginia

19(16).

Anal processes in dorsal aspect broad basally and angulate or pointed apically (Figs. 19.537, 19.539)

20

19'.

Anal processes more slender throughout, and elongate (Fig. 19.53)

21

20(19).

Anal processes pointed apically (Fig. 19.539); head with prominent frontal projection (Fig. 19.540); Northwest

ROSSIANIDAE

20'.

Anal processes angulate apically (Fig. 19.537); head of pupa without prominent frontal projection; widespread

21(19').

Most abdominal segments with dorsal clusters of setae (Fig. 19.541); East, Southeast, Southwest, West Coast Abdominal segments with dorsal setae mostly single and not in clusters(Fig 19.554)

21'. 22(21'). 22'.

^

APATANIIDAE (in part)

LEPIDOSTOMATIDAE (in part) CALAMOCERATIDAE

22

Anal processes closely approximate basally and widely divergent apically (Fig. 19.548)

23

Anal processes not widely divergent but subparallel over entire length (Fig. 19.53), or sometimes crossed over one another (Fig. 19.553)

24

23(22). Mandibles with apices attenuated (Fig. 19.547); widespread 23'.

■

ODONTOCERIDAE (in part)

Mandibles with apices broadly pointed but not attenuated

(Fig. 19.555); widespread BRACHYCENTRIDAE 24(22'). Anal processes closely approximate (Fig. 19.535) 25 24'. Anal processes not as closely approximate (Fig. 19.546) 26 25(24). Anal processes bearing many setae before apex (Fig. 19.535); hook plates Illa-VIIa and Vp present(Fig. 19.535); widespread SERICOSTOMATIDAE 25'. Anal processes with few setae; hook plates Illa-VIIIa and Vp present; Southern Appalachian Mountains ODONTOCERIDAE (in part) 26(24'). Mandibles much longer than labrum and slender, with lateral margins of blades straight and mesal margins bearing minute serrations; labrum with major setae straight apically (Fig.19.545); widespread

^

26'.

MOLANNIDAE

Mandibles short, slightly longer than labrum and more nearly equilateral triangles, with lateral margins indented and mesal margins without minute serrations; labrum with major setae hooked apically (Fig. 19.51)

27(15,26'). Anal processes broad basally, flattened, and angulate apically in dorsal view (Fig. 19.537); widespread

27'.

Anal processes usually slender and elongate (Fig. 19.52), but sometimes

^

28(27').

shorter and/or conical(Fig. 19.556), or with hooked apical spur Abdomen with lateral fringes of slender filaments absent, although isolated patches may occur ventrally on one segment(Figs 19.542, 19.543); West

28'.

N

N

N

27

LEPIDOSTOMATIDAE(in part)

^

^

675

28

UENOIDAE

Abdomen with lateral fringes of filaments continuous over several

segments (Fig. 19.52)

29

preapical spur mesal

Figure 19.557

Figure 19.560

Figure 19.558

Figure 19.561

Figure 19.559

mesal

Figure I9.d63

Figure 19.564 Figure 19.562

preapical spur.

Figure 19.568 mesoscutal setose wart

m

Figure 19.565

Figure 19.557 Rhyacophila sp.(Rhyacophilldae) adult left foretibia and tarsus, anterior.

Figure 19.558 Atopsyche sp.(Hydrobiosidae) adult right maxillary palp, anterior. Figure 19.559 Glossosoma sp.(Glossosomatidae) adult head and pro- and mesonota, dorsal. Figure 19.560 Glossosoma sp.(Glossosomatidae) adult left foretibia and tarsus, anterior. Figure 19.561 Glossosoma sp.(Glossosomatidae)

adult right maxillary palp, anterior. Figure 19.562 Agraylea sp.(Hydroptilidae) adult head and pro- and mesonota, dorsal. Figure 19.563 Agraylea sp.(Hydroptilidae) right wings, dorsal. 676

Figure 19.566

Figure 19.567

Figure 19.569

Figure 19.564 Dolophilodes sp.(Philopotamidae) adult left maxillary palp, anterior. Figure 19.565 Polycentropus sp.(Polycentropodidae) adult head and pro- and mesonota, dorsal. Figure 19.566 Polycentropus sp.(Polycentropodidae) right forewing, dorsal (from H.H. Ross, 1944). Figure 19.567 Polycentropus sp.(Polycentropodidae) adult left foretibia and tarsus, anterior.

Figure 19.568 Nyctiophylax sp.(Polycentropodidae) right wings, dorsal (from H.H. Ross, 1944). Figure 19.569 Psychomyia sp.(Psychomyiidae) adult left foretibia and tarsus, anterior.

Chapter 19 Trichoptera

29(28').

677

Morphological characters diagnostic for pupae of the remaining families have not been resolved, but the cases in which larvae pupate are similar to larval cases and provide some distinguishing features (eases for larvae of North American genera were illustrated by Wiggins, 1996): APATANIIDAE (in part); Appalachian Mtns., North, West Pupal cases entirely of rock fragments, with plant materials sometimes added to basic case in one uncommon western subgenus(Manophylax (M.), mostly in hygropetric habitats); pupal case of common eastern genus {Apatania)strongly tapered and curved with anterior opening nearly on same plane as venter of case. GOERIDAE; East, Midwest, Northwest

Anal processes either more slender and longer than in Fig. 19.53 or shorter and conical (Fig. 19.556). Pupal cases of rock fragments only; cases of common genera with ballast stones fixed symmetrically along sides (similar to larval cases in Figs. 10.179, 19.18, 19.19, 19.173, 19.175, 19.482, 19.483), others lack the ballast stones (similar to larval cases in Figs. 19.174, 19.176, 19.484-19.486). LIMNEPHILIDAE; widespread

Pupal cases typically of plant materials (Figs. 19.22, 19.25, 19.337-19.339, 19.341, 19.342, 19.432-19.434, 19.440, 19.441b, 19.445-19.447, 19.449, 19.450,

19.480, 19.509-19.511) or mineral materials (Figs. 19.23, 19.335, 19.336, 19.340, 19.343, 19.438, 19.441a, 19.442a, 19.448, 19.451, 19.452a, 19.508), or a combination of the two (Figs. 19.24, 19.431, 19.435-19.437, 19.439, 19.442c-19.444, 19.452b), occasionally with mollusk shells (Fig. 19.442b). Anal processes typically long (Fig. 19.53), but shorter in few genera, sometimes conical and pointed or hooked apically (e.g., Ecclisomyia of western North America). THREMMATIDAE; widespread

Pupal cases of rock fragments alone; in cases of most common North American genus {Neophylax), ballast stones fixed asymmetrically along sides (similar to larval cases in Figs. 19.27, 19.494).

^

KEY TO THE FAMILIES OF TRICHOPTERA ADULTS 1.

Body length typically less than 5 mm; mesoscutum without setose warts, mesoscutellar warts transverse and meeting mesally to form angulate ridge (Fig. 19.562); hind wings narrow, apically acute (Fig. 19.563), and often with posterior fringe of long setae; adults as in Fig. 10.194;

1'.

Body length variable, often more than 5 mm; mesoscutum often

2(1').

with setal warts, mesoscutellar warts typically rounded or elongate (Fig. 19.55); hind wings typically broader than above and apically rounded (Fig. 19.589), if posterior fringe present, then with shorter setae Head with 3 ocelli dorsally (Fig.19.55)

2'.

Head without ocelli(Fig. 19.572)

3(2).

Maxillary palps each with 5 segments, terminal segment flexible, curved, of different structure than preceding segments, and typically at least twice as long as segment 4(Figs. 19.564, 19.570); widespread PHILOPOTAMIDAE(p. 712) Maxillary palps each of 2, 3, 4, 5, or 6 segments, segment 5 or 6, if present, similar in structure to preceding segments and about same length as segment 4(Figs. 19.602, 19.603) 4

^

^1^

widespread

' 3'.

HYDROPTILIDAE (in part)(p. 691)

2

3

14

678

Chapter 19 Trichoptera

labial

palp Figure 19.571

maxillary

palp ^ Figure 19.570 Figure 19.572

Figure 19.573

2 preapical spurs

mesoscutal setose wart

Figure 19.574

Figure 19.576

pronotal posterior

setose wart

Figure 19.575

setose wart

median

fissure

mesoscutal setose wart

Figure 19.577

Figure 19.570 Austrotinodes sp.(Ecnomldae) adult head, right lateral (after O.S. Flint, 1973). Figure 19.571 Austrotinodes sp.(Ecnomidae) right wings, dorsal (after O.S. Flint, 1973). Figure 19.572 Hydropsyche sp.(Hydropsychidae) adult head and pro- and mesonota, dorsal. Figure 19.573 Ptiylocentropus sp.(Dipseudopsidae) right forewing, dorsal (from H.H. Ross, 1944). Figure 19.574 Xiphocentron sp.(Xiphocentronidae) adult head and pro- and mesonota, dorsal.

Figure 19.578

Figure 19.575 Banksiola sp.(Phryganeidae) adult head and pro- and mesonota, dorsal. Figure 19.576 Banksiola sp.(Phryganeidae) adult left middle tibia and tarsus, anterior.

Figure 19.577 Helicopsyche sp.(Helicopsychidae) adult head and pro- and mesonota, dorsal. Figure 19.578 Agarodes sp.(Sericostomatidae) adult head and pro- and mesonota, dorsal.

median

vulval lobe A +2+3

Figure 19.581

Figure 19.579

( 8; Figure 19.580 sclerotized lobe

pronotal setose

preapical

warts

Figure 19.584

Figure 19.582

Figure 19.583

Figure 19.585

Figure 19.586

Figure 19.579 Apatania sp.(Apataniidae) right wings,

Figure 19.583 Brachycentrus sp.(Braohycentridae)

dorsai.

adult left middle tibia and tarsus, anterior.

Figure 19.580 Apatania sp.(Apataniidae) female genitaiia, showing semi-membranous median vuivai

Figure 19.584 Brachycentrus sp.(Braohycentridae) adult abdominal sternum V, showing sclerotized lobes associated with internal glands, ventral. Figure 19.585 Lepania sp. (Goeridae) adult head and

iobe, ventral.

Figure 19.581 Moselyana sp.(Apataniidae) female genitaiia, showing semimembranous median vuivai iobe, ventral.

Figure 19.582 Brachycentrus sp.(Braohycentridae) adult head and pro- and mesonota, corsai.

pro- and mesonota, dorsai.

Figure 19.586 Lepania sp.(Goeridae) right wings, with details of posterior margin of forewing and anterior margin of hind wing, dorsai.

679

680

Chapter 19 Trichoptera

aedeagus

median vuival lobe

^phallus Figure 19.587

Figure 19.588

Figure 19.589 Ql^ q,^ scape

pedicel

scape

pedicel

2 preapical*

spurs ^

mesoscutal seta I area

Figure 19.592 Figure 19.593

Figure 19.591

Figure 19.590

3ZI seta I brush

spines

single mesoscutellar setose wart

Figure 19.595 Figure 19.594

Figure 19.587 Goereilla sp.(Rossianidae) male genitalia, ventral, and phallus right lateral, showing spines on aedeagus and absence of parameres. Figure 19.588 Goereilla sp.(Rossianidae)female genitalia, showing rounded apical margin of median

Figure 19.597 Figure 19.596

Figure 19.592 Oecetis sp.(Leptoceridae) adult left middle tibia, anterior.

Figure 19.593 Molanna sp.(Molannidae) adult left middle tibia and tarsus, anterior.

Figure 19.589 Farula sp.(Uenoldae) right wings,

Figure 19.594 Beraea sp.(Beraeidae) adult head and pro- nd mesonota, dorsal. Figure 19.595 Beraea sp.(Beraeidae) adult left

dorsal.

middle tarsus, anterior.

Figure 19.590 Heteroplectron sp.(Calamoceratidae) adult head and pro- and mesonota, dorsal. Figure 19.591 Oecetis sp.(Leptoceridae) adult head and pro- and mesonota, dorsal.

Figure 19.596 Psilotreta sp.(Odontoceridae) adult head and pro- and mesonota, dorsal. Figure 19.597 Psilotreta sp.(Odontoceridae) adult right maxillary palp, anterior.

vulval lobe, ventral.

Chapter 19 Trichoptera

4(3').

681

5

4'.

Maxillary palps each of 5 segments, segment 2 about as long as segment 1 and often rounded (Fig. 19.561) Maxillary palps each of 2, 3, 4, 5 or 6 segments, segment 2 longer than

5(4).

segment 1 and slender (Figs. 19.602, 19.603) Maxillary palps each with segment 2 rounded and globose (Fig. 19.561)

8 6

5'.

Maxillary palps each with segment 2 not globose, but of same general

6(5). 6'. 7(6').

cylindrical shape as segment 1 (Fig. 19.558); Southwest HYDROBIOSIDAE(p. 690) Foretibiae each with preapical spur (Fig. 19.557); adults as in Fig. 10.191; widespread RHYACOPHILIDAE(p. 718) Foretibiae without preapical spurs(Fig. 19.560) 7 Pronotum with mesal setal warts widely spaced (Fig. 19.559); adults as in Fig. 10.196; widespread GLOSSOSOMATIDAE(p. 687)

7'.

Pronotum with mesal setal warts closely approximated (as in Fig. 19.562); Northeast, Northwest HYDROPTILIDAE (in part)(p. 691)

8(4').

11'.

Middle tibiae each with 2 preapical spurs(Fig. 19.576), i.e., spurs typically 2, 4,4 on each pro-, mid-, and hind tibia, respectively; widespread PHRYGANEIDAE(p. 715) Middle tibiae each with 1 (Fig. 19.600) or no preapical spurs, i.e., spurs typically 1, 2-3,4 on each pro-, mid-, and hind tibia, respectively 9 Hind wings each with anterior margin bearing row of stout, apically hooked, setae (Fig. 19.586) 10 Hind wings with anterior margins bearing only straight or slightly curved, normal setae, if any (Fig. 19.579) 12 Hind wings each with vein R1 complete, extending to apical margin of wing (Fig. 19.589); widespread 11 Hind wings each with vein R1 incomplete, not extending to apical margin of wing (Fig. 19.586); Northwest GOERIDAE (in part)(p. 690) Forewings about 2.5 times as long as broad (Figs. 19.758, 19.759); widespread THREMMATIDAE(p.723) Forewings more than 3 times as long as broad (Figs. 19.760, 19.761);

12(9').

West Small black or dark gray insects; length of each forewing 3-8 mm

8'. 9(8'). 9'.

10(9). 10'. 11(10).

12'. 13(12).

13'.

without color pattern 13 Medium or large insects; length of each forewing 12-35 mm with various colors and patterns; adults as in Fig. 10.199; widespread LIMNEPHILIDAE(p. 698) Hind wings each with fork of R2 and R3(Fork I) sessile or rooted, arising at or basal of one or more crossveins; fork of R4 and R5 (Fork 11) distal, arising well beyond any other forks or crossveins (Fig. 19.756); Northwest ROSSIANIDAE(p. 720) Hind wings each with fork of R2 and R3 absent or petiolate and arising beyond all crossveins; fork of R4 and R5 more nearly basal, arising at level of other wing forks and crossveins (Figs. 19.604-19.608); adults as in

Fig. 10.193; Appalachian Mtns., North, West 14(2'). 14'. 15(14). 15'. 16(15).

UENOIDAE (p. 723)

Maxillary palps each with 5 or 6 segments(Fig. 19.54) Maxillary palps each with fewer than 5 segments Maxillary palps each with apparently 6 segments Maxillary palps each with 5 segments(19.54) Forewings each 4.5 mm long; Arkansas, Missouri

(Ozark Mtns.)

APATANIIDAE(p. 686) 15 24 16 17

PSYCHOMYIIDAE(in part) .... Paduniella nearctica Flint^

'This genus is represented in North America by only one species (see also Table 19A).

682

Chapter 19 Trichoptera

posterior

1 preapical ■

setose wart

spur

2 preapical spurs

Figure 19.603

Figure 19.600

Figure 19.598

Figure 19.601

Figure 19.602

Figure 19.599

Figure 19.598 Lepidostoma sp.(Lepidostomatidae) adult head and pro- and mesonota, dorsal. Figure 19.599 Lepidostoma sp.(Lepidostomatidae) adult left middle tibia and tarsus, anterior.

Figure 19.600 Limnephilus sp.(Limnephilidae) adult left middle tibia and tarsus, anterior.

Figure 19.601 Limnephilus sp.(Limnephilidae) adult head and pro- and mesonota, dorsal. Figure 19.602 Limnephilus sp.(Limnephilidae) adult female left maxillary palp, anterior. Figure 19.603 Limnephilus sp.(Limnephilidae) adult male left maxillary palp, anterior.

16'.

Forewings each 11-13 mm long;

17(15').

Maxillary palps each with terminal segment 5 flexible, different in structure from preceding segments (with numerous transverse striae or annulations) and typically at least twice as long as penultimate segment(Fig. 19.570) 18 Maxillary palps each with terminal segment 5 similar in structure to preceding segments and typically about same length as penultimate segment(Fig. 19.54), or palp with some segments bearing long setal brush (Fig. 19.597) 24 Antennae much longer than body; middle tibiae without preapical spurs (Fig. 19.592); adults as in Fig. 10.197; widespread LEPTOCERIDAE (in part)(p. 697) Antennae typically little, if any, longer than body; if longer, then middle tibiae with preapical spurs(Fig. 19.593) 19 Mesoscutum with setose warts(Fig. 19.565) 20 Mesoscutum without setose warts or setae (Fig. 19.572); adults as in Fig. 10.192; widespread HYDROPSYCHIDAE(p. 690)

Southeast

17'.

18(17). 18'. 19(18'). 19'. 20(19).

20'.

21(20).

21'.

CALAMOCERATIDAE (in part) .... Anisocentropuspyraloides)(Walker)'

Mesoscutal setose warts ovoid, their combined area smaller than that

of mesoscutellum (Fig. 19.565) 21 Mesoscutal setose warts quadrate and appressed along median line, their combined area exceeding that of mesoscutellum (Fig. 19.574); Arizona, Texas XIPHOCENTRONIDAE(p. 723) Foretibiae each typically with preapical spur (Fig. 19.567) or, if preapical spur absent, then basal segment of tarsus shorter than twice the length of the longer apical spur (as in Fig. 19.567) 22 Foretibiae without preapical spurs; basal segment of each tarsus at least twice as long as longer apical spur (Fig. 19.569); widespread PSYCHOMYIIDAE(p. 718)

'This genus is represented in North America by only one species(see also Table 19A).

Chapter 19 Trichoptera

683

Figure 19.604

petiole

Figure 19.605

M3+4 Cula+b

Figure 19.606

Figure 19.607

Figure 19.608

© R.W. Holzenthal 2018

Figure 19.604 Apatania zonella (Apataniidae) right forewing, dorsal. Figure 19.605 Pedomoecus sierra (Apataniidae) right wings, dorsal. Figure 19.606 Allomyia cascadis (Apataniidae) right hind wings, dorsal.

Figure 19.607 Manophylax annulatus (Apataniidae) right wings, dorsal. Figure 19.608 Moselyana comosa (Apataniidae) right wings, dorsal.

684

22(21).

Chapter 19 Trichoptera

Forewings each with vein R1 branched near apex (Fig. 19.571); Texas ECNOMIDAE(p. 687) 22'. Forewings each with vein R1 unbranched (Fig. 19.573) 23 23(22'). Forewings each with vein R2 branched from R3 at radial crossvein r (Fig. 19.573); East DIPSEUDOPSIDAE(p. 687) 23'. Forewings each with veins R2 and R3 either fused throughout as vein R2+3 (Fig. 19.568), or with the two veins separated near apex of wing (Fig. 19.566); widespread POLYCENTROPODIDAE(p. 718) 24(14',17'). Mesoscutum without setose warts and setae (Fig. 19.594); tarsal segments 2-5 with spines only near their apices (Fig. 19.595); East(highly localized) BERAEIDAE(p. 686) 24'. Mesoscutum with setose warts (Figs. 19.577, 19.578, 19.582, 19.596, 19.598) or setal areas (Figs. 19.590, 19.591); tarsal segments 2-5 with spines typically arranged irregularly (Fig. 19.583) 25 25(24'). Mesoscutal setae arising in diffuse area over nearly entire length of mesoscutum (Figs. 19.590, 19.591) 26 25'. Mesoscutal setae largely confined to pair of small, discrete, warts (Figs. 19.577, 19.578, 19.582, 19.596, 19.598) 28 26(25). Antennae with scapes about twice as long as pedicels, head typically with posteromesal ridge dorsally (Fig. 19.590); East, Southeast, Southwest, West Coast CALAMOCERATIDAE(p. 687) 26'. Antennae with scapes at least 3 times longer than pedicels, head without posteromesal ridge dorsally (Fig. 19.591) 27 27(26'). Antennae markedly longer than body; pronotum consisting of pair of lateral erect, platelike warts separated by wide, mesal excavated collar usually hidden by produced, angulate anterior margin of mesonotum (Fig. 19.591); middle tibiae without preapical spurs (Fig. 19.592); widespread LEPTOCERIDAE(in part)(p. 697) 27'. Antenna little, if any, longer than body; pronotum with warts much closer together, not platelike; middle tibiae each with 2 preapical spurs (Fig. 19.593); adults as in Fig. 10.198; widespread MOLANNIDAE(p. 711) 28(25'). Head with posterior setose warts relatively large, extending from mesal margins of eyes to mid-dorsal line and anteriorly to middle of head (Fig. 19.577); antennae never longer than forewings; widespread HELICOPSYCHIDAE (p. 690) 28'. Head with posterior setose warts smaller than above (e.g., Fig. 19.596); or antennae about 1.5 times as long as forewings 29 29(28'). Mesoscutellum with single median setose wart extending over most or all of its entire length (Fig. 19.596) 30 29'. Mesoscutellum with pair of setose warts occupying about half its length (Fig. 19.582) and sometimes touching along mid-dorsal line (Fig. 19.598) 31 30(29). Mesoscutellum almost entirely covered by single setose wart, setae arising over most of wart(Fig. 19.596; bare mesally in Pseudogoera singularis); maxillary palps each 5-segmented (4-segmented in male Pseudogoera singularis)-, widespread ODONTOCERIDAE(p. 711) 30'. Mesoscutellum with setose wart narrower, setae mainly confined to periphery (Fig. 19.585); maxillary palps each 5-segmented in female and 3-segmented in male; East, Midwest, Northwest GOERIDAE(in part)(p. 690) 31(29'). Pronotal setose warts fused into single transverse wart on each side of mid-line, median fissure of mesoscutum deep (Fig. 19.578); widespread SERICOSTOMATIDAE(p. 720)

Figure 19.609

Figure 19.610

Figure 19.611

M1+2 M1+2

M3+4

M3+4

Figure 19.612

maxillary palp maxillary palp

labial palp

labial palp

Figure 19.613

Figure 19.614

R.W. Holzenthal 2018

Figure 19.609 Brachycentrus americanus (Brachycentridae) right forewing, dorsal. Figure 19.610 Adicrophleps hitchcocki (Brachycentridae) right forewing, dorsal. Figure 19.611 Micrasema wataga (Brachycentridae) right wings, dorsal.

Figure 19.612 Amiocentrus aspilus (Brachycentridae) right hind wing, dorsal. Figure 19.613 Micrasema wataga (Brachycentridae) maxillary and labial palps. Figure 19.614 Amiocentrus aspilus (Brachycentridae) maxillary and labial palps. 685

686

31'. 32(31').

32'.

Chapter 19 Trichoptera

Pronotum with 2 discrete setose warts on each side of mid-line, median fissure of mesoscutum not as deep as above (Fig. 19.582)

32

Middle tibiae each with 1 or 2 preapical spurs arising at point about one-third distance from apex of tibia (Fig. 19.583), or middle tibiae without preapical spurs; abdominal segment V with internal glands opening in pair of ventral sclerotized lobes(Fig. 19.584); widespread BRACHYCENTRIDAE(p. 686) Middle tibiae each with 2 preapical spurs arising from about mid-point of tibia (Fig. 19.599); ventral abdominal glands not apparent; adults as in Fig. 10.195; widespread LEPIDOSTOMATIDAE(p. 697)

KEYS TO THE GENERA OF TRICHOPTERA ADULTS

Adults of North American genera may be identified also through the use of the keys and references provided by Betten (1934), Ross(1944), Armitage and Hamilton (1990), Armitage (1991), Schmid (1998), and Ruiter (2000).

Apataniidae 1. Forewing Sc ending in transverse crossvein (Fig. 19.604); Appalachian Mtns., North, Southwest 1'. Forewing Sc ending in wing margin (Fig. 19.605) 2(1'). Tibial spurs 1, 3, 4 on fore-, mid-, and hind tibiae, respectively; hind wing Fork Y absent(Fig. 19.606); West 2'. Tibial spurs 1, 2, 2 or 1, 2, 4; hind wing Fork V present (Figs. 19.605, 19.607, 19.608) 3(2'). Tibial spurs 1, 2, 2; hind wing Fork I present (Figs. 19.605, 19.608) 3'. Tibial spurs 1, 2, 4; hind wing Fork I absent(Fig. 19.607); Alaska, Appalachian Mtns., Idaho 4(3). Forewing R2 vein straight (Fig. 19.605); male phallicata and parameres

3-4 times as long as thick; Northwest 4'.

Apatania 2

Allomyia 3 4

Manophylax

Pedomoecus sierra Ross'

Forewing R2 vein sinuous(Fig. 19.608); male phallicata and parameres much longer and more slender; adults as in Fig. 10.193; Oregon,

Washington

Moselyana comosa Denning'

Beraeidae

(Beraea—East, highly localized—is the only genus in North America). Brachycentridae (modified from the work by Schmid, 1998) 1.

Tibial spurs 2, 4, 4 on fore-, mid-, and hind tibiae, respectively;

Northwest r. 2(1').

2'. 3(2').

Tibia! spurs 2, 2, 2 or 2, 2, 3 or 2, 3, 3 Forewings with R1 sinuate subapically, beside pterostigma (Fig. 19.609); tibial spurs 2, 2, 3 or 2, 3, 3; widespread Forewings with R1 only slightly and gradually curved anterad subapically (Fig. 19.610); tibial spurs 2, 2, 2 Forewing with posterior anal vein angled sharply on posterior wing margin before terminating on vein lA (Fig. 19.610);

Northeast 3'.

Eobrachycentrus gelidaeV^iggins^ 2

Brachycentrus 3

Adicrophleps hitchcocki Flint'

Forewing with posterior anal vein gradually curved, not touching posterior wing margin before terminating on vein 1A (Figs. 19.611, 19.612)

'This genus is represented in North America by only one species (see also Table 19A).

4

Chapter 19 Trichoptera

687

R2+3

Figure 19.615

Figure 19.616

Figure 19.615 Heteroplectron americanum (Calamoceratidae) right hind wing, dorsal.

4(3').

4'.

R.W. Holzenthal 2018

Figure 19.616 Phylloicus aeneus (Calamoceratidae) right hind wing, dorsal.

Male labial palps as long as 3-segmented maxillary palps(Fig. 19.613); hind wings with R4+5 apparently unbranched (Fig. 19.611); widespread Male labial palps twice as long as 2-segmented maxillary palps (Fig. 19.614); female hind wings with R4 and R5 distinctly branched

(Fig. 19.612); Northwest

Micrasema

Amiocentrus aspilus(Ross)'

Calamoceratidae

1.

Maxillary palps each 5-segmented

1'.

Maxillary palps each 6-segmented; Southeast

2(1).

Hind wing Fork I present, first fork of Rs originating at r-m crossvein (Fig. 19.615); East, West Coast Hind wing Fork I absent and first fork of Rs originating distal of r-m erossvein (Fig. 19.616); Southwest

2'.

2

Anisocentropuspyraloides(Walker)' Heteroplectron

Phylloicus

Dipseudopsidae (Phylocentropus—East—is the only genus in North America) Ecnomidae

{Austrotinodes texensis Bowles'—Texas—is the only species in North America) Glossosomatidae

(adapted from the works by Mosely 1954; Ross 1956; Schmid 1998; Robertson and Holzenthal 2013) 1. Tibial spurs 2, 4,4 on fore-, mid-, and hind tibiae, respectively 2 1'. Tibial spurs 0, 3, 3, or 0, 4, 4, or 1, 4, 4, with foretibial spur hair-like or absent subfamily PROTOPTILINAE,4 2(1). Hind wing discoidal cell closed and with R1 long, ending on wing margin beyond origin of Fork 1 (Fig. 19.617) subfamily GLOSSOSOMATINAE,3 2'. Hind wing discoidal cell open and with R1 short, simple, often weak, apparently fused with base of R2+3 or apex of Sc well before origin of Fork I(Fig. 19.618); widespread subfamily AGAPETINAE,Agapetus 3(2). Upper and lower parts of mesepisternum on each side separated by constriction (Fig. 19.619); adults as in Fig. 10.196; widespread Glossosoma 3'. Upper and lower parts of mesepisternum on each side separated by transverse suture (Fig. 19.620); West Anagapetus 4(T). Forewing media vein M with only 2 branches. Forks III and IV absent;

Fork V present(Fig. 19.621); tibial spurs 0, 3, 3; Southeast 'This genus is represented in North America by only one species (see also Table 19A).

Paduniajeanae (Ross)'

M3+4

Figure 19.617

mesepimeron

mesepisternum

Figure 19.618 M3+4

constriction mesepimeron mesocoxa

mesepisternum

Figure 19.619

Figure 19.620

transverse mesocoxa su cus

M1+2 M3+4

Figure 19.621

Figure 19.622

Figure 19.623

M3+4

Figure 19.625

Figure 19.624

Figure 19.617 Glossosoma intermedium {Glossosomatidae) right hind wing, dorsai. Figure 19.618 Agapetus walker!(Glossosomatidae) right hind wing, dorsal. Figure 19.619 Glossosoma intermedium (Glossosomatidae) left mesepisternum, left lateral. Figure 19.620 /Anagapefus bemea (Glossosomatidae) left mesepisternum, left lateral. Figure 19.621 Padunia jeanae (Glossosomatidae) right forewing, dorsal. 688

R.W. Holzenthal & D.R, Robertson 2018

Figure 19.622 Cuioptila hamata (Glossosomatidae) right forewing, dorsal. Figure 19.623 Protoptila macuiata (Glossosomatidae) right forewing, dorsal. Figure 19.624 Protoptila erotica (Glossosomatidae) head and pro- and mesonota (including tegulae), dorsal. Figure 19.625 Cuioptila thoracica (Glossosomatidae) head and pro- and mesonota (including tegulae), dorsal.

Chapter 19 Trichoptera

Figure 19.626

689

Cu1

Figure 19.627

cui

a.

vertex

maxillary palp segment 3

Figure 19.628

Figure 19.629

labial palp vertex

maxillary palp segment 3

maxillary palp

Figure 19.631

labial palp

Figure 19.630 R,W. Holzenthal 2018

Figure 19.626 Goerita semata (Goeridae) right wings, dorsal.

Figure 19.627 Goeracea oregona (Goeridae) right wings, dorsal. Figure 19.628 Goera calcarata (Goeridae) right wings, dorsal.

Figure 19.629 Goerita semata (Goeridae) head and palps, frontal. Figure 19.630 Goeracea genota (Goeridae) head and palps, frontal. Figure 19.631 Goera fuscula (Goeridae) head and palps, frontal.

690

Chapter 19 Trichoptera

4'.

Forewing media vein M with 3 or 4 branches(Fork IV sometimes absent), Fork V absent (Figs. 19.622, 19.623); tibial spurs 0, 3, 3 or 0, 4,4 or 1, 4,4 Forewing media vein M with 4 branches. Fork IV present(Fig. 19.622); spurs 0, 3, 3; male tegulae often huge (Fig. 19.625); Northwest, West

Culoptila

Forewing media vein M with 3 branches, Fork IV absent(Fig. 19.623); spurs 0, 4, 4 or 1, 4,4(foretibial spur hair-like); male tegulae of normal size (Fig. 19.624); widespread

Protoptila

5(4'). 5'.

5

Goeridae

(adapted from the works by Ross 1938, 1944; Schmid 1998)

1.

Ocelli present; tibial spurs 1, 2, 3 on fore-, mid-, and hind tibiae, respectively;

1'. 2(1').

Ocelli absent; tibial spurs 2, 3,4 or 2, 4,4 Forewing discoidal cell longer than thyridial cell (Figs. 19.626, 19.627); eyes small, head vertex forming high crown (Figs. 19.629, 19.630) Forewing discoidal cell much shorter than thyridial cell (Fig. 19.628); eyes much larger, head vertex with smaller crown (Fig. 19.631);

Northwest

2'.

East, Midwest, Northwest

3(2).

3'.

Forewing Cul vein straight between thyridial and subthyridial cells (Fig. 19.626); male maxillary palps held before face, shorter than cylindrical labial palps, each with 1st segment wider apically, 2nd segment quadrate, 3rd segment membranous and tapering to fine filament(Fig. 19.629); East Forewing Cul vein sinuous between thyridial and subthyridial cells (Fig. 19.627); male maxillary palps extended beneath head, shorter than labial palps, all segments cylindrical (Fig. 19.630); Northwest

Lepania cascada Ross' 2 3

Goera

Goerita

Goeracea

Helicopsychidae {Helicopsyche—widespread—is the only genus in North America) Hydrobiosidae (Atopsyche—Southwest—is the only genus in North America) Hydropsychidae (adapted from the works by Ross 1944; Schmid 1998; and Schuster 1984) 1.

r.

2(1). 2'.

Antennae 2-3 times as long as forewings, especially in males; hind wing discoidal cell open(= missing)(Fig. 19.634); head usually with anterior warts large and swollen, posterior warts much smaller (Fig. 19.632) subfamily MACRONEMATINAE,2 Antennae about as long as forewings; hind wing discoidal cell closed by rs crossvein (Fig. 19.635); head with anterior warts small or indistinct, posterior warts large (Fig. 19.633) 3 5-segmented maxillary palps each with second segment much shorter than third (Fig. 19.636); Central, East Macrostemum 5-segmented maxillary palps each with second segment distinctly

longer than third (Fig. 19.637); Texas 3(1'). 3'.

Leptonema albovirens(Walker)'

Forewing postcostal region(PCR,region posterior of looped anal veins) half as long as wing and wider than any wing cells (Fig. 19.635) 4 Forewing postcostal region(PCR)2/3 as long as wing and no wider than wing cells (Fig. 19.648) subfamily HYDROPSYCHINAE,9

This genus is represented in North America by only one species (see also Table 19A).

Chapter 19 Trichoptera

4(3).

4'.

5(4').

5'.

6(5). 6'. 7(5'). 7. 8(7').

Forewings and hind wings each with Fork II stalked(R4 and R5 branching from each other beyond discoidal cell) and about 1/4 as broad basally as apically (Fig. 19.638); Southwest subfamily SMICRIDEINAE,Smicridea Forewings and hind wings each with Fork II sessile(R4 and R5 separating at or before end of discoidal cell) and about 1/2 as broad basally as apically (Fig. 19.635) 5 Antennae thick, each with flagellar segment 2(4th antennal segment) and successive segments each only slightly longer than wide (Fig. 19.639); maxillary palp segment 2 much shorter than segment 3 (Fig. 19.641) subfamily ARCTOPSYCHINAE,6 Antennae slender, each with flagellar segment 2 and successive segments at least twice as long as wide (Fig. 19.640); maxillary palp segment 2 as long as or longer than segment 3(similar to Fig. 19.642). . . .subfamily DIPLECTRONINAE,7 Eyes glabrous (Fig. 19.643); widespread Arctopsyche Eyes hairy (Fig. 19.644); widespread Pampsyche Hind wings with apices of Sc and R1 veins deeply bowed, R1 closer to R2+3 than to SC (Fig. 19.635); widespread Diplectrona Hind wings with apices of SC and R1 straight or slightly curved, R1 equidistant between R2+3 and SC (Figs. 19.645, 19.646) 8 Both pair of wings with apical margins incised; forewings each with origin of discoidal cell level with origin of medial cell (Fig. 19.645);

North Carolina 8'.

9(3'). 9'.

10(9').

10'.

691

Oropsyche howellae Ross'

Both pair of wings with apical margins evenly rounded; forewings each with origin of discoidal cell distal of origin of medial cell (Fig. 19.646); East, West Hind wings each with stem of M about as far from stem of Cu as from stem of R and Fork I absent(Fig. 19.647); widespread Hind wings eaeh with stem of M much closer to stem of Cu than to stem of R and Fork I present (Figs. 19.648, 19.649) Forewing crossveins m-cu and cui-cu2 much further apart than length of either crossvein; hind wing median cell closed by crossvein (Fig. 19.648); adults as in Fig. 10.192; widespread Forewing crossveins m-cu and cu^-cui closely approaching one another; hind wing median cell not closed by crossvein

(Fig. 19.649); Central, East

Homoplectra Cheumatopsyche

10

Hydropsyche

Potamyiaflava(Hagen)'

Hydroptilidae (adapted from the works by Ross 1956; Marshall 1979; Blickle 1979; and Mathis and Bowles 1989) 1.

r.

2(1'). 2'. 3(2).

Foretibiae each with 2 apical spurs; forewings broad with rounded apices, sparsely pubescent, fringes relatively short, discoidal cell and all Forks I-V present(Fig. 19.650); Northeast, Northwest

Foretibiae each with 0 or 1 apical spur; forewings usually narrow and acuminate (Fig. 19.651), sometimes with rounded apices (Fig. 19.652), densely pubescent, fringes long, discoidal cell and at least Fork V absent Foretibiae without spurs Foretibiae each with 1 apical spur Midtibiae each with 1 apical spur; hind tibiae each with 1 apical and 1 preapical spur; forewings with venation indistinct,

costal fringe long (Fig. 19.653); Arkansas

Palaeagapetus

2 3 12

Paucicalcaria ozarkensis Mathis and Bowles'

'This genus is represented in North America by only one species(see also Table 19A).

anterior setal warts anterior setal wart

Figure 19.633 Figure 19.632

posterior setal wart

posterior seta! wart

R1 R2

Figure 19.634

R3

Figure 19.635

SC

R1

Figure 19.638

Figure 19.636

pedicle

fo f3

f4

f5

f6

f7

Figure 19.637

Figure 19.639 scape

f6

pedicle_

^

f2 Figure 19.640 R,W. Holzenthal 2018

Figure 19.632 Macrostemum zebratum (Hydropsychidae) head, dorsal. Figure 19.633 Potamyia flava (Hydropsychidae) head, dorsal.

Figure 19.634 Macrostemum zebratum (Hydropsychidae) right hind wing, dorsal. Figure 19.635 Diplectrona modesta (Hydropsychidae) right wings, dorsal. Figure 19.636 Macrostemum zebratum (Hydropsychidae) maxillary palp. Figure 19.637 Leptonema albovirens (Hydropsychidae) maxillary palp. 692

Figure 19.642

Figure 19.638 Smicridea fasciatella (Hydropsychidae) right wings, dorsal. Figure 19.639 Arctopsycfie grandis (Hydropsychidae) basal segments of an antenna. Figure 19.640 Diplectrona modesta (Hydropsychidae) basal segments of an antenna. Figure 19.641 Arctopsyche grandis (Hydropsychidae) maxillary palp. Figure 19.642 Hydropsyche slossonae (Hydropsychidae) maxillary palp.

Figure 19.644

Figure 19.643

incision

Figure 19.645

Figure 19.646

incision

Figure 19.647

Figure 19.649

Figure 19.648

R.W. Holzentha! 2018

Figure 19.643 Arctopsyche grandis (Hydropsychidae) head, dorsal.

Figure 19.644 Parapsyche elsis (Hydropsychidae) head, dorsal.

Figure 19.645 Oropsyche howelae (Hydropsychidae) right wings, dorsal. Figure 19.646 Homoplectra doringa (Hydropsychidae) right wings, dorsal.

Figure 19.647 Cheumatopsyche analis (Hydropsychidae) right wings, dorsal. Figure 19.648 Hydropsyche slossonae (Hydropsychidae) right wings, dorsal. Figure 19.649 Potamyia flava (Hydropsychidae) right wings, dorsai.

693

694

Chapter 19 Trichoptera

Figure 19.650

Figure 19.651

Figure 19.652

Figure 19.653

)D.R. Robertson 2018

Figure 19.650 Palaeagapetus sp.(Hydroptilidae) right wings, dorsal. Figure 19.651 Metrichia sp.(Hydroptilidae) right wings, dorsal.

3'.

Figure 19.652 Dibusa angata (Hydroptilidae) right wings, dorsal. Figure 19.653 Paucicalcaria ozarkensis (Hydroptilidae) right wings, dorsal (after Mathis and Bowles, 1989).

4(3').

Midtibiae each with 2 or 3 spurs, at least 2 of which are apical; hind tibiae each with 3 or 4 spurs, as least 2 of which are apical; forewings with venation distinct and fringe shorter (Figs. 19.651, 19.652) Hind tibiae each with 1 preapical spur and 2 apical spurs; widespread

4'.

Hind tibiae each with 2 preapical spurs and 2 apical spurs

5

5(4').

Midtibiae each with 0 preapical spurs and 2 apical spurs (North American species only)

6

4 Neotrichia

5'.

Midtibiae each with 1 preapical spur and 2 apical spurs

8

6(5).

Ocelli present

7

6'.

Ocelli absent; adults as in Fig. 10.194; widespread

Hydroptila

- mesoscuteilum

metascutellum-

Figure 19.655

Figure 19.654

mesoscuteilum -

• metascutellum

Figure 19.657

Figure 19.656

mesopostscutellum

mesopostscutellum

Figure 19.658

Figure 19.659

Figure 19.660 © D.R. Robertson 2018

Figure 19.654 Alisotrichia sp.(Hydroptilidae) thorax,

Figure 19.658 Agraylea sp. (Hydroptilidae) thorax,

dorsal.

dorsal.

Figure 19.655 Mayatrichia sp.(Hydroptilidae) thorax,

Figure 19.659 Oxyethira sp.(Hydroptilidae) thorax,

dorsal.

dorsal.

Figure 19.656 Orthotrichia sp.(Hydroptilidae) thorax,

Figure 19.660 Ithytrichia sp.(Hydroptilidae) thorax,

dorsal.

dorsal.

Figure 19.657 Ochrotrichia sp. (Hydroptilidae) thorax, dorsal.

695

antenna! scape

Figure 19.665 Figure 19.662

Figure 19.661

antennal scape

Figure 19.664

Figure 19.666

Figure 19.663

Figure 19.661 Nothotrichia shasta (Hydroptilidae) thorax, dorsal (after Harris and Armltage, 1997). Figure 19.662 Stactobia sp. (Hydroptilidae) thorax,

D.R.Robertson 2018

Figure 19.664

Metrichia sp.(Hydroptilidae) head and

thorax, dorsal.

Figure 19.665 Zumatrichia notosa (Hydroptilidae)

dorsal.

head, left lateral.

Figure 19.663 Leucotrichia sp. (Hydroptilidae) head

Figure 19.666 Leucotrichia sp.(Hydroptilidae) head,

and thorax, dorsal.

left lateral.

696

Chapter 19 Trichoptera

7(6).

Mesoscutellum with transverse suture; metascutellum pentagonal

(Fig. 19.654); Southwest

Alisotrichia arizonica (Blickle and Denning)'

7'.

Mesoscutellum without transverse suture; metascutellum triangular

8(5'). 8'. 9(8').

(Fig. 19.655); widespread Ocelli absent; metascutellum rectangular (Fig. 19.656); widespread Ocelli present; metascutellum pentangular or triangular (Figs. 19.657-19.660) Mesoscutellum with transverse suture (Fig. 19.657); widespread

9'. 10(9').

Mesoscutellum without transverse suture (Figs. 19.658-19.660) Mesopostscutellum (sclerotized region behind mesoscutellum) 1/4 to 1/3 as long on midline as mesoscutellum (Fig. 19.658);

Ocelli absent; East

California

14'.

15(14').

11 Oxyethim Ithytrichia

Dibusa angata Ross'

12'. Ocelli present 13(12'). Mesoscutellum without transverse suture (Fig. 19.661); 13'. 14(13').

Agraylea

Mesopostscutellum obscured on midline by mesoscutellum

(Fig. 19.660); widespread

12(2').

Mayatrichia Orthotrichia 9 Ochrotrichia 10

widespread except not deep Southeast 10'. Mesopostscutellum mostly (Fig. 19.659) or entirely obscured on midline by mesoscutellum (Fig. 19.660) 11(10'). Mesopostscutellum visible on midline behind mesoscutellum (Fig. 19.659); widespread 11'.

697

13

Nothotrichia shasta Harris and Armitage'

Mesoscutellum with transverse suture (Figs. 19.662, 19.663) Metascutellum subrectangular and as wide as mesoscutum (Fig. 19.662); widespread Metascutellum subpentangular to triangular and narrower than mesoscutum (Fig. 19.663) Basal segment of each antenna relatively large, nearly twice as large as other segments, broad, covering half of face (Fig. 19.665);

West

14 Stactobiella

15

Zumati'ichia notosa(Ross)'

15'.

Basal segment of each antenna relatively small, similar in size to other segments, and cylindrical(Fig. 19.666) 16(15'). Wings brilliantly colored, sometimes with green and silver; posterior warts of head transversely linear (Fig. 19.663), head of males sometimes modified with large dorsal protuberances; metascutellum pentagonal (Fig. 19.663); widespread 16'. Wings with dull colors; posterior warts of head transversely elliptical; metascutellum convex anteriorly (Fig. 19.664); Central, Southwest Lepidostomatidae (adapted from the work by Schmid 1998) 1. Posteromesal head warts transversely elliptical, about 2/3 as long as broad (Fig. 19.667); adults as in Fig. 10.195; widespread r. Posteromesal head warts transversely linear, about 1/3 as long as broad (Fig. 19.668); East

Leptoceridae (adapted from the works by Ross 1944, and Schmid 1998) 1. Forewing with stem of M vein atrophied so that thyridial cell absent; hind wing Fork V absent (Fig. 19.669); widespread 'This genus is represented in North America by only one species(see also Table 19A).

16

Leucotrichia Metrichia

Lepidostoma Theliopsyche

Triaenodes

698

Chapter 19 Trichoptera

©R.W. Holzenthal2018

Figure 19.667

posteromesal head wart

Figure 19.667 Lepidostoma quercinum (Lepidostomatidae) head, dorsal.

r. 2(1').

2'. 3(2').

Figure 19.668

posteromesal head wart

Figure 19.668 Theliopsyche corona (Lepidostomatidae) head, dorsal.

Forewing with stem of M vein complete so that thyridial cell present; hind wing with Fork V present(Fig. 19.670) Forewing discoidal cell very long and thyridial cell short, discoidal cell arising basal of thyridial cell and terminating beyond it(19.670); foretibiae each with 2 apical spurs; widespread Forewing thyridial cell very long and arising before discoidal cell (Fig. 19.671); foretibiae each with 0 or 1 apical spur Forewing M vein apparently unbranch, Ml+2 protracted as unbranched, rectilinear extension of stem of M (Fig. 19.672); adults as in Fig. 10.197; widespread

2

Ceraclea 3

Oecetis

3'.

Forewing M conspicuously branched, M1+2 not forming rectilinear

4(3'). 4'. 5(4)

Mesopleural postkatepisternum truncate anterodorsally (Fig. 19.673) 5 Mesopleural postkatepisternum acute anterodorsally (Fig. 19.674) 6 Hind wing with bases of Rs and M obscure or absent(Fig. 19.671); wings pale or white; widespread Nectopsyche Hind wing M vein complete basally although Rs may be incomplete (Fig. 19.675); wings dark brown; Central, East Leptocerus americams(Banks)' Forewing R1 forked 90° and anterior branch ending at notch on costal margin; apical cells all sessile, arising at oblique anastomosis (Fig. 19.676); body and wings generally bluish black; widespread Mystacides

extension of M (Fig. 19.671)

5'. 6(4').

6'.

Forewing R1 not forked; at least some apical cells stalked, not all arising at anastomosis(Fig. 19.677); wings burnt gold with small silver spots: Central, East

4

Setodes

Limnephitidae (adapted from the work by Ruiter 2000) 1.

1'. 2(1').

Eorewing anastomosis of apical forks and crossveins forming 1 nearly straight line perpendicular to longitudinal veins (Fig. 19.678); West Forewing anastomosis separated into two parts, posterior part more nearly basal (Fig. 19.679) Hind wing Fork I petiolate (Fig. 19.679); West

'This genus is represented in North America by only one species(see also Table 19A).

Homophylax 2

Cryptochia

Chapter 19 Trichoptera

699

M3+4

Figure 19.669

Ml+2 M3+4

Cu1a+b

Figure 19.670

R4+5

M1+2 M3+4

Figure 19.671

Figure 19.672

mesopleural postkatepisternum

mesopleural

postkatepisternum

Figure 19.673

Figure 19.674

Figure 19.669 Triaenodes tardus (Leptoceridae) right wings, dorsal. Figure 19.670 Ceraclea cancellata (Leptoceridae) right wings, dorsal. Figure 19.671 Nectopsyche albida (Leptoceridae) right wings, dorsal.

© R.W. Holzentha! 2018

Figure 19.672 Oecetis inconspicua (Leptoceridae) right forewing, dorsal. Figure 19.573 Nectopsyche albida (Leptoceridae) mesopleuron, left lateral.

Figure 19.674 Mystacides interjectus (Leptoceridae) mesopleuron, left lateral.

700

Chapter 19 Trichoptera

Figure 19.675

Cu2

Figure 19.676

Figure 19.677

© R.W. Holzenthal 2018

Figure 19.675 Leptocerus americanus (Leptoceridae) right wings, dorsai. Figure 19.676 Mystacides interjectus (Leptoceridae) right wings, dorsai.

Figure 19.677 Setodes oligius (Leptoceridae) right wings, dorsai.

anastomosis

Figure 19.679

Figure 19.678

petiole

anastomosis

anastomosis

M3+4 Cu1a+1b

Figure 19.681 Figure 19.680

M3+4 Cu1a+1b Cu1a+1b

R2+3, R4+5

R2+3, R4+5

fork I root

Figure 19.682

Figure 19.683

©R.W. Holzenthal 2018

Figure 19.678 Homophylax flavipennis (Limnephllidae) right wings, dorsal. Figure 19.679 Cryptochia pilosa (Limnephllidae) right wings, dorsal. Figure 19.680 Limnephilus samoedus (Limnephllidae) right wings, dorsal.

Figure 19.681 Sphagnophylax meiops (Limnephllidae) right wings, dorsal. Figure 19.682 Ecclisocosmoecus scylla (Limnephilldae) right wings, dorsal. Figure 19.683 Ecclisomyia maculosa (Limnephilldae) right wings, dorsal. 701

fork I root

Figure 19.684 Figure 19.685

Figure 19.687

anepisternal

Figure 19.686

mesepisternal

wart

wart

m-cu

infraepisternal wart M3+4 coxa wart

Figure 19.688

mesepisternal

Figure 19.689

wart

second anal cell

=—M3+T infraepisternal m-cu—^

wart

coxa wart M3+4

R.W. Holzenthal 2018

Figure 19.684 Ironoquia punctatissima (Limnephilidae) right wings, dorsai. Figure 19.685 Philocasca rivularls (Limnephilidae) right wings, dorsal. Figure 19.686 Amphicosmoecus canax (Limnephilidae) right wings, dorsal.

702

Figure 19.687 Dicosmoecus atripes (Limnephilidae) left mesopleuron, left lateral. Figure 19.688 Onocosmoecus unicolor (Limnephilidae) left mesopleuron, left lateral. Figure 19.689 Onocosmoecus unicolor (Limnephilidae) right wings, dorsal.

Chapter 19 Trichoptera

703

2'.

Hind wing Fork I sessile or rooted on discoidal cell, with R3-discoidal

3(2').

Hind wing Fork V absent(Fig. 19.680)

4

3'.

Hind wing Fork V present (Fig. 19.683)

5

4(3').

Forewings long, narrow; R1 extending far beyond anastomosis (Fig. 19.680); Northwest, Michigan Limnephilus samoedus(McLachlan)

4'.

Forewings short, broad; R1 ending at wing margin in line with anastomosis (Fig. 19.681); Northwest Territory,

5(3').

Forewing discoidal cell with R2+3 and R4+5 parallel and very close

cell concurrent for variable distance (Fig. 19.680)

Yukon Territory

3

Sphagnophylax meiops Wiggins and Winchester'

together in basal half of cell (Fig. 19.682); Northwest 5'. 6(5').

6'. 7(6').

7'.

8(7).

8'.

9(8'). 9'. 10(9).

Forewing discoidal cell with R2+3 and R4+5 as widely separated as for most other longitudinal veins (Fig. 19.683) Forewing Fork I deeply rooted on discoidal cell, having common vein with apical part of discoidal cell longer than 3-5 times breadth of discoidal cell (Fig. 19.683); West

Forewing Fork I well rooted on discoidal cell, having common vein with apical part of discoidal cell longer than breadth of discoidal cell (Fig. 19.684); Central, East

Forewing Fork I rooted on discoidal cell much less, having common vein with apical part of discoidal cell much less than breadth of discoidal cell (Fig. 19.686) Mesopleuron with anepisternal wart present(Fig. 19.687) Mesopleuron without anepisternal wart(Fig. 19.688) Tibial spurs 1, 2, 2 on pro-, meso-, and metatibiae, respectively,

12(11').

13

Ironoquia

9 10 11

Allocosmoecus partitus Banks' Dicosmoecus

Midtibiae each with 2 apical spurs, but no pre-apical spur Midtibiae each with 2 apical spurs and a pre-apical spur (similar to Fig. 19.583)

Amphicosmoecus canax (Ross)' 12

Forewings orange, with large imprecise brown areas(Fig. 19.689); North, West

12'. 13(7').

8

Tibial spurs 1, 2, 2 or 1, 3, 4; if 1, 2, 2, the spurs and spines dark,

(similar to Fig. 19.560); West 11'.

7

Forewing thyridial cell distal margin strongly oblique to cell length, comprised primarily of m-cu crossvein, this crossvein originating at or slightly basal of fork of M (Fig. 19.684) Forewing thyridial cell distal margin nearly perpendicular to cell length (Fig. 19.685) or m-cu crossvein originating on M3+4 beyond fork of M (Fig. 19.691)

concolorous; West

11(9').

6

EccUsomyia

Forewing Fork I subsessile, having common vein with discoidal cell much shorter (Fig. 19.684)

these spurs paler than leg spines; Northwest 10'.

Ecclisocosmoecus scylla (Milne)'

Forewings uniformly dark brown (Fig. 19.690); Northwest Forewing « crossvein strongly bent(Fig. 19.685); Northwest

13'. Forewing crossvein straight (Fig. 19.691) 14(13'). Hind wing RS + discoidal cell length shorter or equal to distance from discoidal cell to wing tip (Fig. 19.691); Central, East, Northwest 'This genus is represented in North America by only one species(see also Table 19A).

Onocosmoecus

Eocosmoecus Philocasca 14

Pseudostenophylax

M2

second anal cell

Figure 19.690

M3+4

Figure 19.691

R2

R3

Figure 19.692

Figure 19.693

petiole

anterior anastomosis

Figure 19.694 Figure 19.695

© R.W. Holzenthal 2018

Figure 19.690 Eocosmoecus frontalis (Limnephilidae) right wings, dorsal. Figure 19.691 Pseudostenophylax sparsus (Limnephilidae) right wings, dorsal. Figure 19.692 Halesochila taylori (Limnephilidae) right wings, dorsal. 704

Figure 19.693 Phanocelia canadensis (Limnephilidae) right wings, dorsal. Figure 19.694 Glyphopsyche irrorata (Limnephilidae) right wings, dorsal. Figure 19.695 Leptophylax gracilis (Limnephilidae) right wings, dorsal.

R1 R2

Figure 19.696

Figure 19.697

anterior anastomosis

Figure 19.699 Sc+R1

Figure 19.700

Figure 19.701

R.W. Holzenthal 2018

Figure 19.696 Nemotaulius hostilis (Limnephilidae) right wings, dorsal. Figure 19.697 Monophylax mono (Limnephilidae) right wings, dorsal. Figure 19.698 Frenesia missa (Limnephilidae) right wings, dorsal.

Figure 19.699 Chilostigmodes areolatus (Limnephilidae) right wings, dorsal. Figure 19.700 Desmona bethula (Limnephilidae) right wings, dorsal. Figure 19.701 Psychoglypha subborealis (Limnephilidae) right wings, dorsal. 705

706

14'.

Chapter 19 Trichoptera

Hind wing RS + discoidal cell length greater than distance from discoidal cell to wing tip (Fig. 19.692)

15

15(14'). Forewing discoidal cell membrane clear and all veins surrounding it dark (Fig. 19.692); Northwest Halesochila taylori(Banks)' 15'. 16(15').

Forewing discoidal cell usually not clear; if clear, some veins surrounding discoidal cell also clear Forewing Fork III with long petiole, about half as long as Fork III

(Fig. 19.693); North 16'.

16

Phanocelia canadensis (Banks)'

Forewing Fork III petiole absent (Figs. 19.694, 19.695) or much shorter

17

17(16'). Forewing R1 and R2 parallel and strongly bowed posterad around pterostigma (Fig. 19.694)

18

17'.

Forewing R1 and R2 often divergent apically and always more nearly straight

18(17).

Forewing Forks I—III sessile, so that anterior portion of anastomosis linear, perpendicular to wing length (Fig. 19.694) Forewing Forks l-III slightly rooted, so that anterior portion of anastomosis zigzagged (Fig. 19.697) Forewings each with partial or complete r crossvein between R1 and R2

(Fig. 19.696)

18'. 19(18).

25

at base of pterostigma (Fig. 19.694); Central, North 19'.

19 20

Glyphopsyche

Forewings without r crossveins(Fig. 19.698); East

Frenesia

20(18'). Forewing thyridial cell membrane completely darkened (Fig. 19.697), occasionally finely irrorate

20'.

Forewing thyridial cell with two contrasting dark and hyaline areas

21(20).

(Fig. 19.699), dark areas occasionally finely maculate Forewing Fork II membrane hyaline at base (Fig. 19.697);

West

21 23

Monophylax mono(Denning)'

21'. Forewing Fork II membrane completely dark (Fig. 19.700) 22(21'). Forewing Fork III membrane completely hyaline (Fig. 19.700); California

22'.

22 Desmona

Forewing Fork III membrane partially or completely darkened (Figs. 10.199, 19.701); some species with dark areas irrorate; North,

West Psychoglypha Hind wing r-m crossvein present (Fig. 19.702); Northwest Grensiapmeterita (Walker)' Hind wing r-m crossvein absent or very short(Figs. 19.699, 19.703) 24 Hind wing Sc and R1 fused near apex (Fig. 19.699); North Chilostigmodes areolatus (Walker)' Hind wing Sc and R1 separate to wing margin (Fig. 19.703); Minnesota Chilostigma itascae(Wiggins)' 25(17'). Forewings each with postapical margin excised, sinuous (Fig. 19.696); North Nemotaulius hostilis(Hagen)' 25'. Forewings with margins straight or convex (Fig. 19.704) 26 26(25'). Hind wing R5 distinctly darker than other veins (Fig. 19.704); North Grammotaulius 26'. Hind wing R5 not darker than other veins (Fig. 19.706) 27 27(26'). Forewings each with A2 vein atrophied apically so that apical anal 23(20'). 23'. 24(23'). 24'.

cell undivided (Fig. 19.705); wings with distinctive pattern of various shades of brown ranging from almost cream color to chocolate; Central, East 'This genus is represented in North America by only one species(see also Table 19A).

Platycentropus

Figure 19.702

z Figure 19.703

Figure 19.704 2nd & 3rd anal cells undivided

Figure 19.705

2nd anal cell arculus 3rd anal cell

Figure 19.706

2nd anal cell

Figure 19.707

arculus

©R.W. Holzenthal2018

Figure 19.702 Grensia praeterita (Limnephilidae) right wings, dorsal. Figure 19.703 Chilostigma itasca (Limnephilidae) right wings, dorsal.

Figure 19.704 Grammotaulius lorretae (Limnephilidae) right wings, dorsal.

Figure 19.705 Platycentropus amicus (Limnephilidae) right wings, dorsal.

Figure 19.706 Clostoeca disjuncta (Limnephilidae) right wings, dorsai. Figure 19.707 Hydatophylax argus (Limnephilidae) right wings, dorsai. 707

R1

R2

Figure 19.708

Figure 19.709

Figure 19.710

Figure 19.711

Figure 19.712

Figure 19.713

posterior anastomosis

© R.W. Ho zentha 2018

Figure 19.708 Anabolia bimaculata (Limnephilidae) right wings, dorsal. Figure 19.709 Chyranda centralis (Limnephilidae) right wings, dorsal. Figure 19.710 Pycnopsyche scabripennis (Limnephilidae) right wings, dorsal. 708

subradial cell

- posterior anastomosis

Figure 19.711 Arctopora trimaculata (Limnephilidae) right wings, dorsal. Figure 19.712 Limnephilus rhombicus (Limnephilidae) right wings, dorsal. Figure 19.713 Hesperophylax magnus (Limnephilidae) right wings, dorsal.

Chapter 19 Trichoptera

Forewings each with A2 vein complete, defining 2 apical anal cells (Fig. 19.706); wing pattern different 28(27'), Hind wings each with small sc-r crossvein between Sc and R1 near their apices (Figs. 19.706, 19.707) or Sc and R1 fused for short distance

709

27'.

28'. 29(28).

30(28').

30'. 31(30).

30

Clostoeca disjuncta (Banks)'

Forewing second anal cell acute apically, about 25°, reaching beyond half distance to arculus(Fig. 19.707); widespread except not Southwest Forewing R1 straight or nearly so from base of pterostigma to wing margin (Fig. 19.708)

Hydatophylax 31 34

Forewing R1 bowed posterad around pterostigma (Fig. 19.709) Forewing Fork I moderately rooted, so that R3-discoidal cell common boundary at least as long as discoidal cell breadth (Fig. 19.695);

Central 31'.

29

Hind wings with Sc and R1 completely separate (Fig. 19.695) Forewing second anal cell blunt apically, about 65°, reaching less

than half distance to arculus (Fig. 19.706); Northwest 29'.

28

Leptophylax gracilis Banks'

Forewing Fork I only slightly rooted, so that R3-discoidal cell common boundary less than discoidal cell breadth (Fig. 19.708)

32

32(31'). Forewing discoidal cell more than twice as long as RS (Fig. 19.708); widespread

32'.

Anabolia 33

Forewing discoidal cell less than twice as long as RS (Fig. 19.710)

33(32'). Forewings each less than 10 mm long; Northwest

Philarctus bergrothi McLachlan'

33'. Forewings each greater than 10 mm long; Central, East, North 34(30'). Hind wing r-m crossvein distinctly longer than base of Ml (Fig. 19.709);

North, West 34'.

Pycnopsyche

Chyranda centralis(Banks)'

Hind wing r-m crossvein no longer than base of Ml,often shorter

(Fig. 19.711) 35(34'). Forewing discoidal cell shorter than RS (Fig. 19.711); North 35'. Forewing discoidal cell as long as, or longer than RS (Fig. 19.712) 36(35').

Hind wing discoidal cell shorter than RS, or posterior part of hind wing anastomosis nearly perpendicular to wing length,

usually both (Fig. 19.712); widespread except not deep Southeast 36'.

35 Arctopora 36

Limnephilus (in part)

Hind wing discoidal cell as long as, or longer than RS,or posterior part of hind wing anastomosis strongly oblique (Fig. 19.713)

37(36'). Forewing subradial cell with linear stripe of fine, white hairs (Fig. 19.713); North, West

37'. Forewing subradial eell without linear stripe (Fig. 19.714) 38(37'). Hind wing r-m crossvein strongly oblique to rs crossvein (Fig. 19.714) 38'. Hind wing r-m and rs crossveins sub-parallel, although offset(Figs. 19.715, 19.716) 39(38). 39'. 40(39).

37 Hesperophylax

38 39 42

Hind wing r-m crossvein with anterior end at or near fork of R4-I-5, conspicuously basal of crossvein (Fig. 19.714)

40

Hind wing r-m crossvein with anterior end on R5, distal of fork of R4-f5, only slightly basal of rs crossvein (Fig. 19.717)

41

Forewings with large, hyaline blotches (Fig. 19.714); West

'This genus is represented in North America by only one species(see also Table 19A).

Psychoronia

710

Chapter 19 Trichoptera

Figure 19.714

subradiat cell

R3

Figure 19.715

R4

Ml /m

Figure 19.716

Figure 19.717

Figure 19.718 Figure 19.719

©R.W. Holzenthal2018

Figure 19.714 Psychoronia brooksi(LImnephilidae) right wings, dorsal. Figure 19.715 Lenarchus brevipennis (Limnephilidae) right wings, dorsal. Figure 19.716 Limnephilus fumosus (Limnephilidae) right wings, dorsal.

Figure 19.717 Crenophylax sperryi (Limnephilidae) right wings, dorsal. Figure 19.718 Asynarchus montanus (Limnephilidae) right wings, dorsal. Figure 19.719 Clistoronia maculata (Limnephilidae) right wings, dorsal.

Chapter 19 Trichoptera

40'.

Forewings irrorate; Northcentral, Northeast

41(39').

Forewings with large, hyaline blotches (Fig. 19.717);

Asynarchus rossi(Leonard and Leonard)

Southwest

Crenophylax spenyi(Banks)'

41'.

Forewings brown, at most finely maculate (Fig. 19.718); North

42(38').

Flind wing r-m crossvein about half as long as rs crossvein (Fig. 19.715);

Asynarchus (in part)

North, West

42'.

Lenavchus

Hind wing r-m crossvein about as long as rs crossvein (Fig. 19.716)

43

43(42'). Forewing discoidal cell more than 2.5 times as long as RS (Fig. 19.716); Northwest, West 43'.

711

Limnephilus(mpaxt)

Forewing discoidal cell less than 2.5 times as long as RS(Fig. 19.719); West

Clistoronia

Molannidae

1.

Forewing R1 fused with R2+3 near anastomosis; Cu2 terminating before reaching wing margin (Fig. 19.720); adults as in Fig. 10.198; Central, East

r.

Forewing R1 independent to wing margin; Cu2 reaching wing

margin at arculus (Fig. 19.721); far Northwest

Molanna

Molannodes tinctus (Zetterstedt)'

Odontoceridae

1.

Forewings each with 4-6 crossveins in the costal cell between C and Sc(Fig. 19.722)

2

1'.

Forewings each without crossveins in costal cell (Fig. 19.723)

3

2(1).

Forewing R1 fused with R2 before apex (Fig. 19.722); California,

Oregon 2'.

Namamyiaplutonis Banks'

Forewing R1 independent to wing margin (Fig. 19. 724); California,

Oregon 3(1').

Nerophilus californicus (Hagen)'

Forewings 4 times as long as broad (Fig. 19.723); eyes of male very large, nearly touching on vertex; Central, Southwest, and Vermont

3'.

4(3').

Forewings no more than 3.2 times as long as broad (Figs. 19.725-19.727); eyes separated by distance greater than width of one eye in dorsal view

5(4').

5'.

4

Forewing Fork II sessile or rooted on discoidal cell; male with only Forks I and II(Fig. 19.725), female with Forks I-Y;

Southern Appalachian Mtns 4'.

Marilia

Pseudogoem singularis Carpenter'

Forewing Fork II stalked, its pedicel 25-40% as long as Fork (Figs. 19.726, 19.727) Forewings each with R1 and R2 approximate, nearly touching at wing margin, apparently with discoidal cell and looped anal veins present, male with Forks I, II, and V (Fig. 19.726), female with Forks I, IL HI, and V (not illustrated); East Forewings each with R1 and R2 parallel or slightly diverging apically, at least as far apart as apices of other veins, apparently lacking discoidal cell and looped anal veins(Fig. 19.727, male illustrated; female wings unknown); West

'This genus is represented in North America by only one species(see also Table 19A).

5

Psilotreta

Parthina

712

Chapter 19 Trichoptera

R1+2+3

Sc R1

Figure 19.720

Cu1b

R1+2

Figure 19.721

C3D.R. Robertson 2018

Figure 19.720 Molanna flavicornis (Molannidae) male, right wings, dorsal.

Figure 19.721 Molannodes tinctus (Molannidae) male, right wings, dorsal.

Philopotamidae 1. Wings highly reduced, non-functional in flight (Fig. 19.728). Micropterous females of winter populations of D. distincta (Walker); Central, East Dolophilodes distincta (Walker) r

Wings normally developed, functional in flight

2(1').

Tibial spurs 1, 4, 4 on each of the fore-, mid-, and hind tibiae,

respectively; forewing Fork IV absent(Fig. 19.729); widespread 2'.

Tibial spurs 2, 4, 4; forewing Fork IV present(Fig. 19.730)

3(2').

Forewing Fork I petiolate, originating from common stem beyond discoidal cell (Fig. 19.731); widespread Forewing Fork I sessile, originating on apex of discoidal cell (Fig. 19.732)

3'. 4(3').

Hind wings each with 2A atrophied apically so that only 3 veins reaching wing margin posterior of Fork V (Fig. 19.730; veins may be faint, so look closely); widespread

4'.

Hind wings each with 4 veins reaching wing margin posterior of Fork V (Fig.(Fig. 19.731) Forewings each with r-m and m crossveins nearly aligned with each other; also Cu2 and A1+2+3 terminating close together on hind margin; hind wings each with crossvein between veins

5(4').

2A and 3A (Fig. 19.732); Southeast(mountains) 5'.

2

Chimarm 3 Dolophilodes (in part) 4

Wormaldia 5

Fumonta major(Banks)'

Forewings each with r-m and m crossveins separated by distance as great as or greater than length of one crossvein; also veins Cu2 and A1+2+3 terminating separately on hind margin; hind wings lacking crossvein between veins 2A and 3A (Fig. 19.733); West

' This genus is represented in North America by only one species(see also Table 19A).

Sisko

Chapter 19 Trichoptera

713

costal cell crossveins R1+fR2

Figure 19.722

Figure 19.723

Figure 19.724

Figure 19.725

Figure 19.727

Figure 19.726

Cu1bCu1a ©R.W, Holzenthal 2018

Figure 19.722 Namamyia plutonis (Odontoceridae) right wings, dorsal. Figure 19.723 Marilia flexuosa (Odontoceridae) male, right wings, dorsal. Figure 19.724 Nerophilus californicus (Odontoceridae) right wings, dorsal.

Figure 19.725 Pseudogoera singularis (Odontoceridae) male, right forewing, dorsal. Figure 19.726 Psilotreta labida (Odontoceridae) male, right wings, dorsal. Figure 19.727 Parthina linea (Odontoceridae)female, right wings, dorsal.

714

Chapter 19 Trichoptera

Figure 19.729

Figure 19.728

Figure 19.730

Figure 19.731

Figure 19.732

AI+2+3

Cu2

Figure 19.733

A1+2+3 Cu2

© R.W. Holzenthal & D.R. Robertson 2018

Figure 19.728 Dolophilodes distincta (Philopotamidae)female habitus, left lateral. Figure 19.729 Chimarra obscura (Philopotamidae) right wings, dorsal. Figure 19.730 Wormaldia occidea (Philopotamidae) right wings, dorsal.

Figure 19.731 Dolophilodes novusamericana (Philopotamidae) right wings, dorsal. Figure 19.732 Fumonta major (Philopotamidae) right wings, dorsal. Figure 19.733 Sisko sisko (Philopotamidae) right wings, dorsal.

Chapter 19 Trichoptera

715

Phryganeidae (based on the monograph by Wiggins 1998) 1.

Hind wings mostly uniformly dark brown except for transverse yellow band near apex from costal margin to at least Cu2(Fig. 19.734); East except not deep Southeast

r.

2(1').

Oligostomis

Hind wings light brown or mostly light brown and without transverse yellow band near apex (Fig. 19.735)

2

Hind wings narrow (width/length ratio about 0.45), with anterior and posterior margins nearly symmetrical on longitudinal axis and with apices more nearly pointed (Fig. 19.736); California,

Oregon 2'.

3(2')

Yphria californica(Banks)'

Hind wings broader (width/length ratio 0.50 or more), with anterior margins more nearly straight and posterior margins more convex (especially basally), and apices more rounded (Fig. 19.737)

3

Head with pair of posterior setal warts, but no anterior warts (anterior of lateral ocelli and between antennae)(Fig. 19.738);

Northcentral, Northeast

Beothukus complicatus(Banks)'

3'.

Head with both pairs of warts (Fig. 19.739)

4(3').

Forewings with dark speckles and longitudinal brown stripes (Fig. 19.740); widespread

4'. 5(4'). 5'.

6(5'). 6'. 7(6').

Phryganea

Forewings with or without speckles but lacking conspicuous longitudinal stripes (Figs. 19.735, 19.741, 19.744) Forewings with large irregular brown confluent reticulations (Fig. 19.741); widespread

5 Banksiola

Forewings without reticulations (Fig. 19.742) or reticulations smaller and discrete (Figs. 19.735, 19.744)

6

Hind wing m-cu crossvein strongly recurved with its anterior portion long and sometimes with short spur (Fig. 19.737); widespread

Agrypnia

Hind wing m-cu not so strongly curved (Fig. 19.743) or with its anterior portion much shorter than posterior portion, never with short spur (Fig. 19.744)

7

Forewings lacking c-sc crossvein near midlength of wing (Fig. 19.743);

North 7'.

Forewings with c-sc crossvein at midlength (Fig. 19.744)

8(7').

Hind wing m-cu crossvein recurved at least 90°(Fig. 19.744), often with dark V-shaped mark before apex (Fig. 19.744); widespread except not Southwest

8'.

4

Hind wing m-cu crossvein bent less than 90°(m-cu crossvein similar to that of Fig. 19.743)

FaAr/fl moraaffl(Banks)' 8

Ptilostomis 9

9(8').

Northcentral, Northeast

Hagenella canadensis(Banks)'

9'.

Alaska, Yukon Territory

Oligotvicha lapponica (Hagen)'

'This genus is represented in North America by only one species (see also Table 19A).

716

Chapter 19 Trichoptera

Figure 19.734

Figure 19.735

Figure 19.736

Figure 19.737

m-cu

A. pagetana

Figure 19.740

brown stripes anterior

Figure 19.738 posterior setal wart lateral ocellus

posterior setal wart

5R.W. Hoizenthai20i8

Figure 19.734 Oligostomis pardalis (Phryganeidae) female, right wings, dorsal. Figure 19.735 Agrypnia vestita (Phryganeidae) male, right wings, dorsal. Figure 19.736 Yphria californica (Phryganeidae) male, right wings, dorsal. Figure 19.737 Agrypnia coiorata (Phryganeidae) male, right wings, dorsal, with inset of m-cu crossvein.

Figure 19.739

Figure 19.738 Beothukus complicatus (Phryganeidae) head, dorsal.

Figure 19.739 Phryganea cinerea (Phryganeidae) head, dorsal.

Figure 19.740 Phryganea cinerea (Phryganeidae) female, right wings, dorsal.

Chapter 19 Trichoptera

717

Figure 19.741

Figure 19.742

c-sc

Figure 19.743

Figure 19.744

R.W. Holzenthal 2018

Figure 19.741 Banksiola crotchi(Phryganeidae) female, righit forewing, dorsal. Figure 19.742 Agrypnia glacialis (Phryganeidae) male, right wings, dorsal.

V-shaped mark

Figure 19.743 Fabria inornata (Phryganeidae)female, right wings, dorsal. Figure 19.744 Ptilostomis ocellifera (Phryganeidae) male, right wings, dorsal.

718

Chapter 19 Trichoptera

Polycentropodidae (based on revision by Chamorro and Holzenthal 2011) 1. 1'. 2(1'). 2'.

3(2'). 3'.

4(3').

Foretibiae each with two apical spurs, but no preapical spur; Central, East Foretibiae each with preapical spur in addition to two apical spurs Hind wing Fork III present and 3A straight, or nearly so(Fig. 19.745); widespread except not Southwest Hind wing Fork 111 usually absent(Fig. 19.746), but if present, with 3A curved medially, towards 2A (Fig. 19.748) Forewing anal veins fusing at nearly the same point(Fig. 19.746); widespread except not Southwest Forewing veins 2A and 3A fused far basal of their fusion with lA (Fig. 19.747) Forewing Cu2 recurved more than 90° at arculus(Fig. 19.747);

Southwest 4'. 5(4'). 5'. 6(5).

6'. 7(5'). 7.

Cernotina 2

NeurecUpsis 3

Nyctiophylax 4

Polyplectropus

Forewing Cu2 bent less than 90° at arculus (Fig. 19.748) Hind wing Fork I present (Figs. 19.748, 19.749) Hind wing Fork I absent(Figs. 19.750, 19.751) Hind wing discoidal cell closed by rs crossvein (Fig. 19.748); widespread except not Southwest Hind wing discoidal cell open {rs crossvein absent)(Fig. 19.749); widespread Hind wing discoidal cell closed by rs crossvein (Fig. 19.750); Central, North Hind wing discoidal cell open, rs crossvein absent(Fig. 19.751);

5 6 7 Plectrocnemia

Polycentropus Holocentropus

Central, East

Cymellusfraternus(Banks)'

Psychomyiidae (adapted from the works by Flint 1967, and Schmid 1998) 1.

1'. 2(1'). 2'.

3(2').

Maxillary palps each 6-segmented, labial palps each 4-segmented; forewing discoidal cell open, rs crossvein absent(Fig. 19.752); Arkansas, Missouri(Ozarks) Paduniella nearctica Flint' Maxillary palps each 5-segmented, labial palps each 3-segmented; forewing discoidal cell closed by rs crossvein (Fig. 19.753) 2 Hind wing Fork 111 absent, with M forked into 2 branches(Fig. 19.753); widespread Psychomyia Hind wing Fork 111 present, with M forked into 3 branches (Figs. 19.754, 19.755)

3

Forewings each without sc-r crossvein; hind wings each with

Fork 111 petiolate (Fig. 19.754); Central, East 3'.

Lype diversa (Banks)'

Forewings each with sc-r crossvein; hind wings each with Fork 111 sessile (Fig. 19.755); West

Tinodes

Rhyacophilidae (adapted from the work by Schmid 1998) 1. 1'.

Mesoscutellum without long hairs; forewings each less than 20 mm long; adults as in Fig. 10.191; widespread Mesoscutellum with tuft of long, fine hairs; forewings each more

than 20 mm long; Pacific states 'This genus is represented in North America by only one species(see also Table 19A).

Rhyacophila

Himalopsychephryganea (Ross)'

Sc R1 R9

Figure 19.745

arculus

ri

Figure 19.746 Sc

Figure 19.747

R1

arculus

Sc

R1

Figure 19.748 arculus Sc R1

Figure 19.749

arculus

2A

1A

Frgure 19.750 arculus

Figure 19.751

arculus

Sc

RI

Sc__^d2+3

D.R. Robertson 2018

Figure 19.745 Neureclipsis bimaculata (Polycentropodidae) right wings, dorsai. Figure 19.746 Nyctiophylax moestus (Poiycentropodidae) right wings, dorsal. Figure 19.747 Polyplectropus charlesi (Polycentropodidae) right wings, dorsai. Figure 19.748 Plectrocnemia cinerea (Polycentropodidae) right wings, dorsal.

Figure 19.749 Polycentropus colei (Polycentropodidae) right wings, dorsal. Figure 19.750 Holocentropus interruptus (Polycentropodidae) right wings, dorsal. Figure 19.751 Cyrnellus fraternus (Polycentropodidae) right wings, dorsal.

719

720

Chapter 19 Trichoptera

Figure 19.752

Figure 19.753 R4+5

M1+2 M3+4

Figure 19.754

Figure 19.755 M3+4

5 R.W. Holzenthal 2018

M3+4

Figure 19.752 Paduniella nearctica (Pschomyiidae) right wings, dorsal. Figure 19.753 Psychomyia flavida (Psychomyiidae) right wings, dorsal.

Figure 19.754 Lype diversa (Psychomyiidae) right wings, dorsal.

Figure 19.755 Tinodes cascadius (Psychomyiidae) wings, dorsal.

Rossianidae

1.

1'.

Forewings and hind wings each with Fork I deeply rooted on discoidal cell, with R3-discoidal cell concurrent for most of length of discoidal cell, and with discoidal cell lOX as long as broad and originating near first division of M (Fig. 19.756); Northwest

Rossiana montana

Forewings and hind wings each with Fork I subsessile, with R3-discoidal cell concurrent for short distance, and with discoidal cell 6X as long as broad and originating far basal of first division of M (Fig. 19.757); Northwest

Goereilla baumanni

Sericostomatidae

1.

East and Central North America

r. 2(1)

West Male maxillary palps 2-segmented, each with long, clavate process arising apically and tiny second segment arising subapically from large first segment(Fig. 19.764); above 600 m elevation in

Southern Appalachian Mountains 'This genus is represented in North America by only one species(see also Table 19A).

2

Gumaga

Fattigiapele (Ross)^

Chapter 19 Trichoptera

721

Figure 19.756

Figure 19.757

Figure 19.758

Figure 19.759

Cu1a+b M1+2 M3+4

R.W. Ho zentha 2018

Figure 19.756 Rossiana montana (Rossianidae) right wings, dorsal. Figure 19.757 Goereilla baumanni(Rossianidae) right wings, dorsal.

Cu1a+b

Figure 19.758 Neophylax oligius (Thremmatidae) right wings, dorsal. Figure 19.759 Oligophlebodes minutus (Thremmatidae) right wings, dorsal.

722

Chapter 19 Trichoptera

M3+4

Figure 19.760 M3+4

Figure 19.761

Cu1a+b

Cula+b

Figure 19.762

R4+5

Figure 19.763

R1

R.W. Holzenthal 2018

Figure 19.760 Neothremma didactyla (Uenoidae) right wings, dorsal. Figure 19.761 Sericostriata surdickae (Uenoidae) right wings, dorsal.

Figure 19.762 Cnodocentron sp.(Xiphocentronidae) right wings, dorsal. Figure 19.763 Xiphocentron sp.(Xiphocentronidae) right wings, dorsal.

Chapter 19 Trichoptera

Figure 19.764

Figure 19.764 Fattigia pele (Sericostomatldae) male maxillary palp, inner face (after Ross and Wallace, 1974). 2'.

723

Figure 19.765

Figure 19.765 Agarodes griseus (Sericostomatidae) male maxillary palp, inner face (after Ross and Wallace, 1974).

Male maxillary palps 2-segmented, each with two slender processes arising from near base of large first segment, tiny second segment arising near apex of one of them (Fig. 19.765); below 500 m elevation in East and Central North America as far west as Minnesota and eastern Texas

. Agarodes

Thremmatidae

(based on the work by Schmid 1998)

1. 1'.

Hind wing Fork I present(Fig. 19.758); widespread except not Southwest Hind wing Fork I absent(Fig. 19.759); West

Neophylax OUgophlebodes

Uenoidae

(based on the work by Schmid 1998) 1. Tibial spurs 2, 4,4 on fore-, mid-, and hind tibiae, respectively; Pacific states r. Tibial spurs 1, 3, 4 2(1') Forewings each with Fork I subsessile, confluent with discoidal cell for short distance (Fig. 19.760); hind wings each with Fork I stalked, branching beyond discoidal cell, this discoidal cell short, originating in middle of wing (Fig. 19.760); West 2'. Forewings and hind wings each with Fork I rooted, confluent with discoidal cells for about half length of discoidal cell (Fig. 19.761); discoidal cells long and originating near base of wing (Fig. 19.761);

Idaho, Montana

Farula 2

Neothremma

Sericostriata surdickae Wiggins, Weaver, and Unzicker'

Xiphocentronidae (based on the revision by Schmid 1982)

1.

Hind wings each with R1 independently terminating in wing margin

(Fig. 19.762); Arizona T.

Cnodocentron yavapai Moulton and Stewart'

Hind wings each with R1 fused apically with R2+3(Fig. 19.763);

Texas 'This genus is represented in North America by only one species(see also Table 19A).

Xiphocentron messapus Schmid'

724

Chapter 19 Trichoptera

SELECTED ADDITIONAL TAXONOMIC REFERENCES

(L = larvae; P = pupae; A = adults) General Ross (1944)-L, P, A,(1956)-A,(1959)-L; Wiggins(1977, 1996)-L, (1982)-L, A,(2004)-L,P A; McCafferty (1981)-L, P A; Schmid (1980, 1998)-A; Holzenthal et al.(2007, 2015)-L, P A; Ames(2009)-L, P A.

Regional faunas Alberta: Nimmo (1971, 1974, 1977a,b)-A. California: Denning (1956)-L, A, Colorado: Ward and Kondratieff (1992)-L. Florida: Pescador et al.(2004)-L. Illinois: Ross (1944)-L, P, A. Interior Highlands(Ozark, Ouachita, Arbuckle, and Wichita Mountains): Moulton and Stewart(1996)-L, A. Manitoba: Ruiter et al.(2013)-L. Minnesota: Houghton (2012)-A. New York: Betten (1934)-A, North and South Carolina: Unzicker et al.(1982)-L. Southeastern United States: Morse et al. (2017)-L. Wisconsin: Hilsenhoff(1981)-L.

Regional species lists Alabama: Harris et al.(1991); Harris and O'Neil (2015). Alaska: Nimmo (1986b); Kendrick and Huryn (2014). Arkansas; Unzicker et al.(1970); Bowles and Mathis(1989). British Columbia: Nimmo and Scudder (1978, 1983). California: Givens (2014). Colorado: Hermann et al.(1986). Delaware: Lake (1984). Florida: Blickle (1962); Harris et al.(1982b, 2012). Idaho: Smith (1965); Newell and Minshall (1977). Indiana: Waltz and McCafferty (1983); DeWalt et al.(2016). Iowa: Blinn et al.(2009). Kansas: Hamilton and Schuster (1978, 1979, 1980); Hamilton et al.(1983). Kentucky; Resh (1975); Floyd et al.(2012); Evans et al.(2017). Louisiana: Harris et al.(1982a); Holzenthal et al.(1982); Lago et a/.(1982). Maine; Blickle (1964); Blickle and Morse (1966). Manitoba: Flannagan and Flannagan (1982). Massachusetts: Neves(1979). Michigan: Leonard and Leonard (1949); Ellis(1962); Houghton et al.(2018). Minnesota: Etnier(1965); Lager etal.(1979); Houghton etal.(2001). Missouri: Mathis and Bowles(1992); Ferro and Sites (2007). Mississippi; Harris et al.(1982a); Holzenthal et al.(1982); Lago etfl/.(1982). Montana: Roemhild (1982). Nevada: Ruiter et al.(2014). New Hampshire: Morse and Blickle (1953, 1957). New York: Myers et al.(2011). Newfoundland: Marshall and Larson (1982). North Carolina: Denning (1950); Lenat et al.(2010). North Dakota; Harris et al.(1980). Ohio: Huryn and Foote (1983); Armitage et al.(2011). Oklahoma: Bowles and Mathis (1992a). Oregon: Anderson (1976b). Pennsylvania: Mastellar and Flint(1992). Quebec: Roy and Harper (1979). Rhode Island: Hunt(2017). South Carolina: Morse et al.(1980). Tennessee: Etnier and Schuster (1979); Etnier et al. (2000). Texas: Edwards (1973); Moulton and Stewart(1997b).

Utah: Baumann and Unzicker (1981). Virginia: Parker and Yoshell(1981); Flint et al.(2004, 2008, 2009); Flint(2014, 2017). Washington: Ruiter et al.(2005). West Virginia: Tarter (1990); Tarter and Floyd (2016). Wisconsin: Longridge and Hilsenhoff (1973). Wyoming: Ruiter and Lavigne (1985). Yukon: Nimmo and Wickstrom (1984); Wiggins and Parker(1997).

Taxonomic treatments at familial and generic levels Apataniidae; Ruiter (2000)-A; Chuluunbat et al.(2010)-L, P, A. Beraeidae: Wiggins(1954)-L, P, A; Hamilton (1985)-L, P, A. Brachycentridae: Wiggins(1965)-L; Flint(1984)-L, A. Calamoceratidae: Bowles and Flint(1997)-L, P, A; Prather(2003)-A. Dipseudopsidae; Ross(1965b)-A; Schuster and Hamilton (1984)A; Armitage and Hamilton (1990)-A; Sturkie and Morse (1998)-L. Ecnomidae: Bowles (1995)-L, A. Glossosomatidae: Ross(1956)-A; Wymer and Morse(2000)-A; Ruiter (2004)-A; Etnier et al.(2010)-A; Robertson and Holzenthal (2013)-A; Genco and Morse (2017)-P. Goeridae; Parker (1998)-L, A. Helicopsychidae: Johanson (2002)-A. Hydrobiosidae: Ross and King (1952)-A. Hydropsychidae: Denning (1943)-A; Flint(1961, 1974)-L, A; Schmid (1968)-A; Smith (1968a)-L, A; Gordon (1974)-A; Ross and Unzicker(1977)-A; Schuster and Etnier (1978)L; Flint et al.(1979)-A; Givens and Smith (1980)-L, A; Nielson (1981 )-A; Schefter and Wiggins(1986)-L; Schefter et al.(1986)-A; Schmude and Hilsenhoff(1986)-L, A; Nimmo(1987)-A; Korecki and Ruiter (2009)-A; Geraci et al. (2010)-L, A; Harvey et al.(2012)-L; Givens (2015)-L, P, A; Givens and Ruiter (2015)-A. Hydroptilidae; Flint (1970)-L, A; Denning and Blickle (1972)-A; Blickle (1979)-A; Marshall (1979)-L. A; Kelley (1982)-A; Kelley and Morse(1982)-A; Kelley (1984)-A; Moulton et al.(1999)-A; Parys and Harris (2013)-L; Ito et al. (2014)-L, P A; Keth et al.(2015)-A; Thomson and Holzenthal (2015)-A. Lepidostomatidae; Ross(1946)-A; Flint and Wiggins (1961)-A; Wallace and Sherberger (1972)-A; Weaver (1988)-L, A. Leptoceridae: Yamamoto and Wiggins(1964)-L, A; Yamamoto and Ross(1966)-A; Morse (1975, 1981)-A; Resh (1976b)L,P; Haddock (1977)-L, A; Holzenthal(1982)-A; Manuel and Nimmo (1984)-L, A; Floyd (1995)-L; Glover (1996)-L; Glover and Floyd (2004)-L; Carnagey and Morse (2006)A; Manuel(2010)-A; Blahnik and Holzenthal(2014)-A. Limnephilidae; Ross and Merkley (1952)-A; Schmid (1955, includes references to several generic revisions)-A; Flint (1956, 1960)-L; Denning (1964, 1970, 1975)-A; Wiggins and Anderson (1968)-L, A; Wiggins(1973b,c)-L, A; Wiggins and Richardson (1982, 1987, 1989)-L, A; Wiggins and Winchester (1984)-A; Parker and Wiggins (1985)-L, A; Wiggins and Wisseman (1990)-A; Winchester etal.(1993)-L, P, A; Ruiter (1995, 2000)-A; Ruiter and Nishimoto (2007)-L, P, A; Nimmo (2012)-A; Givens (2018)-L,P, A. Molannidae: Sherberger and Wallace (1971)-L; Roy and Harper (1980)-A. Odontoceridae: Parker and Wiggins (1987)-L, A. Philopotamidae: Ross(1956)-A; Lago and Harris(1987)-A; Armitage(1991)-A; Cooper and Morse (1998)-A; MunozQuesada and Holzenthal(2008)-A. Phryganeidae: Wiggins(1956)-A,(1960a)-L,(1962, 1998)-L, P, A; Wiggins and Larson (1989)-L, A. Polycentropodidae: Flint (1964b)-L; Morse(1972)-A; Hudson et al.(1981)-L; Nimmo (1986a)-A; Armitage and Hamilton (1990)-A; Chamorro and Holzenthal (2010)-A; Chamorro and Holzenthal (2011)-L, P, A.

Chapter 19 Trichoptera

Psychomyiidae: Ross and Merkley (1950)-A; Flint(1964b)-L; Armitage and Hamilton (1990)-A. Rhyacophilidae: Ross(1956)-A; Flint (1962)-L; Smith (1968b)L; Schmid (1970, 1981)-A; Peck and Smith (1978)-L, A; Weaver and Sykora (1979)-L; Prather et al.(1997)-A; Prather and Morse (2001)-A. Rossianidae: Ruiter (2000)-A; Wiggins(2004)-L, P, A. Sericostomatidae: Ross(1948)-A; Ross and Scott (1974)-A; Ross and Wallace (1974)-L, A; Keth and Harris(2008)-L, A. Thremmatidae: Vineyard and Wiggins {1987)-A; Vineyard and Wiggins(1988)-L, A; Ruiter (2000)-A; Vineyard et al. (2005)-L, A.

725

Uenoidae: Denning (1958, I975)-A; Wiggins et al.(1985)-L, A, P; Wiggins and Erman (1987)-A; Vineyard and Wiggins (1988)-L, A; Wiggins and Wisseman (1992)-A; Ruiter (2000)-A. Xiphocentronidae: Ross(1949)-A; Edwards(1961)-L, P; Schmid (1982)-A; Armitage and Hamilton (1990)-A; Moulton and Stewart(1997a)-A.

- Arctopsychinae

predators (engulfers) and seasonal scrapers

erosionai

(cool streams)

erosionai

Lotic—

ientic—

facultative

erosionai; some

Generally

retreats)

Ciingers (net spinners, fixed

filterers (coarse particles, animal and plant)

Collectors—

filterers; some

collectors—

Caddisflies

retreat makers)

North

Widespread

Trophic American Relationships** Distribution

spinning

Ciingers (netspinning fixed-

Habit

Generally

Habitat

lotic—

Species

Hydropsychldae

Arctopsyche(5)

Genus

(159)-Net-

Family

*SE = Southeast, DM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Hydropsychoidea

Annulipalpia

Caddisflies

Trichoptera -

Order

of species in parentheses)

Taxa(number

0.0

SE

UM

M

2.0

NW

q

MA*

Tolerance Values

q

6191

114, 434, 2465, 3985, 5536, 6250, 6256, 6270, 3484, 6192, 6328,

979, 1212, 1403, 1576, 2576, 2583, 3179, 4245, 4402, 4599, 5136, 5143, 853, 6270, 6524, 6539, 2087, 6448, 6789, 514, 5542, 6594, 5339, 6527, 1380, 6296, 6371

References**

Ecological

Table 19A Summary of ecological and distributional data for Trichoptera (caddisflies).(For definition of terms see Tables 6A-6C; table prepared by K. W. Cummins, J. R. Wallace, R. W. Merritt, G. B. Wiggins, J. C. Morse, R. W. Holzenthal, D. R. Robertson, A. K. Rasmussen, D. C. Currie, and M. B. Berg.)

) ) ) ) ) ) ; )) ) ) ) ) ) ) )))) ) ) ))) ) ) )

K> ON

- Hydropsychinae

• DIplectroninae

Family

Continued

retreats)

(rock face

retreats)

(especially

rivers)

streams and

warmer

Clingers (net spinners, fixed

Lotic— erosional

unknown

Larvae

springs and seeps)

Lotic—

Clingers (net spinners, fixed

retreats)

Clingers (net spinners, fixed

retreats)

Clingers (net spinners, fixed

Habit

erosional

(headwater streams)

erosional

Lotic—

erosional

Lotic—

Habitat

Cheumatopsyche

howellae

Species

(44)

Oropsyche

Homoplectra(12)

Diplectmna (5)

Parapsyche(7)

Genus

North

detritus, some invertebrates)

filterers (particles include algae,

Collectors—

filterers

Collectors—

(coarse particles, especially detritus)

Collectors—^filterers

(coarse particles)

Collectors—^filterers

Widespread

NC

East, West

Widespread

Widespread

Trophic American Relationships** Distribution

6,6

2.2

0.0

SE

5.0

UM

2.9

M

6.0

0.0

1.0

NW

5.0

MA*

Tolerance Values

Ecological

114, 160, 1073, 1783, 1975, 4071,4474, 4475, 4984, 4500, 4654, 6251, 5544, t

160, 6524, 6539

1296, 2583, 3686, 4077, 4475, 6256, 6270, 6719, t

6719

114, 434, 3640, 3686, 5404, 5536, 6256, 6270, 1528,

References**

Laboratory, and Oregon State and Humboldt State Universities

TRICHOPTERA

(continued)

tUnpublished data, K. W. Cummins, Michigan State University Kellogg Biological Station, University of Pittsburgh Pymatuning Laboratory of Ecology, University of Maryland, Appalachian Environmental

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

of species in parentheses)

Taxa(number

Table 19A

Continued

Potamyia

Hydropsyche(16)

Genus

flava

Species

retreats)

Clingers (net spinners, fixed

North

(detritus, diatoms)

Collectors—^filterers

facultative scrapers and predators

invertebrates):

Central, East

Collectors—^filterers Widespread (particles include diatoms, green algae, detritus,

Trophic American Relationships** Distribution ,.1

17.0

2.5

2.6

4.0

4.0

Tolerance Values

Ecological

1975, 5404

114, 160, 249, 434, 1073, 1077, 1212, 1739, 2015, 2016, 2017, 2150, 2223, 2581, 5475, 2583, 3023, 3271, 3292, 4647, 2020, 3645, 3933, 1047, 3985, 2018, 4071, 4240, 4307, 4402, 3700, 4474, 4475, 4611, 4689, 4522, 4786, 4984, 5372, 5404, 5448, 5456, 5492, 6251, 6775, 1347, 1674, 6256, 6270, 6422, 1535, 3484, 4623, 5544, 5339, 3988, 750, 6191, 6300, 6492, 3010, t

References**

Laboratory, and Oregon State and Humboldt State Universities

tUnpublished data, K. W. Cummins, Michigan State University Kellogg Biological Station, University of Pittsburgh Pymatuning Laboratory of Ecology, University of Maryland, Appalachian Environmental

**Emphasis on trophic relationships

Habit

Lotic— Clingers (net erosional spinners, fixed (larger rivers) retreats)

erosional

Lotic—

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic

Order

of species in parentheses)

Taxa(number

Table 19A

)) ) ) ) ) / )) ) ) ) ) ) ) ) ))) ) )) ) )) )

QO

-4

ve

Lotic—

Generally obligate collectors—filterers

Clingers (sac-like

erosional

Caddisflies

retreats)

Collectors—filterers

particles)

filterers (fine

collectors—

Obligate

Collectors—^filterers

silk net makers)

erosional

Clingers (net spinners, fixed

retreats)

(larger rivers)

Lotic—

Clingers (net spinners, fixed

retreats)

Clingers (net spinners, fixed

Habit

erosional

erosional

Lotic—

Habitat

Generally

albovirens

Species

lotic—

Smicridea (4)

Macrostemum (3)

Leptonema

Genus

North

Southwest

Central, East

Texas

American Trophic Relationships** Distribution

Philopotamidae (48)- Einger-net

- Smicrideinae

- Macronematinae

Family

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Philopotamoidea

Order

of species in parentheses)

Taxa(number

Table 19A

3.6

SE

1.8

M

3.0

NW

3.0

3.0

MA*

Ecological

(continued)

249, 979, 2829, 3179, 6255, 6256, 6270, 6524, 4994, 6370, 6539, 6527, 2958, 1380, 6371

6524, 6539

6492

6256, 6259, 6260, 6270, 4500, 5542, 5544, 6300,

1875, 5285, 6250, 6251,

1861, 1865, 6524, 2890, 446

References**

TRICHOPTERA

3.0

UM

Tolerance Values

J 3 3 J 3 )3 3 33 3 ) 33 3 ) ) ) 3 ) ) 3 ) ) l )

Caddisflies

Dipseudopsidae (5) Pitot-tube

Family

Continued

Phylocentropus(5)

Wormaldia (18)

Sisko (2)

Fumonta

Dolophilodes(8)

Chimatra (20)

Genus

major

Species

streams)

headwater

Borrowers

(branched, silk. buried tubes)

Lotic—

depositional (sand.

Collectors—filterers

filterers

Obligate collectors—

Clingers (sac-like silk nets)

Lotic— erosional

East

Widespread

Southeast, West

Southeast

unknown

filterers

Widespread

Widespread

Larvae

erosional

Lotic—

faces)

on rock

mossy seeps

streams and

(headwater

Obligate collectors—

Clingers (sac-like silk nets)

Lotic—

filterers

erosional

(warmer rivers)

Obligate collectors—

Clingers (sac-like silk nets)

Lotic—

Habit

erosional

Habitat

North

American Trophic Relationships** Distribution

5.6

0.4

1.0

2.8

SE

4.0

4.0

UM 2.6

M

3.0

1.0

NW

5.0

4.0

MA*

Tolerance Values

Ecological

5542

3640, 3648, 6276, 6524,

1380

5475, 6256, 6300, 6492

3700, 5283,

160, 434, 3686, 3985, 5404, 6255, 6270, 6594, 6600, t

249, 1073, 1077, 4402, 5404, 6256, 6270, 6524, 2299, 4522, 6608, 446, 6300, 6492, 3010, t

References**

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships tUnpublished data, K. W. Cummins, Michigan State University Kellogg Biological Station, University of Pittsburgh Pymatuning Laboratory of Ecology, University of Maryland, Appalachian Environmental Laboratory, and Oregon State and Humboldt State Universities

Psychomyloidea

Order

of species in parentheses)

Taxa(number

Table 19A

) ) ) ) ) ) ) )) ) D ) ) ) ) ) ) ) ) ) ) ) ) ) )

(Ji

1

Neureclipsis(5)

Holocentropus(8)

Cyrnellus

Cernotina (7)

fraternus

Collectors—

Lotic and

Lotic—

Collectors—

column)

water

herbivores.

engulfers (predators)

shredders—

facultative

filterers; some

or other

Clingers (trumpet-shaped silk nets)

Collectors—^filterers

supports in

hydrophytes

vascular

erosional (on

Clingers (silk tube retreats)

Predators

(engulfers)

Clingers (silk

predators (engulfers)

facultative

filterers: some

collectors—

Generally

filterers?

tube retreats)

lentic

North

Southwest

except

Widespread,

Central, North

Central, East

Central, East

Texas

Trophic American Relationships** Distribution

Lotic and

Clingers (netspinning retreat makers)

makers)

tube retreat

Clingers (probably silk

Habit

lentic

erosional

Generally

erosional

Lotic—

Habitat

Caddisflies

texensis

Species

lotic—

Austrotinodes

Genus

Polycentropodidae (76)- Trumpet-net

Caddisflies

Ecnomld

Ecnomldae (1)-

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

of species in parentheses)

Taxa(number

Table 19A

4.4

SE

2.7

M

NW

7.0

MA*

4998

(continued)

3944,4402, 4786, 5404, 4999, 5002, 6524, 4654,

6539

2755

4690, 5448, 6524, 6539, 1528, 4212, 4423, 6527, 3261, 1380

3023, 3179, 4402, 4611,

6484

1867, 6304,

6527, 1380

Ecological References**

TRICHOPTERA

7.0

UM

Tolerance Values

diversa

nearctica

Paduniella

erosional

Lotic—

Scrapers

Central, East

Ozarks

Missouri

Arkansas,

6539

3648, 6524,

1380

5143, 6524, 6539, 6527,

gatherers; some facultative scrapers makers)

erosional

Caddisfiies

Lype

979, 1403, 3179, 4599, 5136,

Generally collectors—

Ciingers (silk tube retreat

Generally

Ciingers (silk tube retreats)

Lotic— erosional

6648, 3383

herbivores

shredders—

2.8

160, 1739, 3944, 4402, 5404,

5873, 5874,

5.0

678, 4550

1073, 1077, 1862, 3944, 5542, 6524

6524, 6444,

6.0

Ecological References**

collectors—

2.0

3.4

MA*

fiiterers;

4.3

6.0

NW

iittorai

Southwest

3.5

SE

lentic—

Predators

(enguifers);

Ciingers (silk tube retreats)

Lotic—

herbivores

shredders—

Widespread

Southwest

collectors—

fiiterers;

Widespread, except

erosional;

lentic

Predators

littoral

Southwest

(enguifers);

herbivores

lentic—

Ciingers (silk tube retreats)

shredders—

depositional;

LotIc and

collectors—

fiiterers;

and

Widespread, except

Predators

(enguifers);

Ciingers (silk tube retreats)

Trophic American Relationships** Distribution

Lotic—

Habit

erosional

Habitat

lotic—

Species

Tolerance values

Psychomylidae

Polyplectropus(3)

Polycentropus(29)

Plectrocnemia (13)

Nyctiophylax(10)

Genus

North

(17)- Net-tube

Family

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = ^ Mid-Atlantic **Emphasis on trophic relationships

Order

of species in parentheses)

Taxa(number

Table 19A

) ) ) ) } ) ))) ) ) ) ) ) ))) ) ) ) ) ) ) ) ) )

-4 OJ

-4

Lotic—

Caddisflies

Lotic—

makers)

turtle-shell case

Generally obligate scrapers

Clingers (saddle- or

Collectors—

gatherers

Lotic— erosional

springfed ponds

erosional; lentic—

gatherers

0.0

1604, 5802,

1380

1403, 2576, 3179, 4599, 5136, 5143, 6524, 6539, 3099, 6527,

6539

TRICHOPTERA

(continued)

tUnpublished data, K. W. Cummins, Michigan State University Kellogg Biological Station, University of Pittsburgh Pymatuning Laboratory of Ecology, University of Maryland, Appalachian Environmental Laboratory, and Oregon State and Humboldt State Universities

Texas

Arizona

6527, 1380

325, 1320, 4611, 6524, 391, 1535, 2853, 2854

160, 1073, 1077, 4611, 5404, 6524, 6539,

Scrapers;

2.0

2.0

Ecological References**

collectors—

1.9

NW MA*

Lotic—

2,0

M

erosional

2.6

UM

2431, t West

Widespread

SE

Tolerance Values

scrapers

gatherers and

Facultative collectors—

Clingers (silk tube retreats)

erosional

messapus

XIphocentron

North

Trophic American Relationships** Distribution

Lotic—

Habit

erosional

Habitat

Glossosomatidae

yavapai

Species

Cnodocentron

Tinodes(12)

Psychomyia (3)

Genus

(89)- Saddle-case

Caddisflies

Xiphocentronidae (2) Xiphocentronid

Family

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Basal Lineages

Integripalpia

Order

of species in parentheses)

Taxa (number

Table 19A

■ Glossosomatinae

• Agapetinae

Family

Continued

Glossosoma (23)

Anagapetus(5)

Agapetus(42)

Genus

Species

(including large alpine rivers)

Clingers (turtle shell case, mineral)

Lotic—

erosionai

(including small alpine springs)

Clingers (turtle shell case, mineral)

Lotic—

turtle shell case, mineral)

Clingers (laterally compressed

Habit

erosionai

erosionai

Lotic—

Habitat

American

0.0

164, 201, 3985,

Obligate scrapers

Scrapers

Widespread

West

1.5

0.0

0.0

4357, 4599, 4611, 5136, 5143, 5372, 5845, 5998, 6524, 3099, 4421,4734, 3226, 3227, 6563, 201, 3225, 4420, 251, 3861, 4422, 202, 3323, 4732, 4983, t

2223, 3023, 3179, 3279, 3562, 4077,

160, 164, 249, 979, 1249, 1403,

160, 164, 6539

3010

6300, 6492,

3697, 115,4731,

facultative

gatherers

0.0

Ecological References*

collectors-

Widespread

Distribution

Tolerance values

5448, 202, 1844, 4734, 6328,

Scrapers;

Trophic Relationships

North

*SE = Southeast, DM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships tUnpublished data, K. W. Cummins, Michigan State University Kellogg Biological Station, University of Pittsburgh Pymatuning Laboratory of Ecology, University of Maryland, Appalachian Environmental Laboratory, and Oregon State and Humboldt State Universities

Order

of species in parentheses)

Taxa(number

Table 19A

) ) ) ) > ) ) ) ) ))))) ) )) ) ) ) ))) )) )

u>

Order

Lotic—

gatherers

down; purse- or barrel-case

lentic— littoral

case builders

living, 5th instar

instars free

makers). First 4

collectors—

fasten case

lentic—

scrapers;

herbivores;

erosional;

Generally piercers;

(engulfers)

Predators

Facultative scrapers

Generally clingers or climbers(may

Clingers (free ranging)

turtle shell case. mineral)

Clingers (laterally compressed

Generally

erosional

Lotic—

erosional

turtle shell case)

Scrapers

Southwest

Widespread

Southeast

erosional

Lotic—

Clingers (depressed

Northwest, West

Scrapers

Clingers (turtle shell case. mineral)

Lotic—

Habit

erosional

Habitat

lotic and

jeanae

Species

Hydroptilidae

Atopsyche(3)

Protoptlla (13)

Padunia

Culoptila (5)

Genus

North American Trophic Relationships** Distribution

(309)Microcaddlsflies

Caddisflies

Hydrobiosidae (3) Hydrobiosid

- Protoptilinae

Family

Continued

2.8

0.0

SE

1.0

UM

M

4.0

1.0

0.0

NW

MA*

Tolerance Values

1380

1403, 2576, 3179, 4347, 4599, 5136, 5143, 6524, 6539, 6527,

6524, 6539

6527, 1380

5404, 6524, t

6524, 6539

2721

6524, 6539,

Ecological References** 1

TRICHOPTERA

(continued)

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships tUnpublished data, K. W. Cummins, Michigan State University Kellogg Biological Station, University of Pittsburgh Pymatuning Laboratory of Ecology, University of Maryland, Appalachian Environmental Laboratory, and Oregon State and Humboldt State Universities

1

of species In parentheses)

Taxa(number

Table 19A

J3 ))>^33J3)3J :) )

Family

Continued

Mayatrichia (5)

Leucotrichia(3)

Ithytrichia (3)

Hydroptila (120)

Dibusa

Alisotrichia

Agraylea (4)

angata

Lotic—

erosionai

Lotic—

erosionai

Clingers (pursetype case, fixed)

silk)

Clingers (pursetype case of

mineral)

depositional (including seeps) Lotic—

and fine

and

and Lemanea)

Clingers (pursetype case of silk Clingers (pursetype case of silk

erosionai (on rocks)

American Trophic Relationships** Distribution

Scrapters (red alga.

6.0

160, 1866, 5404,

6524

6524, 3089, 574

160, 1077, 4347, 4611, 1535,

Scrapers

gatherers Widespread

3860, 3862, 2428, 2431, 3099, t

2.0

6.0

4551

4959, 6524,

544, 545

160, 306, 4071, 4307, 4347, 5159, 5404, 5448, t

collectors—

4.3

2.6

8.0

Ecological References**

6524, 2436,

Widespread

Widespread

Widespread

East

5.7

Tolerance Values

scrapers;

Facultative

Scrapers

(Cladophora); facultative scrapers

herbivores

Piercers—

Lemanea)

Southwest

Widespread, Climbers Piercers— except deep (purse-type case herbivores of silk and algal (filamentous algae); Southeast and plant stem collectors— strands) gatherers

Habit

erosionai

Lotic—

erosionai

Lotic—

erosionai

Lotic—

hydrophytes)

(vascular

erosionai

lotic—

algae);

filamentous

(with

hydrophytes

vascular—

Lentic—

Habitat

North

) ) ) ) ) ) ) )) )

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships tUnpublished data, K. W. Cummins, Michigan State University Kellogg Biological Station, University of Pittsburgh Pymatuning Laboratory of Ecology, University of Maryland, Appalachian Environmental Laboratory, and Oregon State and Humboldt State Universities

Order

parentheses)

Taxa(number of species in

Table 19A

)o ) ) I ) ) ) ) ) ) )) ) )

OS

Family

Continued

Oxyethira (46)

Orthotrichia (6)

Ochrothchia (57)

Nothotrichia

Neotrichia (40)

Metrichia (3)

Genus

shasta

Species

of silk)

Piercers—

herbivores;

Climbers

(purse-type

case, flat, flask- collectors— shaped, open gatherers; scrapers at back. (?) primarily of silk)

hydrophytes

herbivores

Piercers—

gatherers; scrapers

collectors—

facultative

herbivores;

Piercers—

Scrapers

Lentic—

hydrophytes)

depositlonal (vascular

and

erosional

lotic—

algae);

filamentous

(with

North

SE

UWI

5.2

3.6

M

6.0

6.0

4.0

NW MA*

Tolerance Values

4347, 6524, 3091, t

6524

160, 1077, 4347, 4611, 1535, 6524, 3089

4535

6524, 6539

Ecological References**

)):>

Laboratory, and Oregon State and Humboldt State Universities

TRICHOPTERA

(continued)

tUnpublished data, K. W. Cummins, Michigan State University Kellogg Biological Station, University of Pittsburgh Pymatuning Laboratory of Ecology, University of Maryland, Appalachian Environmental

Widespread

Widespread

Widespread

California

Widespread

Southwest

Central,

Trophic American Relationships** Distribution

vascular

algae)

filamentous

(with

vascular

hydrophytes

mineral)

depositlonal Clingers? (purse-type case

and fine

and

Lentic—

Clingers (pursetype case of silk

Lotic— erosional

mineral)

Clingers (case a tube of fine

Lotic—

Habit

erosional

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

of species in parentheses)

Taxa(number

Table 19A

) ) J3 )

U)

Caddisflies

Rhyacophllidae (130)- Free-living

Family

Continued

Himalopsyche

Zumatrichia

Stactobiella (5)

Paudcalcaria

Palaeagapetus(2)

Genus

phryganea

notosa

ozarkensis

Species

erosional

Lotic—

erosional

Lotic—

erosional

Lotic—

streams)

depositional (small rapid

and

erosional

Lotic—

(cold springs and seeps)

erosional

Lotic—

Habitat

Habit

Shredders

detritivores

Shredders—

Trophic Relationships**

Clingers (free ranging) (alpine)

Clingers (free ranging)

scrapers

(engulfers),

Predators

Generally predators (engulfers)

Clingers (purse- Scrapers; type case, fixed) collectors— gatherers

strands)

Clingers? (purse-type case of silk and algal and plant stem

liverwort)

fragments, especially

of small leaf

Sprawlers (purse-type case

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

of species In parentheses)

Taxa (number

Table 19A

North

Pacific states

West

Widespread

Arkansas

Northwest

Northeast,

Distribution

American SE

UM

M

0.0

2.0

NW

MA*

Tolerance Values

Ecological

6524

1380

1403, 3179, 4599, 5136, 5143, 1528, 6524, 6539, 6753, 6527,

6524

160, 6524, 6539

2418

160, 6539, 2863

References**

) ) ) I ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) )) )

00

-4

Caddisflies

Spring-loving

Beraeidae (3) -

Family

Continued

Beraea (3)

Rhyacophila (129)

Genus

Species

fine mineral)

(detritus,

gatherers

curved, smooth, collectors-

springs)

Sprawlers (case

Probably

(chewers)

herbivores

shredders—

gatherers.

collectors—

scrapers.

Predators (engulfers); a few

depositional

Ciingers (free ranging)

Habit

Trophic Relationships**

Lotic—

erosional

Lotic—

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Brevitentoria

Infraorder

Order

of species in parentheses)

Taxa (number

Table 19A

North

East

Widespread

Distribution

American

o

o

o

1,0

Ecological

(continued)

2348, 6510, 6524, 1380

6527

801, 3562, 3640, 3648, 3716, 3985, 4357, 4611, 5372, 5404, 5448, 5492, 1347, 5990, 6524, 6753, 1535, 1534, 6328, 1528, 1534, 3740, 3366, 3484, 3700, 4415, 4212, 4804, 961, 3383, 3697, 3699, 6143, 4442, 1677, 3010

References**

TRICHOPTERA

o

Tolerance values

)

■tx

o

)

Sprawlers

Habit

Generally

Trophic Relationships**

Helicopsychidae

Caddisflies

(7) - Snail-case

Phylloicus (2)

Heteroplectron (2)

large leaf pieces

(detritus)

Lotic

detritivores; facultative scrapers

projection)

and dorsal

with large leaf and bark pieces

Shredders—

Sprawlers? (case a flat tube

facultative scrapers

bark)

and

(detritus)

litter and gougers

stick or piece of

(detritus) of wood);

(chewers of leaf

a hollowed out

depositional

detritivores

Sprawlers (case

Lotic—

erosional

projection) Shredders—

detritivores

(case flat of

depositional with dorsal

shredders—

Sprawlers?

Lotic—

Generally

scrapers

detritivores and

Lotic

Habitat

Caddisflies

pyraloides

Species shredders—

Anisocentropus

Genus

(5) - Comb-lipped

Calamoceratidae

Family

Continued

Southwest

Cost

East, West

Southeast

Distribution

American

North

2.9

3.0

1.0

3.0

Tolerance Values

Ecological 3179, 5136, 6524, 1380

1380

1403, 4599, 3373, 5542, 3370,

3179, 3680, 5143, 6524, 4964,

3118, 6524, 2890, 2211, 4392, 573, 574

160, 167, 3562, 4540, 168, 6524, 6648, 5845, 6527, 4442, t

6257, 6524

1403, 4599, 5143, 6527,

References**

)

)

)

)

)

)

)

)

)

)

)

)

)

)

)

)

)

)

)

)

)

)

)

)

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships tUnpublished data, K. W. Cummins, Michigan State University Kellogg Biological Station, University of Pittsburgh Pymatuning Laboratory of Ecology, University of Maryland, Appalachian Environmental Laboratory, and Oregon State and Humboldt State Universities

Order

of species in parentheses)

Taxa (number

Table 19A

)

Genus

mineral)

projection and flanges)

a fine mineral or dorsal

Widespread

6.4

0.0

3.0

3.0

3.1

1.8

5.0

4.0

3.0

3.0

3944,4934, 4936, 4937, 4938, 4940, 4965, 5136, 2549, 5404, 6524, 5489

1380

1403,3179, 4599, 5136, 1951, 5143, 5830, 6524, 6527, 1204,

2800, 4963, 6581, 1777, 6153, 6154, 4964, 6155, 6157, 2890, 3010, t

1776, 1884, 5404, 6524,

1073, 1077, 3984, 3985,

Laboratory, and Oregon State and Humboldt State Universities

TRICHOPTERA

(continued)

tUnpublished data, K. W. Cummins, Michigan State University Kellogg Biological Station, University of Pittsburgh Pymatuning Laboratory of Ecology, University of Maryland, Appalachian Environmental

**Emphasis on trophic relationships

predators (engulfers of sponge)

(chewers);

herbivores

shredders—

Collectors— gatherers;

limnetic)

in sponges)

herbivores

(chewers); scrapers; predators (engulfers)

wide variety)

(including

Climbers—

shredders—

case makers of

habitats

sprawlers,(case

filterers;

swimmers(tube

lentic

horned Caddisflies

Lotic and

gatherers and

lentic(some

Collectors—

Climbers—

sprawlers— clingers—

Cerac/ea (39)

MA*

Widespread

NW

Obligate scrapers

lotic and

springs)

thermal

shaped, fine

erosional

(including

Clingers (case snail shell

lentic—

Habitat Lotic and

All types of

Species

M

Ecological UM

References**

SE

Distribution

Leptoceridae

Helicopsyche(7)

Habit

American

North

Tolerance Values

Trophic Relationships**

(123)- Long-

Family

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic

Order

of species In parentheses)

Taxa(number

Table 19A

Family

Continued

Oecetis(26)

Nectopsyche {]5)

Mystaddes(3)

Leptocerus

long and

hydrophytes

mineral and

vegetation pieces may

vascular

hydrophytes

herbivores

long, slender of (chewers); collectors—

gatherers; (predators [engulfers])

mineral and

vegetation pieces, may have long balance sticks)

hydrophytes: lotic— erosional

herbivores of coarse

littoral

mineral or plant fragments)

shredders—

often tapered.

lentic—

Widespread

Widespread

Widespread

Central, East

5.7

4.2

3.5

SE

3.0

4.0

UM

2.4

M

8.0

3.0

4.0

NW

8.0

3.0

4.0

MA*

Tolerance Values

Ecological

481, 3944, 5136, 5404, 6328

449, 2576, 3944, 3984, 3985, 4307, 5136, 5404, 4337, 6328, 6524

6776, t

160, 2588, 3562, 3944, 6524,

6524

3648, 4396,

References**

Laboratory, and Oregon State and Humboldt State Universities

00 b

tUnpublished data, K. W. Cummins, Michigan State University Kellogg Biological Station, University of Pittsburgh Pymatuning Laboratory of Ecology, University of Maryland, Appalachian Environmental

facultative

a curved tube.

depositionai;

and

Predators

(engulfers);

Climbers—

clingers— sprawlers,(case

Lotic—

erosional

hydrophytes)

(vascular

depositionai

and

Shredders—

Climbers—

swimmers (case

Lentic—

(chewers)

herbivores

shredders—

facultative

gatherers;

Collectors—

(chewers)

herbivores

Shredders—

Trophic American Relationships** Distribution

vascular

sticks)

have balance

sprawlers (case a rough tube of

depositionai; lentic—

Climbers—

Lotic—

primarily of silk)

slender.

Climbers—

swimmers (case

Lentic—

Habit

vascular

Habitat

*SE = Southeast, DM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

parentheses)

of species in

Taxa(number

Table 19A

UJ

) ) ) ) ) ) > ) ) ) )) ) ) ) ) 1 ) ) ) ))))))

-4

) )

OS

:)

leaf and bark

fragments, or cylindrical, of

streams and

springs)

Climbers—

scavengers on

shredders—

detritlvores; instars

case makers of

great variety)

lentic habitats

Caddlsflles

East

Widespread

Distribution

American

North

1.0

1.0

4.0

1.0

4.0

1.0

Tolerance values

Ecological

6524, 6539, 6527, 1211, 1204, 1380

5143, 5542,

979, 1403, 1858, 2576, 3179, 4599, 5136,

6524

160, 163, 167, 979, 2214, 2361, 3640, 5394, 5404, 5438, 6607, 6648, 400, 5001, 6450, 446, 4047, 3220, 3235, 2270, t

References**

Laboratory, and Oregon State and Humboldt State Universities

tUnpublished data, K. W. Cummins, Michigan State University Kellogg Biological Station, University of Pittsburgh Pymatuning Laboratory of Ecology, University of Maryland, Appalachian Environmental

herbivores

shredders—

gatherers; some

collectors—

some facultative

facultative scrapers;

mineral cases

shredders, with

facultative

with organic cases

with organic cases, generally obligate

sprawlers— clingers (tube

All types of

When all Instars

detrltivores

Shredders—

salmon carcasses)

lotic and

gravel)

Climbers—

sprawlers (case cylindrical of sand)

Lotic—

erosional (in

sand or silk)

"rough log cabin" type of

(detritus) (headwater

(chewers)(also reported as

detrltivores

square or

depositional

and

Obligate shredders—

Climbers—

sprawlers— clingers (case

Trophic Relationships**

Lotic—

Habit

erosional

Habitat

LImnephilidae

Theliopsyche(6)

• Theliopsychinae

Species

(252)- Northern

Lepidostoma (66)

Genus

■ Lepidostomatinae

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

of species in parentheses)

Taxa(number

Table 19A

) ) ) ) ) ) ) ))) ) ) ) ) ) ))) ) ) ) ) ) ) ) )

©

(71

-J

Family

Continued

canax

Amphicosmoecus

Arctopora (3)

Anabolia (5)

partitus

Species

Allocosmoecus

Genus

gatherers

Climbers—

sprawlers (case a smooth tube

of long leaf pieces)

Lentic— littoral

(including temporary ponds)

temporary ponds)

collectors—

pieces; may be three-sided)

depositionai (including

facultative

a rough tube of (chewers); leaf and wood

hydrophytes:

detritivores

lotic—

Shredders—

Climbers—

sprawlers (case

vascular

wood pieces)

littoral

Shredders

Lentic—

bark pieces or entire case of

and

ientic—

with anterior

erosionai

depositionai;

Sprawlers (case a hollowed twig

rough mineral)

Scrapers; shredders?

Sprawlers (case curved, flattened.

Lotic—

North

North

Widespread

West

Northwest

American Trophic Relationships** Distribution

Lotic—

Habit

erosionai

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

of species in parentheses)

Taxa (number

Table 19A

Ecological

6524

2906

(continued)

2359, 3023, 5492, 6524, 476,

6539

4365, 6524,

31,6524

References**

TRICHOPTERA

0.0

Tolerance Values

Family

Continued

centralis

Chyranda

Clostoeca

disjuncta

areolatus

Chilostigmodes

Clistoronia (4)

itascae

Species

Chilostigma

Asynarchus(13)

Genus

temporary

bark and leaf

(detritus)

detritivores

Shredders—

a flattened tube

depositional (detritus. spring seepage)

of large leaf pieces with flanges)

Sprawlers (case

Lotic—

Lentic—

Northwest

SE

UM

M

1.0

NW

MA*

Tolerance Values

Ecological

6524, 6539

159, 160, 4880, 6524, 890, 6648, t

6607

160,6524,6539,

3299, 6524

6524

1841, 6522,

160, 1858, 4365, 6524, 513, 6607, 6693, 3598

References**

Laboratory, and Oregon State and Humboldt State Universities

tUnpublished data, K. W. Cummins, Michigan State University Kellogg Biological Station, University of Pittsburgh Pymatuning Laboratory of Ecology, University of Maryland, Appalachian Environmental

detritivores

Shredders—

(chewers)

West

North, West

North

Minnesota

North

American Trophic Relationships** Distribution

Sprawlers (case Collectors— littoral a rough tube of gatherers; (sediments twig and bark facultative and detritus) pieces arranged shredders— longitudinally) detritivores

pieces)

Sprawlers (case a flat tube of

Lotic—

leaves and bark)

depositional

unknown

Larva

seeps

Lotic-spring

depositional

lotic—

Sprawlers (case irregular tue of small pieces of

mineral and

plant pieces)

(including ponds);

Climbers (case variable tube of

Lentic—

Habit

littoral

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

of species in parentheses)

Taxa(number

Table 19A

) ) ) ) ) ) ))) ) ))) )) ))) ) ))) ) ) ) )

K)

■~4

Family

Continued

Desmona (2)

Cryptochia (7)

Crenophylax

Genus

sperryi

Species

(terrestrial

herbivory reported

of coarse sand

and/or organic debris)

(small spring streams)

California

West

Southwest

Distribution

American

North SE

UM

M

0.0

NW

MA*

Tolerance Values

Ecological

1757

3783, t

160, 6522, 73,

6547, 5197

References**

Laboratory, and Oregon State and Humboldt State Universities

TRICHOPTERA

(continued)

tUnpublished data, K. W. Cummins, Michigan State University Kellogg Biological Station, University of Pittsburgh Pymatuning Laboratory of Ecology, University of Maryland, Appalachian Environmental

**Emphasis on trophic relationships

in D. bethula)

herbivores

Sprawlers (case

Shredders—

or shredders

Facultative scrapers

Scrapers

Trophic Relationships**

Lotic—

wood and bark)

Sprawlers (case flat, tapered, of

particles

mineral

slightly tapered posteriorly, slightly curved; pupal case of

and leaves,

irregular bark

constructed of

Sprawlers (prepupal case rough,

Habit

erosional

springs and seeps)

streams,

(detritus; small, cool

depositional

Lotic—

brooks

Spring

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic

Order

of species in parentheses)

Taxa (number

Table 19A

)))))3 )) )))) )))) ))) ) ) )) ) ))

Family

Continued

Eocosmoecus(2)

Ecdisomyia (4)

Ecdisocosmoecus

Dicosmoecus(4)

Genus

scylla

Species

may have long plant pieces)

(cold alpine

streams)

(small spring

erosional

Lotic—

Northwest

West

Northwest

West

M

2.0

0.0

NW MA*

6464, 6535

6524

160, 3985, 4365, 5141, 5001,

6539

5001, 6524,

6533, 2466, 2926, 3372, 1400, 3373, 3370, 4047, 367, 1366, 4968, t

160, 763, 2434, 6524, 2202,

Ecological References**

Laboratory, and Oregon State and Humboldt State Universities

tUnpublished data, K. W. Cummins, Michigan State University Kellogg Biological Station, University of Pittsburgh Pymatuning Laboratory of Ecology, University of Maryland, Appalachian Environmental

Shredders

in cases

significant amount of organic matter

shredders if

facultative

with mineral cases;

gatherers; facultative scrapers

mineral tube,

streams)

Collectors—

Clingers (case a

Scrapers; shredders

detritivores,(also reported as predators and scavengers)

shredders—

cases are

Scrapers; early instars in organic

Lotic—

Borrowers (case tapered, curved, smooth mineral)

Sprawlers (case curved, flattened, rough mineral, early instars with organic case)

Habit

North

Tolerance Values

Trophic American Relationships** Distribution SE UM

erosional

(sand)

depositional

Lotic—

erosional

Lotic—

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

of species in parentheses)

Taxa(number

Table 19A

)) ) ) ) ))) ) ) )) ) )) )))))))) )))

(71

'M

Continued

Grensia

Grammotaulius(4)

Glyphopsyche(3)

Frenesia (2)

Genus

praeterita

Species

of twig and bark pieces)

mineral)

fragments and

Lentic— Sprawlers? (case a curved littoraK?) (tunda lakes) tube of plant

streams

temporary

hydrophytes), including

(vascular

depositional

lotic—

Climbers (case

a tube of long leaf pieces)

Lentic—

littoral and

depositional (detritus)

lotic—

(detritus);

depositlonal (detritus; including springs) a smooth tube

(chewers)

of minerl and

wood pieces)

and

Sprawlers (case

detrltivores

a smooth tube

littoral

Shredders—

Sprawlers (case

lentic—

North

Northwest

North

Central, North

East

Trophic American Relationships** Distribution

Lotic—

Habit

eroslonal

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

of species in parentheses)

Taxa(number

Table 19A

UM

IVI

4.0

1.0

NW MA*

6524

6524

(continued)

160, 1858, 2588,

6524

1756, 3562, 5143, 6524

Ecological References**

TRICHOPTERA

SE

Tolerance Values

Continued

Ironoquia (5)

Hydatophylax(4)

Homophylax(10)

Hesperophylax(6)

Halesochila

Genus taylori

Species

shredders— detritivores

pieces)

mineral tube)

depositional (detritus),

a curved tube of leaf or bark

pieces, or mineral)

depositional; lentic— littoral

(temporary

ponds)

streams and

Sprawlers (case

rough cylinder of wood, bark, mineral, with balance sticks)

(detritus)

Lotic—

detritivores; some facultative

climbers (case a

Central, East

Southwest

except

Widespread,

West

North, West

Northwest

7.3

2.3

3.0

2.0

1.0

0.0

5.0

Tolerance Values

Ecological

2386

302, 1857, 3640, 5136, 6519, 6524, 6599, 948,

160, 167, 1858, 3562, 6206, 6524, t

6524, 6524

160, 2223, 3562, 3756, 5404, 6524, 6607, 513, 3118, 4517

6648

2115, 6524,

References**

Laboratory, and Oregon State and Humboldt State Universities

tUnpublished data, K. W. Cummins, Michigan State University Kellogg Biological Station, University of Pittsburgh Pymatuning Laboratory of Ecology, University of Maryland, Appalachian Environmental

Shredders

gatherers

collectors—

shredders—

Sprawlers—

Obligate

(chewers)

Lotic—

of bark pieces)

a smooth tube

detritivores

Shredders—

gatherers)

collector—

(chewers); scrapers;

herbivores

detritivores and

Shredders—

depositional

(sediments and detritus)

erosiona!

Lotic—

streams

temporary

including

and

Clingers— sprawlers (case

Sprawlers (casea slightly curved, slightly rough, coarse

Lotic—

erosiona!

(chewers)

gatherers;

leaf and wood

Collectors—

(sediments)

littoral

Sprawlers (case a rough tube of

Habit

North

American Trophic Relationships** Distribution

Lentic—

Habitat

*SE = Southeast, DM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

of species in parentheses)

Taxa(number

Table 19A

) ) ) ) ) ) ) )) ) ) ) ) ) )) !) ))))) ) ))

o\

-4 (41

'J\

Family

Continued

mono hostilis

Monophylax

gradlis

Species

Nemotaulius

Limnephilus(85)

Leptophylax

Lenarchus(9)

Genus

tube of long leaf pieces or

(including

Larvae

gatherers (and probably others)

temporary

(detritus)

depositional

lotic—

(detritus):

littoral

Lentic—

ponds and streams)

5.0

NW

MA*

Ecological

1858,3562,

6548

5284, 5404, 5492, 6206, 6519, 6524, 6607, 476, 4423, 6684, 2133, 4337, t

3271,3562, 3944, 4045,

160,302, 1858, 2359, 2360,

6524

6648

6524, 6607,

References**

5492,6524

M

(chewers)

UM

5136, 5404, 476,

North

SE

Tolerance Values

detritivores

Shredders—

West

collectors—

or sand

construction. variable)

(including

Sprawlers (case a flat "log cabin" type, of leaf pieces)

Southeast

herbivores;

habitats

facultative

detritivores and

lentic

Widespread, except deep

Shredders—

Climbers—

sprawlers— clingers (case of stick, leaf and/

lotic and

Central

North, West

All types of

unknown

pieces)

bark and leaf

climbers (case a

temporary ponds)

Collectors—

gatherers

Sprawlers—

Trophic American Relationships** Distribution

Lentic—

Habit

littoral

Habitat

North

TRICHOPTERA

(continued)

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships tUnpublished data, K. W. Cummins, Michigan State University Kellogg Biological Station, University of Pittsburgh Pymatuning Laboratory of Ecology, University of Maryland, Appalachian Environmental Laboratory, and Oregon State and Humboldt State Universities

Order

parentheses)

Taxa(number of species in

Table 19A

)))))))

Family

Continued

bergrothi

Philarctus

Philocasca (7)

canadensis

Phanocelia

Onocosmoecus(2)

mineral portion)

terrestrial)

semi-

(detritus-

depositional

and

(sediments and detritus)

erosional

Lotic—

coarse mineral)

Ciingers— sprawlers (case rough, curved.

pieces)

small shells. seeds, and leaf

lotic—

depositional

Sprawlers (case a tube with

Lentic—

littoral;

Sprawlers (case

of short pieces of sphagnum arranged transversely)

Lentic—

sphagnum bog pools

(detritus)

littoral

or with some

lentic—

Sprawlers (case of wood, bark.

Habit

depositional (detritus);

Lotic—

Habitat

detritivores

shredders—

Probably

larvae, crustaceans)

engulfers (insect

herbivores. predators—

Shredders—

Northwest

Northwest

North

North,West

Obligate shredders

American Distribution

Trophic Relationships** SE

UM M

1.5

NW

MA*

Tolerance Values

Ecological

6537

156, 160, 6524,

6524

1742

4365,6524, 6539, 6607, 6534, 6648, 6688, 3506, t

References**

Laboratory, and Oregon State and Humboldt State Universities

tUnpublished data, K. W, Cummins, Michigan State University Kellogg Biological Station, University of Pittsburgh Pymatuning Laboratory of Ecology, University of Maryland, Appalachian Environmental

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

Taxa(number of species in parentheses)

Table 19A

) ) ) ) )) )))) ))))) ) ) ) )))) ) )))

U\ 00

ui

Family

Continued

Psychoronia (2)

Psychoglypha (25)

gatherers(some scavengers on

and wood

pieces)

depositional

Lotic—

(alpine streams?)

erosional

tube)

Sprawlers (case a rough mineral

detritivores and collectors—

mineral, bark

1.0

West

North, West

2.0

157, 160, 163,

6524, 1841

160, 763, 1858, 6524, 6648, 961, 35

1858, 3640, 6524, 6537

Laboratory, and Oregon State and Humboldt State Universities

TRICHOPTERA

(continued)

tUnpublished data, K. W. Cummins, Michigan State University Kellogg Biological Station, University of Pittsburgh Pymatuning Laboratory of Ecology, University of Maryland, Appalachian Environmental

salmon carcasses)

shredders—

and

erosional

Lotic—

(detritus)

Facultative

curved, smooth, (chewers); collectors— mineral) gatherers

Northwest

Sprawlers— clingers (case a type of mixed

temporary streams)

depositional (including seeps and

and

(detritus)

Lotic—

erosional

detritus)

1858, 6524, 6607, t

Widespread

Ecological References**

Central, East,

hydrophytes,

Tolerance Values

Distribution

detritivores

pieces)

(vascular

North American

Shredders—

other fine

depositional

Sprawlers (case tapered,

detritivores (and herbivores)

I otic—

Shredders—

Climbers (case

a rough log cabin type of plant stems and

Lentic—

Habit

Trophic Relationships*

littoral;

Habitat

Pseudostenophylax

Species

(3)

Platycentropus(3)

Genus

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

Taxa (number of species in parentheses)

Table 19A

depositional

lotic—

160, 6513, 6524, t

Laboratory, and Oregon State and Humboldt State Universities

tUnpublished data, K. W. Cummins, Michigan State University Kellogg Biological Station, University of Pittsburgh Pymatuning Laboratory of Ecology, University of Maryland, Appalachian Environmental

(chewers); scrapers(?)

herbivores

detritivores and

(ponds);

Shredders—

Climbers (case a

spirally arranged. tapered cylinder of leaf pieces)

Lentic— littoral

Widespread

Terr.

1247, 1770, 2219, 2731, 3562, 3642, 3643, 3649, 4071, 6524, 6601, 512, 513, 5542, 3742, 513, 5542, 3742, 6062, 6281, 1270, 4151, 5477, 5777, 4162, 1609, 2810, 5111, 1366, t

1073, 1077,

tundra pools

4.0

Ecological References*

6538, 6641,

3.3

NW MA*

6642

4.0

M

Northwest

Shredders

2.5

UM

Terr., Yukon

Sprawlers

North

Central, East,

SB

Tolerance Values

transient

Lentic—

(detritus)

Generally

Agrypnia (10)

or like

Hydatophylax)

smooth mineral,

depositional; lentic— littoral

clingers (case

and

facultative scrapers

detritivores;

climbers or

Obligate shredders—

Sprawlers—

Lotic—

Habit

erosional

Habitat

shredders; some predators

meiops

Species

Phryganeidae (28)

Sphagnophylax

Pycnopsyche(18)

Genus

North

Trophic American Relationships** Distribution

- Giant Caddisflies

Family

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

of species in parentheses)

Taxa (number

Table 19A

) ) ) )) ) ) ) )) ) ) > ) ) ))) ) )) 1 ) ) ) 1

0\

OS

Continued

inornata

canadensis

Fabria

Hagenella

Oligostomis(2)

complicatus

Species

Beothukus

Banksiola (5)

Genus

"Christmas tree

hydrophytes

herbivores and detritivores

(chewers)

(detritus and vascular

shredders—

Hagenella)

and

depositional

Predators

(engulfers);

Climbers (case similar to

Southeast

East except deep

Northcentral, Northeast

North

Northeast

Northcentral,

Widespread

Distribution

Climbers(case a

Lotic—

hydrophytes)

North American

slightly curved cylinder of leaf pieces)

strands)

erosional

Lentic-pools

herbivores

shaped" tube of long vegetation

Shredders-

Climbers (case a

rough,

Lentic—

vascular

Oligostomis)

Climbers (case similar to

Lentic—

sphagnum bog pools

predators (engulfers)

two instars

instars, filamentous

green algae); last

irregular strands of vegetation)

deposltional

(chewers; early

Agrypnia but with some

hydrophytes:

herbivores

lotic—

Shredders—

Climbers(case similar to

Lentic—

Habit

Trophic Relationships

vascular

Habitat

*SE = Southeast, DM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

of species in parentheses)

Taxa(number

Table 19A

MA*

Ecological

6524

(continued)

3229, 5508,

6524

6524

6532

160, 3944, 6513, 6524, 5778, 6648, 6649

References**

TRICHOPTERA

NW

Tolerance Values

))))))))) J ) ) ) D ) ) )))) )))) ) ) J

~4

Caddisflies

Western

Rossianidae (2) -

Family

Continued

Yphria

Ptilostomis(4)

Phryganea (2)

Oligotricha

(chewers):

constructed of

predators (engulfers)

a curved

cylinder of

and erosional

wood pieces)

mineral and

Clingers— sprawlers (case

depositional

(engulfers)

Predators

facultative

hydrophytes) Lotic—

(chewers);

vascular

herbivores and detritivores

Hagenella)

and

depositional (detritus and

shredders—

Climbers (case similar to

Lotic—

Facultative

predators (engulfers)

facultative

erosional

hydrophytes) leaf pieces)

herbivores and detritivores

(detritus and vascular

and lentic

Facultative shredders—

Climbers (case

a spirally arranged. tapered cylinder

depositional

(engulfers)

Predators

California, Oregon

except Southwest

Widespread

Widespread

Alaska, Yukon

Trophic American Relationships** Distribution

Lotic—

Agrypnia)

Climbers(case similar to

Lentic—

Habit

littoral

Habitat

North

6.7

SE

5.0

UM

M

NW

5.0

MA*

Tolerance Values

Ecological

6527, 1380

160, 6516, 6524

3562,6513, 6524, 6648, t

481, 2060, 2300, 3229, 3562, 5448, 6513, 4337, 4423, 6524, t

6524

References**

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships tUnpublished data, K. W. Cummins, Michigan State University Kellogg Biological Station, University of Pittsburgh Pymatuning Laboratory of Ecology, University of Maryland, Appalachian Environmental Laboratory, and Oregon State and Humboldt State Universities

Order

of species in parentheses)

Taxa(number

Table 19A

) ) ) > ) ) ) )))) ) ) ) ) ) ) ) ) ) ) ) ) > ) )

ON

o\ w

Caddisflies

(32)- Little

Thremmatidae

Family

Continued

montana

Rossiana

Oligophlebodes(7)

Neophylax(25)

baumanni

Species

Goereilla

Genus

erosional

Lotic—

erosional

Lotic—

depositional (especially In moss)

and

erosional

Lotic—

depositional (springs)

Lotic—

Habitat

Habit

mineral)

Clingers (case strongly tapered and curved, rough

side)

stones on each

with ballast

curved, mineral

Clingers (case tapered, slightly

minerals)

Clingers (case tapered, curved, rough

curved, mineral)

Clingers (case tapered,

North

gatherers

collectors-

Scrapers;

Obligate scrapers

(chewers)

herbivores

and shredders—

Probably scrapers

gatherers

Collectors—

West

Southwest

Widespread except

Northwest

Northwest

Trophic American Relationships** Distribution

1.6

SE

3.0

UM

M

1.0

3.0

NW

MA*

Tolerance Values

Ecological

6524

4430, 4561,

5393, 5404, 6524, 372, 2592, 3841, 1534, 2594, 3742, 2593, 3741, 3010, 2676, 3992, 3993, t

160, 1073, 1077, 3640, 4077,

6524

6523, 6524

References**

Laboratory, and Oregon State and Humboldt State Universities

TRICHOPTERA

(continued)

tUnpublished data, K. W. Cummins, Michigan State University Kellogg Biological Station, University of Pittsburgh Pymatuning Laboratory of Ecology, University of Maryland, Appalachian Environmental

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

of species in parentheses)

Taxa (number

Table 19A

Caddisflies

Sericostriata

Neothremma (7)

Farula (12)

surdickae

Habitat

erosional (on rocks)

Lotic—

erosional (on rocks)

Lotic—

including seeps)

erosional (on rocks,

Lotic—

Lotic—

Species erosional

Genus

Uenoidae (20)

Family Habit

slender, mineral)

Clingers (case tapered, slightly curved, long,

slender mineral)

Clingers (case tapered, slightly curved, long,

mineral)

Scrapers;

Clingers (case tapered, curved, long, very slender,

Montana

Idaho,

West

Pacific States

Distribution

American

North SE

UM

M

0.0

0.0

0.0

NW

MA*

Tolerance Values

Ecological

6543

6524, 4430

160, 1858, 3985,

160, 6524

160,6524,6543, 6527, 1380

References**

I ) ) ) ) ) > ) ) ) ) ) ) ) ))) ) )) > ) )

gatherers

collectors—

Scrapers;

gatherers

collectors—

Scrapers;

gatherers

collectors—

gatherers

collectors—

Scrapers;

Trophic Relationships**

slender, curved, mineral)

Clingers (case tapered,

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

of species in parentheses)

Continued

- Stone-case

Table 19A

Taxa(number

)) J

ON

-4

jjHtt *r'-

^ '7"'

^ • -

AQUATIC AND

SEMIAQUATIC LEPIDOPTERA M. Alma Soils

Systematic Entomology Laboratory, ARS, USDA,Smithsonian Institution

Washington, D.C.

INTRODUCTION

Aquatic Lepidoptera, moths with one or all stages morphologically adapted for living in water, occur primarily in the subfamily Acentropinae (Pyraloidea; Crambidae). The larvae have developed various methods for respiration in water, from being hydrophilic in early instars to hydrophobic in later instars, either involving plastron-like layers of air or oxygen within cases made from their host plants, or from the interstitial spaces of the host plant, or uniquely in the Lepidoptera, from tracheal or blood gills (Welch 1922)(Fig. 20.1). In North America, most acentropine larvae are polyphagous feeders on floating and emergent plants in lakes or ponds, and algae on rocks in fast and slow-moving rivers and creeks. Other acentropine host plants and feeding habits have been observed or postulated in other parts of the world, such as feeding on mosses and liverworts in Japan (Yoshiyasu 1980), and on black flies in Brazil (Gorayeb and Finger 1978). In the noctuid Bellura the first three instars are gregarious leafminers, and in later instars the larvae bore into stems of emergent

plants (Fig. 20.12)(Levine and Chandler 1976). Most acentropines have immatures adapted for survival in aquatic habitats, but the adult is not aquatic. The adults fly and mate terrestrially, feeding on flowers. They often come to collecting lights in great numbers in North America, especially Petrophila, near bodies of water.

The Acentropinae has over 700 species worldwide in temperate and tropical regions, and reaches its greatest diversity in the Old World tropics. Only 50 species in 15 genera, including two exotic species, are found in the United States (Scholtens and Solis 2015). Munroe(1972-1973) updated the taxonomy of

the North American Acentropinae (as the subfamily Nymphulinae)and included three tribes, Ambiini ter restrial species that feed on ferns, and Nymph ulini and Argyractini aquatic species. In his study of the Palearctic Acentropinae, Speidel (1984) resurrected the subfamily name Musotiminae for the Ambiini, and removed the terrestrial, fern-feeding tribe from the Acentropinae. In addition, Speidel showed that Acentria belonged in the Acentropinae, not the Schoenobiinae. Several other taxonomic changes have been made in the North American acentropines.

These include the largest genus in the United States, Petrophila (Fig. 20.29), being found to be the senior name and Parargyractis, thejunior synonym(Munroe 1983). The North American Munroessa (Fig. 20.28) was shown to belong to the Old World genus Elophila by Yoshiyasu(1985)in his excellent study ofJapanese acentropines, and Speidel (2005) synonymized Synclita with Elophila (Fig. 20.27). The introduced species, Petrophila drumalis (Fig. 20.23), was trans ferred to the genus Argyractis based on the similar larval morphology and biology with A. subornata, which feeds on the roots of water hyacinth in Brazil (Forno 1983; Habeck and Solis 1994). A recent pre liminary study of all the species in Argyractis showed that these two species are not congeneric with the type species of Argyractis, and will require a new genus. According to Lange(1956a) the aquatic acentro pines were clearly separated into two tribes based on morphology and biology. The Nymphulini that live in standing or slowly moving water, and may or may not have gills at some stage, and the Argyractini that live in slow or fast moving water under silken webs on rocks scraping algae and have gills. The separation of the Acentropinae into two tribes on a worldwide level

765

766

Chapter 20 Aquatic and Semiaquatic Lepidoptera

is no longer valid as shown morphologically by Yoshiyasu (1985). Munroe (1995, pers. comm.)in his checklist of Neotropical Acentropinae chose not to separate the genera into tribes. Additionally, in the Western Hemisphere the discovery of three argyractine genera that feed and live on aquatic plants nulli fies the Argyractini biological definition by Lange (e.g., Forno 1983; Fiance and Moeller 1988; Dray

tunnels into the leaf, sometimes boring into the peti ole. Later instars develop tracheal gills and create cases in various ways. The larvae feed on leaves near their cases and some were seen to leave their case and

et al. 1989). The immatures of Oxyelophila and Usingeriessa were described recently (Soils et al.

feed on the leaf surface. Buckingham and Bennett (2001)also verified that in P. maculalis larvae, move ments replenish oxygen(Welch and Sehon 1928). The pupa ties the case completely closed with a silk cocoon in the case. Respiration is through "spots" on the surface of the leaf. The sexually dimorphic males

2018). In addition, nothing is known about the

and females mate outside the water for about an

biology and immature morphology of Chrysendeton, Contiger, and Oligostigmoides. Langessa nomophilalis has been reared on a wide variety of aquatic plants (Herlong 1979, Stoops et al. 1998), but its biology and its immatures have not been described. The biology of Neocataclysta magnificalis is unknown. It was pre sumably described by Forbes(1911)(as Elophila sp.), but the species identity is in doubt and it is now believed to be £ obliteralis(Munroe 1972). The most biologically fascinating species in the Acentropinae is Acentria ephemerella, a species that was accidentally introduced into the United States and first reported here by Forbes in 1938(Figs. 20.24

hour. Parapoynx has Old World affinities and there is much literature associated with them from Europe

and 20.25). This species occurs in the Palearctic region and there is much literature associated with it from

Europe. The larvae are case makers in standing water (Fig. 20.16). The parthenogenetic females are aquatic with reduced wings and rely on a plastron for respira tion. Winged, reproducing females swarm with winged males to mate and then disperse. First instar larvae bore into the stems, and later instars overwin ter in a shelter made with silk and bits of leaves. The

pupa breathes oxygen that is released into the cocoon by the host plant. Acentria ephemerella feeds on Myriophyllum, Elodea, Hydrilla, Potamogeton, and Ceratophyllum species, among others (Berg 1942; Batra 1977; Buckingham and Ross 1981; Wichard et al. 2002). Acentria ephemerella is important to lake community dynamics. It was shown to affect entire macrophyte populations in Cayuga Lake, New York due to its preference for Myriophyllum (Gross et al. 2001).

and Asia.

Species with known biologies of Nymphuliella and Elophila also make cases, but lack gills in all instars (Figs. 20.2 and 20.3). They are completely submerged in standing or slowly moving water and feed on floating or emergent vegetation (Neunzig 1987; Habeck 1991) and presumably breathe with a plastron type ofrespiration(Berg 1949,1950a;Thorpe 1950; Wesenberg-Lund 1926). Early instar larval res piration may be cuticular as in some Elophila (Wesenberg-Lund 1943), and later instars become surface feeders(Welch 1924). Nymphuliella daeckealis

lives in a completely submerged case in water holes of sphagnum bogs in Maine (Heinrich 1940; Munroe 1972). Elophila, with Old World affinities, has much European and Asian literature associated with them.

In Elophila, although two species are known to lack gills, they reportedly have very different habitat and feeding behaviors. In E. icciusalis the larvae make cases, but in E. gyralis early instars feed on leaves, scraping the lower epidermis, and later instars bore into the stem of water lilies (Neunzig 1987). The 17 species of Petrophila live in fast-flowing streams, intermittent streams, and stagnant pools, and are scrapers of algae and diatoms on the surfaces of rocks (Lange 1956a; Tuskes 1977, 1981; Neunzig 1987)(Figs. 20.1, 20.5, 20.6). The larvae lack gills in the first instar, but tracheal gills are present in subsequent instars. In the western United States, immatures of Petrophila confusalis are found in

Parapoynx larvae are also case makers, but

well-oxygenated streams and lakes where the water

although they lack gills in the first instar, tracheal gills are present in subsequent instars (Fig. 20.7). Known species of Parapoynx are leafminers first and in later instars they create a case from the young leaves ofthe host plant and are completely submerged (Forbes 1910; Welch 1916). Buckingham and Bennett (2001) published an excellent study of P. seminealis

velocity is between 0.4 and 1.4 m/sec (Tuskes 1981).

found in eastern United States. The female sits on the

edge ofthe leaf, the abdomen is curled under, and the eggs are laid under floating leaves. The first instar

The larvae construct silk tents on rock under which

they feed and in Montana larvae are parasitized by an ichneumonid wasp (Jamieson and Resh 1998). The pupal cocoon consists of an inner cocoon surrounded by an outer cocoon with holes to allow water circula

tion (Fig. 20.11). Prior to pupation, the larva cuts an opening in the inner cocoon to allow adult emergence. The emerging adult reaches the surface where the wings expand. After mating,the adult female deposits

Chapter 20 Aquatic and Semiaquatic Lepidoptera

eggs on a rock (Fig. 20.1 a, b). In northern California there are two or three generations per year of P. confusalis, but in western Montana McAuliffe and Williams (1983) report only one generation a year. Population numbers and distribution are dependent on water temperature, concentration of dissolved oxygen, substrate texture, and algal growth (Lange 1956a; Tuskes 1977, 1981). Other species with gills are Argyractis drumalis caterpillars that are found in lakes, canals, and slow streams, and feed on the rootlets of floating Pistia (Dray et al. 1989). Eoparargyractis plevie caterpillars are found in lakes and feed on emergent aquatic plants, Lobelia and Isoetes species (Fiance and Moeller 1988)(Figs. 20.4 and 20.8). Neargyractis slossonalis caterpillars are found in rivers feeding on the roots ofemergent plant species,such as Vitis,Fraxinus, and Taxodium, and in lakes feeding on floating Eichornia species (Habeck 1988). Usingeriessa onyxalis caterpillars feed on Hygrophila and Eleocharis and Oxyelophila callista feeds on five species of aquatic plants (Soils et at. 2018). Much ofthe modern literature on the biology and ecology of plant-feeding aquatic and semi-aquatic Lepidoptera concerns the need to control invasive noxious aquatic weeds(Center et al. 1999; Harms and Grodowitz 2009). For example, the studies on the following aquatic plants elucidated the biology of the lepidopterans that feed on them: water hyacinth by crambids Argyractis subornata (Acentropinae) (Forno 1983), NiphograptaalbiguttalisiSpWomeMnsLc), Samea multiplicalis (Spilomelinae), and the noctuid, Bellura densa; water lettuce by crambids Sameodes albiguttalis (Spilomelinae)(Center 1983; Dray et al. 1989), Samea multiplicalis (Spilomelinae), Argyractis drumalis(Acentropinae), and the noctuid Spodoptera pectinicornis; hydrilla by Parapoynx diminutalis (Crambidae: Acentropinae) (Balciunas and Center 1981; Balciunas and Minno 1985; Batra 1977; Buckingham and Bennett 1989); alligator weed by Arcola malloi (Pyralidae: Phycitinae); smartweeds or speciesofPolygonum,by Ostriniapenitalis(Crsimhidae: Pyraustinae) and the noctuid, Simyra insularis (Herrich-Schaffer). Recently, studies in Hawaii have discovered over 15 species of Hyposmocoma (Cosmopterigidae) larvae to be aquatic, although many more are terrestrial. These larvae have evolved a diversity of case architecture and are herbivorous (Schmitz and Rubinoff 2008; 201 la, b). Some lepidopteran families have immatures that are associated with floating or emergent aquatic plants such as cattail {Typha spp.), bulrush (Scirpus spp.), sundew (Drosera sp.), or other vascular hydrophytes, but the immatures have not developed morphological

767

adaptations for underwater respiration. The larval feeding habits of these species include leafmining, foliage feeding under webs on the surface of leaves, stem or root boring, and behaviors such as movement to flower or seed structures for pupation above the waterline and host switching in later instars. Caterpillars in this category belong to the families Crambidae(Crambinae: Crambus,Chile-, Schoenobiinae: Donacaula; Pyraustinae: Ostrinia(Welch 1919; Ainslie and Cartwright 1922)), Nepticulidae, Coleophoridae (Ellison 1991), Cosmopterigidae, Gelechiidae, Tortricidae, Olethreutidae, Noctuidae(Claassen 1921; Wagner etal.2011), Arctiinae in Erebidae(WesenbergLund 1943; MacKay and Rockburne 1958; Vogel and Oliver 1969; Levine 1974; Levine and Chandler 1976), Cossidae, Hesperiidae, and Sphingidae(Hagen 1880). The literature for these species is extensive and is not treated here, but some citations for common species are included in the summary of the ecological and distributional data for Lepidoptera.

EXTERNAL MORPHOLOGY

Eggs The eggs of Lepidoptera are laid singly or in clusters or groups. In many crambids, the eggs are often flattened and deposited in groups of overlap ping layers, and some eggs are ovoid (Peterson 1963). In Petrophila confusalis, 85 to almost 300 eggs are laid in a cluster; each egg is smooth and light colored with the micropyle (minute opening) toward the edge of the cluster (Tuskes 1977)(Fig. 20.1b). In Petrophila bifascialis 300-550 eggs in a cluster have been reported to be laid by a single female (Kubik 1981). In the Acentropinae, females lay eggs on exposed or submerged vegetation, as in Parapoynx,or on rocks as in Petrophila. Commonly, such as in noctuids, eggs are upright, ribbed with a depressed micropyle, and are laid singly. In con trast, Bellura gortynoides females lay egg masses, each with about 14 eggs each, and are covered with dark brown scales (Levine and Chandler 1976). Bellura females lay eggs on exposed leaves and occa sionally on petioles. Larvae

Lepidopterous larvae (Fig. 20.3) are character ized by: (1) a distinct head with stemmata or simple eyes;(2)chewing mouthparts;(3)spinneret;(4) a thorax, each segment with a pair of legs; (5) 10 abdominal segments with prolegs on segments 3,4, 5,6, and 10 (anal prolegs); and (6) spiracles on the prothoracic segment and abdominal segments 1-8

768

Chapter 20 Aquatic and Semiaquatic Lepidoptera

(Lange 1996; Stehr 1987). The position of larval setae is used for identification and nomenclature

(Heinrich 1916; Hinton 1946; Common 1970; Stehr, 1987). Secondary setae, setae from a flattened, pigmented area, called a pinaculum, or a chalaza, if

elevated, are present in some families of the Lepidoptera, but they do not occur in the Acentropinae and Bellura. Most recently, six species of Acentropinae larvae are figured and described by Neunzig (1987). Bellura gortynoides is figured and described by Godfrey (1987). In the Acentropinae, larval respiration may be cuticular as in Elophila (Wesenberg-Lund 1943), which is without gills, or with open or closed tracheal gills, usually filamen tous processes with a network of tracheoles just under the cuticle. Tracheal gills and/or blood gills, hollow filaments through which blood circulates, are

present only in certain instars, and can occur in the abdomen as well as the thorax, are compound or simple, and occur dorsally, laterally, and/or ventrally (Figs. 20.4, 20.5, 20.6, 20.7, 20.8, 20.9, 20.20a). Species of Parapoynx and Petrophila have a nongilled first larval instar, and after molting to the second instar develop tracheal gills. Other genera, such as Elophila and Neocataclysta lack gills in all instars (Berg 1949, 1950a), and a plastron type of

respiration has been suggested for E. icciusalis(Berg 1949, 1950a; Thorpe 1950; Wesenberg-Lund 1926). The cuticle oflarvae in later instars can be composed of microtrichia that create the plastron (Fig. 20.21b); sometimes the plastron is incomplete and larvae obtain oxygen from their host plant (Petrischak 2000; Wichard et al. 2002). Later instars of B. gor

tynoides bore into petioles of water lilies, and use the dorsally located spiracles of the 8th segment for res piration above the water line (Fig. 20.12). Head: In most Lepidoptera and some Acentropinae, such as E. obliteralis, the head capsule is vertical (hypognathous) with the mouthparts directed downward (ventrally)(Fig. 20.11). In some leaf mining Lepidoptera and many Acentropinae the mouthparts are directed forward and the head cap sule can be horizontal (prognathous) as in E. plevie (Fig. 20.10), or at an angle (semiprognathous) as in Parapoynx. Most acentropine species have progna thous (e.g., Petrophila) or semiprognathous heads and lack or have a very short epicranial suture. But the ratio of the length of the adfrontal suture to the epicranial suture when present can sometimes be used to distinguish Lepidoptera species. The stemmata are usually six in number but in acentropines can be reduced in number, poorly developed, or lost (Fig. 20.3). The clypeus and labrum are ventrad or anterad to the frons(Fig. 20.2Id). In acentropines the labrum

may have brush-like or spatulate setae as in N. slossonalis(Habeck 1988), E. plevie (Fiance and Moeller 1977), and A. subornata (Habeck and Solis 1994). These modified setae have been postulated to indicate

feeding on periphyton, but this has not been docu mented. The antennae are usually short and bear sev eral sensillae in Lepidoptera, but in acentropines the second segment can be three times or more as long as thick. A spinneret (silk-producing structure) (Figs. 20.10, 20.11, 20.20d, 20.21d) is used to build path ways along leaves, rocks, and cocoons and their cov ers, e.g., Petrophila. The cover is created by the final instar by detaching the edges of the webs, pulling them inwards, and spinning them together to form a very tough cover, underneath which they next build a cocoon (Kubik 1981). There are unattached portions of the cover for water circulation (Fig. 20.11). Mouthparts are the chewing type with opposable toothed mandibles,and very toothed in Acentropinae. Thorax: The prothorax (Tl) usually bears a prothoracic shield, often sclerotized or patterned, with one pair of thoracic legs (Fig. 20.3). The mesothorax (T2) and metathorax (T3) also have thoracic legs each five-segmented with claws. Thoracic seg ments are usually without gills, but present in some species for example, compound gills arising from a single base (Fig. 20.7) as in Parapoynx allionealis, or simple gills in Petrophila santafealis and E. plevie (Figs. 20.5 and 20.6). Abdomen:In most Lepidoptera the abdomen con sists of 10 segments, with pairs of prolegs on segments 3, 4, 5, 6, and 10, sometimes reduced or absent (Fig. 20.3). Each proleg has curved hooks called crochets (Figs. 20.13, 20.14, 20.15, 20.20c, 20.21c) for traction during movement. Each crochet is usually circular (Figs. 20.13b, 20.14b), but in Bellura they appear as hooks(Fig. 20.15b). The length, pattern, and arrange ment may vary and therefore be useful for classifica tion. The crochets are uniordinal if they are the same length, if two or three lengths, they are biordinal or triordinal as in most acentropines. Crochets can be in a complete circle, in an interrupted circle or penellipse as in many crambids, in a single band extending longi tudinally on the mesal side, or a mesoseries, as in noctuids. Uncharacteristically for the Crambidae, in acentropines the crochets are elliptical, but they can be an incomplete ellipse that is open laterally and mesially and therefore appear like transverse bands.

Pupae The typical lepidopteran pupa is of the obtect type (appendages and body compactly united) and may or may not be enclosed in a cocoon (Figs. 20.16,

Chapter 20 Aquatic and Semiaquatic Lepidoptera

20.17, 20.18). In Acentropinae species spiracular

openings can be reduced, or enlarged and protruding (Figs. 20.Id and 20.2d). Some acentropine species may have external spiracular openings on abdominal segments 3 and 4 that are greatly enlarged and pro truding, and at times spiracles on abdominal seg ments 1 and 2 less so. Enlarged and protruding spiracles also occur in terrestrial forms, e.g., Musotiminae, and Noctuidae, e.g., Spodoptera pecticornis. The cremaster, or last segment of the abdo men,is adapted to firmly anchor the pupa to its silken case (Fig. 20.18) (Mosher 1916). Petrischak (2000)

reported sexual dimorphism in pupae and stridulation in a European acentropine species.

Adults

Lepidoptera adults are characterized by a pro boscis or haustellum and the presence of overlapping scales (modified setae) on two pairs of wings, body, and legs (Figs. 20.19 and 20.32)(Scoble 1992). In the adults the ribbed scales are what create the plastron and allow the adults to emerge from water unscathed, and allow females to enter the water wholly or partly to oviposit(Tuskes 1977; Wichard et al. 2002). Head: The Pyraloidea has scales at the base of the proboscis (Fig. 20.19), and in the Noctuoidea scales at the base of the proboscis are absent. The compound eyes are well developed. Ocelli are prominent in most Acentropinae, but absent in Acentria, and in the noctuid Bellura. Chaetosemata, sensory organs with spe cialize setae behind the compound eye, are prominent in Acentropinae, and absent in Bellura. In the Lepidoptera, antennae are clubbed, hooked,or serrate (saw-like), pectinated in Bellura and ciliated or annulated in Acentropinae. Maxillary palpi are prominent and easy to see in acentropines. The overall position of labial palpi is variable in Lepidoptera, but in most Acentropinae labial palpi are usually upright (Fig. 20.32), but porrect(pointing forward)in, for example, some Elophila,and decumbent in Acentria(Fig. 20.33).

769

Thorax: The prothorax possesses a pair of over lapping plates, the patagia, and the well-developed mesothorax has a pair of laterally placed tegulae that cover the base of the wings. The metathorax is usually inconspicuous, and the legs are long and thin in the Acentropinae. In some aquatic Acentropinae, the hind legs possess an oar-like fringe of hairs used in swimming. Wings: In the Lepidoptera wing colors and patterns are useful for identification of species (Figs. 20.22-20.31). Some families may have brachypterous forms in one or both sexes, and in a few species (e.g., Acentria ephemerella) both winged and brachypterous females occur (Figs. 20.24 and 20.25). Many acentropines and a few other crambids have "a row of black spots with contrasting white, blue, or metallic pupils or interspaces on or near and parallel to the terminal margin of the hind wing" that Munroe (1991) termed "cataclystiform" (Figs. 20.22, 20.23, 20.29). The biological significance of these beautiful spots in Neargyractis, Usingeriessa, Petrophila,Eoparargyractis,Neocataclysta,Chrysendeton, and Argyractis is unknown. Abdomen: Tympanal organs (hearing) occur at the base of the abdomen in the Pyraloidea; in crambids there is a unique flap over the tympanal organ called a praecinctorium. The praecinctorium in acen tropines is simple, and not bilobed as in other crambids. In the Noctuoidea tympanal organs are located on the metathorax. The genital structures at the ter minal end of the 10-segmented abdomen are used for generic and species classification. In most lepidopteran groups males and their variation in of the valvae, phallus, juxta, tegumen, and uncus are very useful. Unlike most groups, the acentropine males are not very variable, but the female genitalia have strong variation in an ostium bursae (copulatory opening), ductus bursae (egg canal), and especially the corpus bursae, with a great diversity in patterns of scobinations or spines, can be diagnostic for species(Habeck and Solis 1994).

KEY TO LARVAE OF AQUATIC LEPIDOPTERA

The following genera have unknown immatures in North America: Neocataclysta, Chrysendeton, Contiger, and Oligostigmoides. This key was developed from parts of Lange (1996), Habeck, unpublished, and recent studies (Solis et al. 2018). Note: The Pyraloidea and Noctuoidea can be separated from most other Lepidoptera by the presence of two prespiracular setae present on the prothorax. 1. Proleg with crochets in a mesoseries, or if an incomplete circle, crochets hook-like (Fig. 20.15).... 2 1'. Proleg with crochets in a circle or incomplete circle, an ellipse or sometimes appearing as transverse lines

2(1).

Crochets in a mesoseries, spiracles on 8th segment normal

3

NOCTUOIDEA,not Bellura

770

Chapter 20 Aquatic and Semiaquatic Lepidoptera

2'.

Crochets in an almost incomplete circle, spiracles on 8th segment specialized (Fig. 20.12)... .Bellura

3(1').

Body without Filamentous gills (Fig. 20.3)

3'. 4(3).

Body with filamentous gills (Fig. 20.5) Crochets in a circle or incomplete circle

4'.

Crochets in an ellipse or an elliptical circle, sometimes appearing as transverse lines, larvae living in cases (Fig. 20.16), case made of leaves or plant material, usually associated with lakes, ponds, or quiet water (Fig. 20.17) 6 With a membranous sac or protuberance anterior to prothoracic coxae; without a transverse plate posterior to dorsal pinacula on mesothorax (Fig. 20.34) SCHOENOBIINAE

5(4).

4 9 CRAMBIDAE(in part), 5

5'.

Without a membranous sac anterior to prothoracic coxae; with a single transverse plate posterior to dorsal pinacula on mesothorax (Fig. 20.35) CRAMBINAE

6(4').

Lateral setae of T1 on ventral extension of prothoracic shield

6'.

Lateral setae of T1 not on prothoracic shield (Fig. 20.3)

7(6').

Crochets triordinal

Langessa

T.

Crochets biordinal

8

8(7').

Cephalic and caudal rows of crochets same size (Fig. 20.14) or cephalic row of crochets distinctly larger than caudal row (Fig. 20.13)

Elophila

Caudal row of crochets distinctly larger than cephalic row; larvae in retreats made by fastening together aquatic plants

Acentria

8'.

Nymphuliella 7

9(3').

All gills branched (Fig. 20.7); larvae living in cases cut from leaves of aquatic plants

10

9'. 10(9).

Without branched gills or with only a few (Figs. 20.5 and 20.6) With abdominal spiracles

10'.

Without abdominal spiracles

11(9'). 11'.

Head prognathous, body dorso-ventrally flattened, gills lateral (Fig. 20.10) Head hypognathous, gills generally distributed (Fig. 20.11)

12(11). 12'.

Gills on A9 and AlO extremely dense and long, forming a fan (Figs. 20.4 and 20.8)...Eopamrgymctis Gills uniformly distributed on segments, primarily lateral (Figs. 20.5 and 20.6); head flattened dorsoventrally; prognathous(head horizontal and mouthparts directed forward)(Fig. 20.10); thorax and abdomen with numerous blood gills (Figs. 20.1c, 20.5, 20.6); mandibles prominent, adapted to scrape algae and diatoms from rocks in streams, lakes, and springs Petrophila

11 Parapoynx Usingeriessa 12 13

13(11'). With numerous ventral short gills (Fig. 20.9) Argyvactis 13'. Without ventral gills 14 14(13'). A1-A7 with a transverse row of dorsal gills; without modified labral setae (Fig. 20.36). .Oxyelophila 14'. AI-A7 without a transverse row of dorsal gills; with modified labral setae (L2 and L3 brush-like; M3 spatulate)(Fig. 20.37) Neargymctis

KEY TO ADULTS OF AQUATIC LEPIDOPTERA

This key is developed from parts of Lange (1996), Munroe (1972), and Munroe and Soils (1999). 1. r. 2(1').

Adult with metathoracic tympanal organs, without scales at the base of the proboscis NOCTLOIDEA,Bellura Adults with abdominal tympanal organs, with scales at the base ofthe proboscis.... PYRALOIDEA,2 Tympanal organ without praecinctorium PYRALIDAE

2'.

Tympanal organ with praecinctorium

3(2').

Chaetosema present, forewing with distal part of CuP developed as a tubular vein

CRAMBIDAE,3

4

Chapter 20 Aquatic and Semiaquatic Lepidoptera

3'. 4(3). 4'. 5(4).

771

Chaetosema absent, forewing with CuP absent ... CRAMBIDAE,not Acentropinae or Schoenobiinae Rs[ offorewing stalked with RS2+3, proboscis normal, larvae in silk webs on rocks or feeding on submerged plant tissues ACENTROPINAE,5 Rsi offorewing most often separate from RS2+3, proboscis reduced, larvae borers in semiquatic graminaceous plants SCHOENOBIINAE Proboscis short, rudimentary, or missing; species with brachypterous wings in some females; male with normal wings

Acentria

5'.

Proboscis normal length; wings usually fully developed in males and females

6(5'). 6'.

Hind wing with M2 present Hind wing with M2 absent

7(6).

Forewing with veins M3 and Cui stalked

7.

Forewing with veins M3 and Cui separate

8(7').

Forewing and hind wing with margin excavated behind Mj; eighth sternite of male produced backward in a broad triangular process Wings with margin at most weakly excavated behind Mj; posterior triangular process of eighth sternite of male abdomen absent or very narrow

19

9(8).

Outer margin of hind wing with a series of black and metallic spots

10

9'.

Outer margin of hind wing without such spots

11

10(9).

Black spots distinctly separate, with pupillate bluish centers; CuP of hind wing complete

Neocataclysta

Black spots not distinctly separate, metallic spots not pupillate; CuP of hind wing vestigial

Chrysendeton

8'.

10'.

6 7 14 Langessa 8 9

11(9').

Apex offorewing angular

12

11'.

Apex offorewing rounded

13

12(11).

Size small; color blackish; second segment of labial palpus long, with rough thick scaling, third segment relatively short and slender, the palpus weakly ascending

Nymphuliella

12'.

Size medium; color white, with black-borders, straw-yellow bands; labial palpus short, moderately upturned, with relatively short, curved second segment, its rough irregular scaling grading to the slender scaling of the third segment Elophila(= ekthlipsis)

13(11').

Antennae strongly annulated

13'.

Antennae with at most weakly raised scale-rows on each segment dorsally

14(6').

Forewing with apex falcate

14'.

Forewing with apex not falcate

15

15(14').

Hind wing with discal cell open

16

15'.

Hind wing with discal cell closed

17

16(15).

Wings with lines bright yellow

16'.

Wings with most lines brown

17(15').

Hind wing with discal cell more than half length of wing; outer margin of hind wing not emarginated behind apex Hind wing with discal cell at mostjust over half length of wing; outer margin of hind wing emarginate behind apex

17'.

Pampoynx Elophila Oxyelophila

Argyractis Eoparargyractis Petrophila 18

18(17'). 18'.

Veins Sc+R and Rs of hind wing separating halfway between cell and apex of wing ... Usingeriessa Veins Sc+R and Rs of hind wing separating just before apex of wing Neargymctis

19(8').

Termen of forewing excavated; discocellular (or cross) vein of hind wing almost straight....Contiger

19'.

Termen of forewing not appreciably excavated; discocellular (or cross) vein of hind wing strongly curved

Oligostigmoides

112

Chapter 20 Aquatic and Semiaquatic Lepidoptera

Figure 20.1

Figure 20.2

Figure 20.1 Life stages of Petrophila confusalis (Walker)(Crambldae)(after Lange 1956a). a, adult male; b, eggs; c, mature larva; d, pupa; e, larval web; f, cocoon cover.

Figure 20.2 Life stages of Elophila occidentalis Lange (Crambidae)(after Lange 1956a). a, adult male; b, eggs; c, mature larva; d, pupa.

metathorax

spiracles

mesothorax

-prothorax

head

caudal

prolegs stemmata

crochets

abdominal prolegs thoracic legs

Figure 20.3

filamentous gills

Figure 20.4 Figure 20.5

Figure 20.6

gill tuft

Figure 20.7 spinneret

Figure 20.8

antenna

Figure 20.9

Figure 20.10

Figure 20.3 Lateral view of caterpillar Elophila obliteralis (Walker)(Crambidae). Figure 20.4 Lateral view of larva Eoparargyractis plevie Dyar (after Fiance and Moller 1977). Figure 20.5 Dorsal view of gilled larva of rockdwelling type, Petrophila confusalis (Walker)(Crambidae)(after Lange 1996). Figure 20.6 Gills of Petrophila confusalis (Walker) (Crambidae)(after Lange 1996). Figure 20.7 Compound gill of Parapoynx sp. (Crambidae)(after Lange 1996).

spinneret

Figure 20. spinneret

Figure 20.8 Gill tufts on posterior segments of Eoparargyractis plevie Dyar (after Fiance and Moller 1977).

Figure 20.9 Ventral short gills of Argyractis sp. (after Habeck and Soils 1994). Figure 20.10 Lateral view of prognathous head of Eoparargyractis pievie Dyar (Crambidae). Figure 20.11 Lateral view of hypognathous head of Elophila obliteralis (Walker)(Crambidae).

773

spiracle

Figure 20.12

"

Figure 20.13

Figure 20.14

"X..

Figure 20.15

(

(

is?

I \

x;.

/ ^

a

antenna

chaetosema

Figure 20.16

maxillary palp

Figure 20.17

spiracles

labial palp

Figure 20.18

Figure 20.19 haustellum(= proboscis)

Figure 20.12 Eighth and ninth abdominal segments of Bellura obliqua (Waiker)(Noctuidae). Figure 20.13 Crochets of Acentria ephemerella Denis and Schiffermuller (Crambidae). Figure 20.14 Crochets of Elophila sp.(Crambidae). Figure 20.15 Crochets of Bellura obliqua (Walker) (Noctuidae)(on abdominai segments 3-6). Figure 20.16 Larval retreat of Acentria sp. (Crambidae) on Ceratophyllum sp. (after Lange 1996). 774

Figure 20.17 Larval case of Elophila occldentalis Lange (Crambidae)(after Lange 1996). Figure 20.18 Lateral view of pupa of Petrophila confusalis (Waiker)(Crambidae) showing enlarged spiracular openings on specific abdominal segments (after Lange 1996). Figure 20.19 Lateral view of aduit Lepidoptera head (after Lange 1996).

Chapter 20 Aquatic and Semiaquatic Lepidoptera

-

775

V sa'

B5x?3S5 o

5.,t\,\>..iJt'4

-> -a • •,. 1

i

MSt

lOOHM

lOKV

00

001

S

Figure 20.20 SEMs of Parapoynx diminutalis Snellen (Crambldae) larvae, a, lateral view of head and thorax; b, integument; c, proleg and crochets; d, mouthparts.

Figure 20.21 SEMs of Elophila obliteralis (Walker) (Crambldae) larvae, a, lateral view of head and thorax; b, integument; c, proleg and crochets; d, mouthparts.

'j

±

X Figure 20.22

■•Si

Aa

Figure 20.24

Figure 20.23

Figure 20.25

j -sy

w •-■-•- < -i:dt^-' .u

i ■ :

Figure 20.26

Figure 20.27

Figure 20.28

5.0 mm

Figure 20.29

Figure 20.30

Figure 20.32

Figure 20.31

Figure 20.22

Chrysendeton medicinalis (Grote)

(Grambidae).

Figure 20.23 Argyractis drumalis Hampson (Grambidae). Figure 20.24 Male Acentria ephemerella (Denis and Schiffermulier) (Grambidae). Figure 20.25 Female Acentria ephemerella (Denis and Schiffermulier) (Grambidae). Figure 20.26 Parapoynx alllonealls (Walker) (Grambidae). Figure 20.27 776

Elophlla obllteralls (Walker) (Grambidae).

Figure 20.33

Figure 20.28 Elophlla Icclusalls (Walker) (Grambidae). Figure 20.29 Petrophlla jallscalls (Schaus) (Grambidae). Figure 20.30 Male Bellura obllqua (Walker) (Noctuidae). Figure 20.31 Female Bellura obllqua (Walker) (Noctuidae). Figure 20.32 Lateral head of Elophlla with upturned labial palpi (I.p.). Figure 20.33 Lateral head of Acentria with downcurved labial palpi (I.p.).

Chapter 20 Aquatic and Semiaquatic Lepidoptera

111

Figuni 30.34

Figure Ml.35

Ml M2

LI L2

Figure 20.37

V

'V ^•

Figure 20.36

Figure 20.34 Membranous sac anterior to prothoracic coxae in Schoenobiinae (Crambidae). Figure 20.35 Transverse plate posterior to dorsal pinacula on mesothorax in Crambinae (Crambidae).

Figure 20.36 Transverse row of dorsal gills of Oxyelophila callista (Forbes).

Figure 20.37 Modified labral setae of Neargyractis slossonalis (Dyar) modifed from Habeck (1988).

778

Chapter 20 Aquatic and Semiaquatic Lepidoptera

ADDITIONAL TAXONOMIC REFERENCES General taxonomic Muller (1892); Quail (1904), Portier (1911, 1949)Tsou (1914); Fracker (1915); Heinrich (1916); Mosher (1916); Gerasimov (1937); Frohne (1939b); Wesenberg-Lund (1943); Hinton (1946, 1948, 1956); Peterson (1948); Chu(1949, 1956); Patocka (1955); Capps(1956); Lange (1956a,b); Haggett (1955-1961); Speyer (1958); Welch (1959); Okumura (1961); MacKay(1963a, 1964,1972); Klots (1966); Common (1970); Pennak (1978); Lehmkuhi (1979a); Brigham and Flerlong (1982); Scoble (1992); Gra^a and Soils (2018)'.

Regional faunas California: Lange (1956a); Okumura (1961); Powell(1964); Jackson and Resh (1989). Florida: Fleppner (1976); Minno(1992). Indiana and adjacent areas: McCafferty and Minno(1979). New York: Forbes(1923, 1954, 1960).

Cosmopterigidae:(Cosmopteriginae): Walsingham (1907)-A; Hodges(1962)-A; Zimmerman (1978)-A. Nepticulidae: Braun (1917)-L,A; Braun (1925)-L,A. Noctuoidea: Crumb (1929, 1956)-L; Forbes(1954)-L., A.; MacKay and Rockburne (1958)-L; Beck (1960)-L;

Fibiger et al.(2005)-A; Lafontaine and Schmidt (2010)A; Pyraloidea: Forbes (1910)-L, A; Heinrich (1940)-A; Lange (1956a,b)-L, A; Hasenfuss(1960)-L; Munroe

(1972-1973)-A; Heppner (1976)-L, A; Kubik (1981)L,A; Yoshiyasu (1985)-L,A.; Soils et al.(2004)-L,A; Speidel (2005)-A; Boughton and Pemberton (2012)-L, A; Scholtens and Soils (2015)-A; Soils et al.(2018)-L, A; Tortricidae: MacDunnough (1933)-L; Swatchek (1958)-L; MacKay (1926b, 1963b)-L; Powell (1964)-L, A Tortricidae (Olethreutinae): MacKay and Rockburne (1958)-L; MacKay (1959,1962a)-L; Beck (1960)-L.; Gilligan et al. (2008)-L., A.

Taxonomic treatments at the family and generic levels(L = larvae; A = adults) Coleophoridae: Bucheli et al.(2002)-L,A; Cosmopterigidae: Hodges(1962)-A; Hodges(1978)-A; Koster (2010)-A; Rubinoff(2008)-L,A; Schmitz and Rubinoff(2008)-L,A; Rubinoff and Schmitz(2010)-L,A; Schmitz and Rubinoff(201 la,b)-L,A; Haines et ai.(2014)A; Dupont and Rubinoff (2015)-L.

'Graca, M. B., and M. A. Solis. 2018. Order Lepidoptera. Chapt. 11, pp. 325-337. In: Thorp and Covich's Freshwater Invertebrates, Vol. IV: Keys to Neotropical Hexapoda. N. Hamada, J. H. Thorp, and D. C. Rogers (eds). 4th edition. Academic Press, San Diego, CA.

3); table prepared by K, W. Cummins, R. W. Merritt, W. Fl. Lange, and M. A. Solis)

Table 20A Summary of ecological and distributional data for Lepidoptera.(For definition of terms

(Continued)

1519, 1520, 1938, 2308

324, 459, 771, 1929, 3033, 3397, 3900, 4203, 4207, 4359, 6040, 6041, 1201, 1203,2260, 4339

2547, 3065, 3100, 3179, 3397, 3441, 3442, 4207, 6061, 6410

3396

33, 3065, 3179, 3397, 4599, 4789, 4790, 5983, 950, 6061, 6410, 6423,4076, 669, 2310,

References**

Ecological

) J )) ))))))) ) ) J ) ) ) )) ) ))) ) ) )

Family

Langessa

(3)

Eoparargyractls

Elophila

Nymphuta, Syndita)

Elophila (9) (= Munroessa,

nomophllalls

ekthllpsis

leaves or

Lemna)

Brasenla.,etc.)

hydrophytes (Nymphaea, Nympholdes)

Lentic—vascular

Lentic—littoral?

hydrophytes

herbivores

Shredders—

(chewers)

herbivores

Shredders—

Climbers

sedges. Cyperaceae)

herbivores

Shredders—

(case (chewers and constructed of miners)

Climbers— swimmers

Lentic—vascular

pieces of

constructed of

(cases

Climbers— swimmers

Lentic and lotic—

vascular hydrophytes (Potamogeton, Valllsneria, Nuphar, Nymphaea, Ludwigia, Lemna,

Widespread (common in Florida)

Texas

US west to

Canada and

Eastern

Canada

Michigan and

west to

Eastern US

Widespread

US

Southeastern

vlttatalls

Contiger

Habit Eastern US

Habitat

west to Texas

Species

Chrysendeton

Genus

North M

NW MA*

Tolerance Values

Trophic American Relationships Distribution SE UM

(4)

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

parentheses)

(number of species in

Taxa

Table 20A

Ecoiogical

2547, 3397, 4207, 5773

1809, 3397, 4207

297,449, 450, 1928, 1995, 2451, 2961, 3397, 3944, 3945, 4204,4207, 5371, 5479,6423, 6572

3900, 3944, 4207, 5773

1928, 2547, 3397,

3397, 4207

1938, 2756, 3397,4207

References**

) ) ) ) ) ) ) ) ) )) ) ) ) ) ) ) ) ) ) ) ) ) )!)

~4 00 o

00

Family slossonalis

magniflcalis

daeckealis

Neargyractis

Neocatadysta

Nymphuliella

Oligostigmoides cryptalis

Species

Genus

Continued

constructed of

Lemna)

herbivores;

herbivores

fruitans)

Cephalozia

Shredders-

Climbers (case constructed of

(periphyton)

scrapers

facultative

Shredders—

Climbers—

swimmers

herbivores(on Lemna)

Shredders—

Lentic and lotic

Lentic—bog poois

LEPIDOPTERA

North

Texas

Eastern US

US

Southeastern

Florida

Quebec to

and southern

Nova Scotia

Trophic American Relationships Distribution SE

(streams and lakes)

Lemna)

Climbers—

swimmers (case

Lentic and lotic—

Habit

vascular hydrophytes (floating zone:

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

species in parentheses)

(number of

Taxa

Table 20A

NW MA*

Tolerance Values

3397, 4207

(Continued)

2509, 3397, 4207

2306, 3193, 4207

1927, 3397,4207, 6061, 2756, 3900

References**

Ecological

J ) 3 1 ) 3 3 J ) ) )) J 3 )) ) } 3 ) > )3 ) ) )

Climbers—

Habit

Oxyelophila

herbivores

Climbers— swimmers

Lentic—vascular

hydrophytes (Hygrophila, Heteranthera, Ceratophylium, Myriophyllum, Hydriiia)

plant material)

shredders—

makers, shelters of

(algae, diatoms, roots of submerged plants)

Scrapers:

Texas

Widespread

Widespread

Widespread

2.7

M

5.0

5.0

NW MA*

Tolerance Values

'Solis, M. A., and P. Tuskes. 2018. Two new species oi Petrophila Guilding (Lepidoptera: Crambidae)from Arizona, USA. Proc. Ent. Soc. Wash. 120: 593-604. See also: Tuskes, P. M., and A. M. Tuskes. 2019. Aquatic moths of the genus Petrophila and their biology in Oak Creek, Arizona (Crambidae), J. Lep. Soc. 73: 43-53.

tUnpubiished data, W.H. Lange, Department of Entomology, University of California

tlotal number of North American (or Hawaiian for Hyposmocoma)taxa including an unknown number of semiaquatic taxa

herbivores

Shredders-

facultative

retreat

Lotic—erosional;

Clingers (silk-

herbivores

Shredders-

lentic— erosional

Omnium, etc.)

Vallisneria, Nymphoides,

hydrophytes swimmers (Nuphar, Nymphaea, (cases of a Brasenia, wide variety Potamogeton, of aquatic Bacopa, plants) Myriophyllum,

Lentic—vascular

Habitat

Petrophila (17)'

callista

Species

North

Trophic American Relationships Distribution SE

(=Parargyractis)

[sic])

Parapoynx(7) (=Paraponyx

Genus

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

parentheses)

(number of species in

Continued

Family

Table 20A

Taxa

Ecological

3397, 4207, 5569

3397, 3441, 3442, 3443, 3559, 3900, 4207, 1938, 6061, 6067, 6068, 1241, 1519, 1520, 4962, 3851, 6328, 3318, t, 4

2425, 2548, 3065, 3397, 3900, 4207, 6606, 1520, 277, 2307, 3150, 2540, 5773

278,450, 1926,2425, 2547, 3900, 3944, 3945, 5255, 6404, 6407, 6408, 6410, 1520, 277, 772, 2303, 4175, 491, 4175, 5773

References**

) ) ) ) ) ) ) ) ) ) ) ))) ) ) ) ) ))) ) ) } ) ) )

00 to

00

Family

Schoenobiinae

Generally

Generally Generally lentic— vascular hydrophytes burrowers (miners—stem (emergent zone) Donacaula (13)

herbivores

Scirpus)

Eleocharis, Carex,

(miners)

stem borers

below water)

Shredders— herbivores

Burrowers

(miners—

Lentic—^vascular

herbivores

hydrophytes (semiaquatic) (emergent zone;

borers)

Shredders—

Climbers

Lentic—margins(on ferns)

Neomusotima

(1) shredders—

Florida

herbivores

LEPIDOPTERA

Widespread

Florida

Southern

Southern

Shredders—

Texas

California, Arizona,

Climbers

herbivores

Shredders—

Lentic—margins(on ferns)

Eleocharis)

Climbers— swimmers

hydrophytes (Hygrophila,

Habit

Lentic—^vascular

Habitat

North M

NW MA*

Tolerance Values

Trophic American Relationships Distribution SE UM

Undulambia (3)

Species

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships ***Estimated number of aquatic and/or semiaquatic taxa

(25)***

Genus

Usingeriessa (2)

Continued

Musotiminae (4)

species in parentheses)

(number of

Taxa

Table 20A

(Continued)

1995, 1998, 3900, 6403

4207

5568, 621

2547, 4207

3397, 4207, 5569

References**

Ecological

J ))))))) ) ))) 3 )))) ) J )) ) ) ) ) )

Pyraustinae Ostrinia (4)

(=Pyrausta)

penitalis

comptulatalis

hydrophytes (Potamogeton penitalis)(emergent and floating zone; Eupatorium, Poiygonum, and Nymphaceae)

Lentic—vascular

hydrophytes (emergent zone; Scirpus)

Lentic—^vascular

locomotion)

(adapted for aquatic

borers)

and stem

(leaf miners

Borrowers

(miners— stem borers)

Borrowers

Borrowers

(miners— stem borers)

Lentic—vascular

hydrophytes (emergent zone; Scirpus, Juncus, Eleocharis, Otyza)

*SE = Southeast, DM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Err)phasis on trophic relationships ***Estimated number of aquatic and/or semiaquatic taxa

Spilomelinae (30)***

(5)***

i=Adgona)

Occidentalia

Chilo (3)

herbivores

Shredders—

(miners)

herbivores

Shredders—

(miners)

herbivores

Shredders—

(miners)

stem borers

below water)

herbivores

Shredders—

Widespread

United States

Eastern

Widespread

M

NW MA*

Tolerance Values

Trophic American Relationships Distribution SE UM

Borrowers

Habit

(miners—

Habitat Lentic—vascular

Species hydrophytes (emergent zone)

Genus

Crambinae

Family

Continued

(8)***

parentheses)

(number of species in

Taxa

Table 20A

Ecological

33, 2540, 3900, 6406

3205

1998, 3900

1995, 1997, 3900

1998, 3900, 4207

References**

) ) ) ) j V ) ) ) ) ) ) ) ) ) ) ) ) ) ))) ) I )) )

00

00 cn

Noctuinae

(100)***

Noctuidae

Family albiguttalis

multiplicalis

Niphograpta

Samea

Capsula (4) (=Archanara)

Bellura (6) (=Arzarna)

Species

Genus

Continued

Borrowers

(miners— stem borers)

Lentic—vascular

hydrophytes (emergent and floating zones; Typha, Scirpus, Juncus, Sparganium, Phragmites)

Sympiocarpus, Sagittaria, Sparganium)

(later instars) Pontederia, Eichhornia, Nelumbo,

LEPIDOPTERA

(chewers)

herbivores

Shredders—

(miners)

(early Instars); petiole and stem borers

herbivores

Borrowers—

leaf miners

Shredders—

herbivores

Shredders—

herbivores

Shredders—

herbivores

Shredders—

hydrophytes (emergent and floating zones; Typha, Nuphar,

climbers

borrowers and

Generally

North

Widespread

US

and southern

Central, East,

US

Southeastern

US

Southeastern

M

NW MA*

Tolerance Values

American Trophic Relationships Distribution SE UM

Lentic—^vascular

Generally lentic— vascular hydrophytes

of plant

zone, Pistia, Eichhornia, Salvia, Azoila)

material)

Clingers (silk retreat made

Lentic—^vascular

Borrowers

Habit

hydrophytes (floating

zone, Eichhornia)

hydrophytes (floating

Lentic—vascular

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships ***Estimated number of aquatic and/or semiaquatic taxa

Order

parentheses)

(number of species In

Taxa

Table 20A

(Continued)

1231, 1995, 3397, 3900, 6219

450, 1040, 1231, 3511 3900, 3944, 3945, 6193, 2302, 2513, 6401, 6732, 6219

2310, 3639

3205

950, 952, 1394

References**

Ecological

J > 3 ))))) ) ) ) J ))3 )) )))) ) ) ) ) )

Condicinae

Family Genus

Homophoberia (2)

Xylena

Papaipema (2)

Meropleon (6) (=Oligia)

Hemipachnobia (2)

Continued

nupera

diversicolor

Species

(chewers)

(early instars),

hydrophytes (emergent zone; Nuphar, Polygonum)

Lentic—vascular

Midwestern US

(chewers)

Eastern US, herbivores

Shredders—

Midwestern US

(chewers)

etc.)

Northeastern

US,

Shredders—

endemic

locally

(chewers)

Widespread, some spp.

herbivores

herbivores

Climbers

Midwestern US

hydrophytes {Scirpus,

Lentic—vascular Borrowers

hydrophytes (emergent zone; Decodon, Saururus, Sarracenia)

Borrowers—

stem borers

Lentic—vascular

(later instars) Shredders—

herbivores

Borrowers—

leaf miners

Shredders—

locally endemic

(chewers)

Eastern US, some spp.

herbivores

Shredders—

Lentic—vascular

stem borers

North

M

NW MA*

Tolerance Values

Trophic American Relationships Distribution SE UM

hydrophytes (emergent zone; Scirpus)

Dionaea, Drosera)

Borrowers—

climbers

Lentic—vascular

Habit

hydrophytes (emergent zone;

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

parentheses)

(number of species in

Taxa

Table 20A

Ecological

2304, 2540, 3900, 6219

6219

3351, 6219

3900

5369, 3350, 6219

References**

I ) ) ) ; ) ) )) ) ) ) ) > )) ) ) ))) ) ) ))) )

00 On

00

Tortricinae

(20)***

Tortricidae

Acronictinae

Family Genus

Burrowers-

climbers

Lentic—^vascular

hydrophytes (emergent zone; Polygonum)

Platynota (1)

rostrana

Burrowers-

climbers

Lentic—vascular

hydrophytes (emergent zone; Polygonum)

Sparganothis (1)

Burrowers-

climbers

hydrophytes (emergent and floating zones; Typha, Polygonum)

Climbers

Habit

Lentic—^vascular

Salix, Gramlnaceae)

hydrophytes (emergent and floating zones; Typha, Polygonum,

Lentic—vascular

Habitat

Choristoneura

sulfureana

henrld

Species

(2)

Simyra (1)

Continued

LEPIDOPTERA

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships ^Unpublished data, W.H. Lange, Department of Entomology, University of California

Order

parentheses)

(number of species in

Taxa

Table 20A

North

herbivores (leaf rollers)

Shredders—

herbivores (leaf rollers)

Shredders—

herbivores (leaf rollers)

Shredders—

(chewers)

herbivores

Shredders-

U.S.

Midwestern

Eastern,

U.S.

Midwestern

Eastern,

Widespread

Widespread

Trophic American Relationships Distribution SE UM M

NW MA*

Tolerance Values

Ecological

2540

2540

1040, 2540

(Continued)

930, 1231, 6219, +

References**

(180)t

Cosmopterigidae

Coleophoridae (144)t

Olethreutinae

Family

Coleophora (16)

Bactra (2)

Argyrotaenia

Continued

ivana

Generally lentic— vascular hydrophytes (miners)

Generally borrowers

Borrowers—

seed capsules

Lentic—salt marsh;

Juncus, Salicornia, Polygonum

Juncus)

Burrowers-

climbers

Lentic—vascular

hydrophytes (emergent and floating zones; Scirpus, Cyperus,

etc.)

Burrowers-

climbers

hydrophytes (emergent zone; Cyperus, Paspalum,

Habit

Lentic—vascular

Habitat

North

Shredders— herbivores

herbivores

Shredders—

(borers)

herbivores

Shredders—

herbivores (leaf rollers)

Shredders—

Widespread, New England

Widespread

U.S.

Southeastern

M NW MA*

Tolerance va ues

Trophic American Relationships Distribution SE UM

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships tlotai number of North American (or Hawaiian for Hyposmocoma)taxa including an unknown number of semiaquatic taxa

Order

parentheses)

(number of species in

Taxa

Table 20A

Ecological

1040, 1995, 2310

1659, 2540, 765

2122, 2507, 2508, 1434, 3141, 3082, 1983, 1982

2084, 3141

References**

) ) ) ))) ) ) ) ) ) ) ) ) ) )) ) ) ) ) ) ) ) ) )

00

-4 00

90 xo

(=Stigmellidae)

(48)t

Nepticulidae

Family

Lentic —margins(on lichens, algae, and mosses); predator (1 sp.)(unknown number of species semi-aquatic)

scirpi

Shredders—

Climbers (in cases of sand

Lotic—erosional;

Hyposmocoma (>15)***

Acalyptris (=Nepticula)

herbivores

portion of

Borrowers

(miners— stem borers)

hydrophytes (emergent zone; Scirpus, Eleocharis)

basalt rocks)

(miners)

herbivores

Shredders—

scrapers)

emergent

(chewers and

particles, silk,

herbivores

Shredders—

(miners)

and debris on

Lentic—vascular

hydrophytes (emergent and floating zones; Typha)

Borrowers (in heads, seeds, or stems)

Lentic—vascular

phragmitella

Lymnaecia

herbivores

Shredders—

Borrowers

(miners— stem borers)

Habit

Lentic—vascular

Habitat

hydrophytes (emergent zone)

Species

Cosmopterix

Genus

LEPIDOPTERA

North

Widespread

Hawaii

Endemic to

Widespread

Widespread

M NW MA*

Tolerance Values

Trophic American Relationships Distribution SE UM

(5)***

Continued

*SE = Southeast, DIM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships ***Estimated number of aquatic and/or semiaquatic taxa tlotal number of North American (or Hawaiian for Hyposmocoma)taxa including an unknown number of semiaquatic taxa tUnpublished data, W.H. Lange, Department of Entomology, University of California

Order

parentheses)

(number of species In

Taxa

Table 20A

Ecological

1995, 1998, 657

6295, 6855, 5184, 5335, 5185, 5336, 2321, 1555, t

1040

1040

References**

nwi

"< =■

*'r

rj*f.•,**«.ff- . *%'-"-'(i^--:' 't

,'t'tJ :\^ \ J L -•*'%«

• ^y»^.

-

atofltefcw-^"

AQUATIC COLEOPTERA* Andrew E. Z. Short

David S. White

University of Kansas

Hancock Biological Station Murray, Kentucky

Lawrence, Kansas

INTRODUCTION

Coleoptera, in addition to being the most diverse order of insects, ranks as one of the major groups of freshwater arthropods, with more than 13,000 aquatic species (Short 2018). Beetles occupy a broad spectrum of aquatic habitats, ranging from cold mountain streams to brackish waters of estu

aries and salt marshes, to forested vernal pools. Beetles are important in some aquatic food webs, and a number of taxa are consumed by fish and waterfowl. While they may be dominant in some lentic habitats, beetles rarely reach the high popu lation densities or biomass levels in lotic habitats

exhibited by some Ephemeroptera, Trichoptera, and Diptera. The general systematics, biology, and ecology of North American Coleoptera are summarized in the two volume American Beetles (Arnett and Thomas 2001; Arnett et al. 2002), and Bousquet (1991) provides a very useful checklist to the species of Canada and Alaska. Detailed, modem summaries on all water

beetle families can be found in the Handbook ofZoology series (Rolf and Leschen 2016). Hilsenhoff (2001) and Hershey and Lamberti (2001) also provide excellent discussions of the families of aquatic Coleoptera. Epler (2010) and Ciegler (2003) give keys to genera and spe cies of the aquatic Florida and southeastern fauna, respectively, along with extensive summaries of ecol ogy. Hilsenhoff (1996) provides keys to Wisconsin taxa that are useful for much of the Midwest. Intertidal

species are treated by Doyen (1975, 1976) and White and Nelson (2007). Stehr (1991) and the worldwide work by Bertrand (1972) are the most comprehensive available on aquatic immatures. General treatments of aquatic Coleoptera, as well as references to more detailed ecological and taxonomic works, are given at the end of the key and in Table 21 A.

It is difficult to make broad generalizations about the life histories and ecology of aquatic Coleoptera, even at the family level, because the order has invaded aquatic habitats many times and in many ways. Jach (1998) provides a survey of water beetle diversity in the context of habitat and life his tory variability. For a general discussion of many of the adaptations see chapter 2.8 in Wichard et al. (2002). Several families of the suborder Adephaga (e.g., Dytiscidae, Haliplidae, Gyrinidae) are wholly aquatic, leading some to speculate that the hardened elytra and heavy sclerotization may have evolved as a mechanism to keep water out. The suborder Myxophaga (e.g., Hydroscaphidae) is primarily aquatic living in madicolous habitats (thin sheets of flowing water). Only a few of the numerous families in the suborder Polyphaga are wholly aquatic, and adaptations to aquatic existence are quite varied. For example, Heteroceridae exist only at the water's edge; Psephenidae adults are terrestrial, whereas the larvae are aquatic; Dryopidae adults are primarily aquatic and the larvae are primarily terrestrial; and both the adults and larvae of most Flmidae are

aquatic. Therefore, the introductory materials given below are quite broad and general, and more specific information for individual families has been included within the tables and text.

Many aquatic beetles are substrate dwellers. Some notable exceptions, including pelagic larvae of some Dytiscidae, adult Dytiscidae, and Hydrophilidae, are efficient swimmers that must return to the surface periodically to renew their air supply (see Chapter 4). Numerous species, includ ing virtually all marine and littoral forms, inhabit cracks, crevices, or self-constructed burrows and

seldom, if ever, venture into open water. Adults of most aquatic species leave the water temporarily on

* Some of the references cited in this chapter are presented as "Additional Coleoptera References Not Included in Bibliography Above" following reference No. 6908 in the Bibliography (Page 1453)". 791

792

Chapter 21

Aquatic Coleoptera

dispersal flights, which may occur once (Elmidae) or repeatedly(many Dytiscidae and Hydrophilidae). There are a few taxa known only from subterra nean habitats (e.g., Stygoporus, Haideoporus), and undoubtedly more genera and species will be described when other ecosystems (e.g., aquifers, groundwater, hyporheic) are better studied. In Australia,for example,large numbers ofspecies are known only from these subterranean habitats and

many of them were discovered only recently (Leys et al. 2003). Most Adephaga are predators either engulfing their prey or injecting digestive enzymes through piercing mouth parts (especially larvae of Dytiscidae). Some large dytiscids attack small fish or tadpoles. The Myxophaga are scrapers, and hydroscaphids {Hydroscapha) are unusual in that their diet consists largely of bluegreen algae. Feeding habits of aquatic Polyphaga are extremely diverse (Table 21A) and include most of the categories described in Chapter 6. Respiration in aquatic beetles conforms to four major modes(Chapter 4):(1)reliance on self-contained air reserves(e.g.,Dytiscidae,Haliplidae,Hydrophilidae, Hydraenidae); (2) transcuticular respiration, with or without tracheal gills(larvae of most families);(3)plas tron respiration (adult Dryopidae, Elmidae, etc.); and (4) piercing plant tissues (e.g., larval donaciine Chrysomelidae).In most adults having an air reservoir, it occupies the space beneath the elytra, and these bee tles must regularly return to the surface to renew depleted air supplies. Adults using plastron respiration mostly occupy fast-moving, well-oxygenated water and can be quite sensitive to pollutants that act as wetting agents. Type 1 adults must return to the sur face periodically to renew their oxygen supplies. Representatives of types (2), (3), and (4) may remain submerged indefinitely. Adults of most aquatic beetles probably survive a single season or part of a season, but some dytiscids, hydrophilids, and elmids have been maintained for years in aquariums. Species with terrestrial adults (Psephenidae, Scirtidae, Ptilodactylidae) frequently reproduce and die after a short time, sometimes with out feeding. Although Gyrinus species form schools of thou sands of individuals, aquatic beetles generally do not form mating aggregations. Species with short lived adults, however, often emerge synchronously and may be very abundant locally at specific times of the year. Copulation most often occurs with the male mounted dorsally on the female and may be preceded by stroking, stridulation, or other courtship

behaviors. The eggs, numbering from one (Hydroscaphidae) to hundreds (Psephenidae), are deposited singly or in masses in diverse situations. Most species with truly aquatic larvae oviposit underwater, but scirtids apparently oviposit in the damp marginal zone, which is used also by georissids, heterocerids, and other families with sub-

aquatic larvae. Some dytiscids insert the eggs into plant tissue, and a few hydrophilid females carry the egg mass beneath the abdomen. Eggs are generally simple ovoids without a thickened or sculptured chorion. Most are laid naked, but those of hydroph ilids and some hydraenids are enclosed in silken cases. Eggs usually hatch in about 1-2 weeks, but eclosion may be delayed for many months in excep tional cases.

As with most other aquatic insects, at least one stage is terrestrial at some point, which may have limited colonization of the open oceans. The life his tories of aquatic Coleoptera are quite variable. The larvae of some families (e.g., Psephenidae) can be col lected the year round, but the adults are present for only a short period of time during summer. Both adults and larvae of Elmidae can be collected

together most times of the year. In other families (e.g., Hydrophilidae) it is the adult that is most often collected, whereas the larvae exist for only a few weeks in the summer. Most aquatic Coleoptera pass through from three to eight larval instars, requiring an average of6-8 months to develop, with a single generation per year in temperate regions. In almost all species pupation is terrestrial, usually in cells excavated by the larvae under stones, logs, or other objects or occasionally in mud cells on aquatic vegetation (some Gyrinidae). Noteridae, Curculionidae, Chrysomelidae, and possibly some Hydrophilidae spin silk cocoons enclosing a bubble of air, within which they pupate while submerged. Psephenidae pupae are pharate, forming beneath the cuticle of the last larval instar, usually under stones at the water's edge. Just prior to pupation, the last larval instar becomes a quiescent prepupa, when diapause may occur. Few Coleoptera dia pause as pupae and generally transform to adults in about 2-3 weeks.

LITTORAL AND SHORELINE COLEOPTERA

Of all the orders, except perhaps Diptera and Hemiptera, Coleoptera is unique in including many semi-aquatic species that inhabit shoreline environments. Fresh, brackish, and saline waters

all support shoreline faunas, similar in family

Chapter 21 Aquatic Coleoptera

representation but differing in species composition. Aquatic marine Coleoptera are almost entirely members of families that commonly frequent the shoreline and are discussed separately below. In freshwaters, however, families well represented at the shoreline are almost entirely absent from truly aquatic habitats. For example, the large families Staphylinidae, Curculionidae, Chrysomelidae, and Carabidae contain numerous genera and hundreds of species that occur regularly, if not exclusively, around water. Yet only a few could be called aquatic, even in the loosest sense. Some species may submerge temporarily to escape predators but require terrestrial situations to survive and repro duce. Because of the sheer numbers of taxa, we

have not included these and other strictly littoral and intertidal families in the generic level keys. We have, however, included them in the family level keys, in the text, and in listings of genera in

793

littoral aquatic macrophytes harbor a number of Chrysomelidae and Curculionidae, many of which feed on a specific species or genus of plant. Indeed some beetles have been introduced for biological control of invasive plants (e.g., Agasicles on the plant Alternanthera and Galerucella on the plant Lythrum). In their peripheral areas, shoreline habitats merge with purely terrestrial ones. Along steepbanked streams the transition may be abrupt, but along swampy streams and ponds it is almost imperceptible. In addition to the shoreline inhabi tants, many terrestrial taxa are often regular mem bers ofsuch intermediate communities.In particular, members of the terrestrial soil fauna (Pselaphidae, Ptiliidae, Leiodidae, Scydmaenidae, etc.) are likely to occur along with Lampyridae, Cantharidae, and others.

Table 21 A.

The shoreline comprises several distinct habi tats. The interstices in gravel or coarse sand sub strates alongside bodies of water support a fauna of minute beetles such as Sphaeriusidae, Flydraenidae, and Hydrophilidae. Many of these species are actu ally aquatic, living in the water held in interstitial spaces in the substrate. A much larger number of species frequent the surfaces of damp sand or mud adjacent to standing or running water. Adult and larval Carabidae and Staphylinidae are the most obvious and diverse element in this situation, often

very dense on mud or sandbars, especially around drying ponds or intermittent streams. Limnichidae and Georissidae also frequent this zone and may be locally abundant. Adult and larval Heteroceridae

MARINE COLEOPTERA

Although numerous families of Coleoptera inhabit coastal environments, most of these occupy beach or intertidal situations. In particular, the families Staphylinidae and Carabidae are often the dominant coastal Coleoptera in numbers

of individuals and species. Other terrestrial families with coastal representatives include Histeridae, Anthicidae, Melyridae, Ptiliidae, Salpingidae, Tenebrionidae, and Curculionidae. Some of these coastal forms inhabit air-filled crevices in rocks, whereas others live under debris and washed up sea

locations, but some species dig in mud, and a few

weeds(wrack). Many Staphylinidae(and a few exotic Carabidae) inhabit self-constructed burrows in sandy beaches. In most cases these burrows probably entrap air. While there are no open-ocean Coleoptera as far as is known, a number of freshwater aquatic lineages

live on intertidal mud flats. Larval Helichus and

have invaded estuarine and other brackish water

Postelichus(Dryopidae)burrow through moist sand adjacent to streams and most likely are submerged during spring floods. Early larval instars of Acneus (Psephenidae) are apparently aquatic, as with other members of the family. Later instars cling to moist stones just above the waterline and drown if sub merged (G. Ulrich, pers. comm.). Undersides of stones and logs, including those partially sub merged, provide shelter for many shoreline inhabi tants that may emerge nocturnally to forage. Accumulations of water-borne organic debris also shelter many shoreline species. Sampling with Berlese funnels is effective, especially in obtaining minute forms such as Ptiliidae. Emergent and

habitats. Such brackish water taxa are known in the

inhabit burrows they excavate beside streams. Recently emergent sandbars and banks are favorite

Hydrophilidae, Limnichidae, and

Hydraenidae.

Some of these swim openly in brackish waters (e.g., a few species of the hydrophilid genera Berosus,

Tropisternus, and Enochrus), whereas others cling to rocks in intertidal zones (e.g., the hydraenid, genus Neochthebius).

DEFENSE MECHANISMS

Defense mechanisms against predators include behavioral patterns, cryptic coloration, speed, con cealment, holdfasts, hardened bodies, spines and claws, and distasteful chemicals. Most Coleoptera

794

Chapter 21 Aquatic Coleoptera

larvae rely on some form of concealment, residing in cracks and crevices, in debris, or remaining buried in bottom or shoreline sediments, particularly during daylight hours. Larval Psephenidae adhere tightly to rocks and other smooth surfaces and are difficult

to dislodge. Coleoptera and Hemiptera are the only two major orders where most of the adults are both longlived and also aquatic, and both orders appear to rely heavily on chemical defense mechanisms (reviews in

EXTERNAL MORPHOLOGY

Adults

Most adult beetles, including nearly all aquatic members, are characterized by a heavily sclerotized, usually compact body. The combination ofelytra(see below) and antennae with 11 or fewer segments will separate nearly all beetles from other insects with only cursory examination. General features and terminol ogy of beetle structure are illustrated in Figures 21.28,

Dettner 1987; Scrimshaw and Kerfoot 1987; White

21.127, and 21.128.

1989). In the suborder Adephaga, several families have specific glands that produce distasteful and irri tating compounds. Dytiscidae and Hygrobiidae pos sess both pygidial glands that secrete aromatic compounds and prothoracic glands that secrete steroids, giving them a distinctive odor and making them unpalatable. Defensive chemicals in Gyrinidae, Carabidae, Haliplidae, Noteridae, and possibly Amphizoidae are produced primarily in the pygidial glands. Among these aquatic families, the chemistry of gyrinid secretions is best known and consists pri marily of terpenes that are aromatic and extremely distasteful to most predators. Chemical defenses of the aquatic families in the suborder Polyphaga are less well understood, although some Chrysomelidae are known to possess defensive glands and secretions. No glands or chemi cals have been identified, but adult Elmidae appear to be distasteful and are rejected by fish and other pred ators. One South American elmid species apparently is used by people as a spice. Short lived aerial adults, such as limnichids and psephenids, seem to have no chemical defenses but rely on rapid movements and will enter the water when disturbed. Although dytiscids often are distasteful to predators, hydrophilids apparently are not, which seems to have led to mim icry that is particularly common in lentic species. Some adult Hydrophilidae are rejected by fish, but this may be a result of the hard exoskeleton rather

Head and Mouthparts: Coleoptera are mandibulate insects, and the mouthparts are usually visi ble without dissection (Figs. 21.3 and 21.7). Mandibular structure is broadly indicative of feed ing habits. In herbivore scrapers (Dryopidae, Elmidae), the mandible bears a basal flattened molar or grinding lobe as well as a sharp, anterior incisor lobe. In predaceous forms the molar lobe is usually absent. Posteriad of the labium, the head capsule con sists of a gula, delimited by paired gular sutures, except in Curculionidae, where the sutures are coalesced. Configuration of the antennae is used extensively in identification at all levels; important types are illustrated in the keys (Figs. 21.34-21.37, 21.52-21.54). Thorax and Legs: In most beetles the pronotum has encroached ventrally to the region of the procoxae where notum and sternum are separated by a suture (Fig. 21.61). In Dytiscidae and related families the lateral prothoracic region is occupied by the pleuron so that both notopleural and sternopleural sutures are present(Fig. 21.28). Dorsally, much of the thorax and abdomen are concealed by the elytra in most beetles. Morphologically the elytra represent heavily sclero tized forewings without crossveins. Flying wings (hind wings), present in most aquatic beetles, are folded beneath the elytra when the adults are at rest. The number of tarsal segments on each leg is crit ical in identification to family. By convention, these numbers are designated by a three-digit tarsal for mula (e.g., 5-5-5), indicating the number of tarsal segments (tarsomeres) on the anterior, middle, and posterior legs, respectively. The claws are not counted as separate segments. The tibiae and/or tarsi may bear fringes of long, slender swimming hairs (Fig. 21.135), which adhere to the leg in dried specimens and may not be visible without wetting. Abdomen: Normally, only the abdominal sternites are visible. In Dytiscidae and related families the posterior coxae are greatly enlarged, dividing

than chemicals.

In addition to chemical defenses,adult Dytiscidae, Haliplidae, Gyrinidae, and Hydrophilidae are fast swimmers,whereas terrestrial Staphylinidae,Carabidae, and Chrysomelidae are swift fliers and runners. Adult Amphizoidae, some Carabidae, Noteridae, Dryopidae, Elmidae, Sphaeriusidae, Melyridae, Salpingidae, Hydraenidae, and Curculionidae live concealed within crevices, sediments, or vegetation. Further, most adult Coleoptera are hard bodied with sharp claws and many have legs with stiff spines.

Chapter 21 Aquatic Coleoptera

the basal sternite into two separate sclerites (Figs. 21.28, 21.128, 21.134). In counting the number of abdominal sternites (Fig. 21.128), those seg ments associated with the genitalia and normally

elongate and filamentous (Figs. 21.125 and 21.267), also may be present and presumably are homologous to the cerci of many other insects.

drawn into the abdomen should not be tallied. They

KEYS TO THE FAMILIES OF AQUATIC

almost always differ markedly in sculpturing, color, and degree of sclerotization from the external

COLEOPTERA

sternites.

Larvae

Larval Coleoptera are exceedingly diverse. Among aquatic forms, the presence of a distinct, sclerotized head capsule with mandibles, maxillae, labium, and two- or three-segmented antennae

(except in Scirtidae that have long, multisegmented antennae) is a reliable distinguishing feature. All but curculionids(Fig. 21.17) and a few hydrophilids have three pairs of thoracic legs bearing one or two apical claws. Beetle pupae are exarate (appendages not fused to body) and bear a general similarity to adults.

Cranial Structures: Mouthparts are extremely important in larval classification and identification, and careful dissection may be necessary to view crit ical structures. As in adults, the mandibles may pos sess or lack a molar lobe. In some larvae a fleshy cushion-like or digitate prostheca is inserted on the mandible just anterior to the molar lobe (Fig. 21.8B). The maxilla may bear a separate galea and lacinia or these may be united as a single lobe, the mala. The maxillary palp is inserted on the palpifer, a projection of the stipes, which is greatly enlarged in Hydrophilidae, appearing as a separate segment of the palp (Fig. 21.220). Eyes often occur in clus ters of 5 or 6 and in larvae are termed stemmata

(singular, stemma). Abdominal Structures: A variety of gill-like appendages may be present laterally or ventrally on any segment (Figs. 21.79, 21.229, 21.314). "Gills" may be simple or branched, dispersed or clustered, unsegmented or articulated. In some families the gills are concealed in a pocket beneath the terminal abdom inal sternite (Figs. 21.322 and 21.363), which forms a lid or operculum, and dissection is frequently neces sary for examination of the gills. Diverse unsegmented,immovable dorsal, or dorsolateral appendages occur on the apical or preapical abdominal segment (Figs. 21.21 and 21.23). These terminal appendages are usually relatively short and strongly sclerotized. They are termed urogomphi(sin gular, urogomphus) regardless of position or origin. Movable urogomphi,sometimes segmented and often

795

Aquatic beetles are not a natural group but represent multiple independent invasions of aquatic habitats within the suborders Adephaga,Myxophaga, and Polyphaga. The following keys are designed primarily for identification of beetles that exclusively or primarily occur in aquatic habitats. Coleoptera that regularly inhabit shorelines including marine forms, although not strictly aquatic, are included at the family level because they frequently appear in aquatic samples. Many members of families that are not aquatic may occasionally appear around water, and those that have merely fallen into the water or have been caught in floods cannot be identified with the keys. If one is unsure whether a specimen is aquatic or not, it is wise to consult some of the gen eral references listed above in the Introduction.

With minor exceptions, there are no species level keys to aquatic Coleoptera larvae, and rearing (see Chapter 3) often is necessary for species level identifications.

Taxonomic usage of family names follows Bouchard et al. (2011), with the exception of the Hydrophiloidea, which we treat as six separate fam ilies as in the prevailing usage (Hansen 1991, 1999; Short 2018). Keys to genera are not provided for the families that are primarily shoreline and/or intertidal. Many of these families are large with only a few marginally aquatic taxa, and it would be difficult to separate the aquatics from the terrestrials that could occur accidentally near water. They include the Carabidae, Lampyridae, Staphylinidae, Ptiliidae, Salpingidae, Anthicidae, Melyridae, Salpingidae, Tenebrionidae, Flisteridae, Chrysomelidae, and Curculionidae. References to appropriate generic keys are given in the section on Selected Additional Taxonomic References. Most all Heteroceridae do

occur in mud and sand along streams and lakes, and genera are best identified with Katovich (2002). All of the above families are included in the text, and all except Heteroceridae, Lampyridae, and Anthicidae are included in Table 21A. Table 21A details those families that are of numerical or

economic importance or that are intertidal. The annotations are followed by a simple listing of

other genera that are primarily limited to aquatic margins.

796

Chapter 21

Aquatic Coleoptera

1.

Mesothoracic wings (elytra) present, usually covering entire abdomen (Fig. 21.41), sometimes only its base (Fig. 21.40) or elytra absent (e.g., Thinopinus—Staphylinidae); antennae with at least 4 segments, usually 6 or more (Figs. 21.37 and 21.51); tarsus with

r.

Mesothoracic wings absent(Figs. 21.1-21.2); antennae with 3 or fewer segments'"(Fig. 21.3);

at least 3 segments (Figs. 21.30 and 21.48)"

ADULTS

tarsus with a single segment(Figs. 21.4 and 21.12)

LARVAE

Larvae

1.

Legs absent(Fig. 21.17)

9

T.

Legs sometimes small, but always with 3-6 clearly defined segments(Figs. 21.12, 21.88,21.90,21.119,21.234)

2(1')

2

Legs (excluding claws) with 5 segments; tarsi with 2 claws (Fig. 21.4)(exception: HALIPLIDAE with single claw. Figs. 21.88 and 21.90)

3

2'

Legs with 3-4 segments (Figs. 21.12 and 21.234); tarsi with single claw (Figs. 21.12 and 21.234)

9

3(2)

Abdomen with 2 pairs of stout, terminal hooks on segment 10(Fig. 21.80); abdominal segments 1-9 bearing lateral gills (Fig. 21.79)

3'

GYRINIDAE(p. 809)

Abdomen without hooks on terminal segment(Fig. 21.89); abdominal segments usually without lateral gills, occasionally with ventral gills (Fig. 21.314)

4

4(3')

Abdomen with 8 segments (Fig. 21.101)

6

4'

Abdomen with 9 or 10 segments(Figs. 21.1, 21.2, 21.10)

5

5(4')

Tarsus with single claw (Figs. 21.88 and 21.90); mandibles grooved internally (Fig. 21.92); at least last larval instar with erect, dorsal projections from thoracic and abdominal tergites (Figs. 21.87 and 21.89) HALIPLIDAE (p. 812)

5'

Tarsi with 2 claws(as in Fig. 21.4); mandibles not grooved; tergites

without projections(Fig. 21.1)

CARABIDAE'*^'^'''

6(4)

Urogomphi slender, longer (usually much longer) than 1st abdominal segment (Fig. 21.125) DYTISCIDAE(in part)(p. 815)

6'

Urogomphi stout, shorter than 1st abdominal segment(Fig. 21.2) or rudimentary or absent

7

7(6')

Thorax and abdomen strongly flattened; tergites expanded laterally as thin, flat projections (Fig. 21.2); gular suture single (Fig. 21.3) AMPHIZOIDAE Amphizoa (p. 809)

T

Thorax and abdomen round or subcylindrical in cross section; tergites not expanded as flat, plate-like projections; gular suture double (Fig. 21.14)

8(7')

Legs short, stout, adapted for digging (Fig. 21.201); mandibles with enlarged molar portion (Fig. 21.202) NOTERIDAE(p. 833)

8'

Legs long, slender, adapted for swimming (Fig. 21.101); mandibles falcate (sickle-shaped), without enlarged molar portion (Fig. 21.120) .... DYTISCIDAE (in part)(p. 815)

9(1,2') 9'

Labrum separated from clypeus by distinct suture (Fig. 21.13) Labrum not represented as separate sclerite (Figs. 21.22, 21.218, 21.222, 21.225) (the ventral labium may be visible dorsally. Fig. 21.228)

10

10(9')

Body round or subcylindrical in cross section; head projecting anteriorly from prothorax and visible from above (Fig. 21.229); movable urogomphi often visible (Fig. 21.267)

11

® Lepiceridae, not occurring in United States, have a single tarsomere. '' Larvae of Scirtidae have numerous, filiform antennal segments. 'Riparian or inhabiting littoral region. ^ Includes intertidal species in North America. ^ Larvae of Brachinus spp.(Carabidae), which are ectoparasitic on pupae of Hydrophilidae, have 3 leg segments and a single claw. " Keys to genera not given.

8

16

Chapter 21

Aquatic Coleoptera

797

labial palp maxillary palp antenna

0|

gular — suture

tarsus

claws

Figure 21.4 Figure 21.3 pronotum lateral

projection

cercus

head

Figure 21.2

Figure 21.5

Figure 21.6

Figure 21.1 maxillary pulp galea stipes

mandible

palpifer

Figure 21.11 Figure 21.13

cardo

gula Figure 21.7

claw

Figure 21.8

Figure 21.12

Figure 21.9

Figure 21.1

Chlaenius sp.(Carabidae) larva, dorsal

Figure 21.14

Figure 21.10

Figure 21.6 Lampyridae larva, dorsal aspect of head

Figure 21.8 Oxytelus sp. (left) and Piestus sp. (right) larvae (Staphylinidae), mandible. Figure 21.9 Thinopinus sp.(Staphylinidae) larva, dorsal aspect. Figure 21.10 Georissus sp.(Georissidae) larva, dorsal aspect (after Van Emden 1956). Figure 21.11 Georissus sp.(Georissidae) larva, abdominal apex. Figure 21.12 Georissus sp.(Georissidae) larva, leg (after Van Emden 1956). Figure 21.13 Idealized dorsal aspect of a coleopteran

and thorax.

larva head.

Figure 21.7 Piestus sp. (Staphylinidae) larva, ventral aspect of head.

larva head.

aspect.

Figure 21.2 Amphizoa sp.(Amphizoidae) larva, dorsal aspect.

Figure 21.3 Amphizoa sp.(Amphizoidae) larva, ventral aspect of head. Figure 21.4 Amphizoa sp.(Amphizoidae) larva, metathoracic leg. Figure 21.5 Lampyrldae larva, lateral aspect of head and thorax.

Figure 21.14 Idealized ventral aspect of a coleopteran

798

10'

Chapter 21

Aquatic Coleoptera

Body dorsoventrally flattened, with large, transverse thoracic and abdominal tergites; pronotum expanded anteriorly, usually concealing head from above

(Figs. 21.5-21.6) 11(10)

11'

Maxilla with palpifer appearing as a segment of palpus (Figs. 21.220 and 21.231); spiracles biforous(having 2 openings) (Fig. 21.209)

12

Maxilla with palpifer appearing as part of stipes (Fig. 21.7); spiracles annular (ring-shaped); various marginal habitats

including intertidal 12(11)

LAMPYRIDAE'-^ (p.

STAPHYLINlDAE''^''*(in part)(p. 847)

Abdomen with 9 complete segments, segment 10 terminal, small but distinct no spiracular atrium or cavity present

13

12'

Abdomen with 8 complete segments, segments 9 and 10 reduced and modified into a spiracular atrium or cavity (atrium absent in Berosus)

15

13(12)

Legs short, 3-segmented (Fig. 21.12)

13'

Legs long, 5-segmented (Figs. 21.210, 21.218, 21.229, 21.235)

14(13')

Urogomphi long, 3-segmented; abdominal tergites and sternites present; integument noticeably chitinized HELOPHORIDAE Helophoms(p. 845)

14'

Urogomphi short, 1-segmented, abdominal tergites and sternites absent; segments 8 and 9 with pairs of fleshy projections (Fig. 21.208) EPIMETOPIDAE Epimetopus

15(12')

Antennae with points of insertion nearer anterolateral angles of head than are insertion points of mandibles; labium and

GEORISSIDAE' Georissus(p. 845) 14

maxillae inserted in furrow beneath head; lacinia

present but small

HYDROCHIDAE Hydrochus(p. 845)

15'

Antennae with points of insertion further from anterolateral angles than those of mandibles; labium and maxillae inserted at anterior margin of ventral side of head; lacinia absent HYDROPHILIDAE(p. 836)

16(9)

Thorax and abdomen short, obese, without distinct sclerites (Figs. 21.15 and 21.17); legs reduced (Fig. 21.15) or absent (Fig. 21.17)

16'

17

Thorax and abdomen cylindrical, flattened, or fusiform (spindle-shaped)(Fig. 21.18), but not markedly obese; thoracic and

abdominal tergites clearly defined (Fig. 21.18); legs adapted for walking 17(16)

18

Legs very small but complete and visible (Fig. 21.15); spiracles on 8th abdominal segment forming large, sclerotized dorsal hooks

(Fig. 21.16)

CHRYSOMELIDAE''"(p. 868)

17'

Legs entirely absent(Fig. 21.17); spiracles sometimes set on tubercles, but 8th segment never with sclerotized

18(16')

At least 8th abdominal tergite bearing pairs of fleshy, articulated, finger-like lobes(Fig. 21.18); antennae very short, 2-segmented (Fig. 21.19); minute larvae, less than 2 mm long

dorsal hooks

'Riparian or inhabiting littoral region. ^ Includes intertidal species in North America. Keys to genera not given.

CURCULIONIDAE"'" (p. 869)

19

Chapter 21

18' 19(18)

Aquatic Coleoptera

Abdominal tergites without dorsal lobes; antennae 3-segmented or more (Fig. 21.26)

799

20

Finger-like articulated lobes present on abdominal segments 1-8 ; antenna with 2nd segment about 2-3 times as long as broad and bearing minute, lateral appendage

(Fig. 21.19)

SPHAERIUSIDAE' ApW/Hs(p. 836)

19'

Finger-like lobes present only on abdominal segments 1 and 8 (Fig. 21.18)

20(18')

Abdomen with 10 segments; 9th segment bearing articulated, 1- or 2jointed urogomphi(Fig. 21.267)

21

Abdomen with 9 segments; 8th or 9th segment sometimes bearing immovable urogomphi(Figs. 21.21 and 21.23), but articulated urogomphi never present

23

21(20)

Mandibles with large, asperate (roughened) molar lobe

22

21'

Mandibles falcate (sickle-shaped), without molar lobe (Fig. 21.8) various marginal habitats

22(21)

Tenth abdominal segment with pair of recurved ventral hooks; urogomphi with 2 segments

20'

including intertidal

HYDROSCAPHIDAE Hydroscapha(p. 835)

STAPHYLINIDAE''^'''(in part)(p. 847)

(Fig. 21.267) 22'

HYDRAENIDAE' (p. 845)

Tenth abdominal segment without hooks; urogomphi with single segment

(as in Fig. 21.9) 23(20')

PTILIIDAE'--"(p. 845)

Antennae much longer than head (Fig. 21.284), multiarticulate

(many jointed)

SCIRTIDAE''(p. 849)

23'

Antennae short

24(23') 24'

Body cylindrical, subcylindrical, or fusiform; head and legs visible in dorsal aspect 25 Body extremely flattened, with thoracic and abdominal tergites expanded laterally as thin laminae concealing head and legs from above (Figs. 21.296-21.299, 21.301, 21.302) PSEPHENIDAE(p. 849) Ninth abdominal segment with a lid-like operculum covering the anal region ventrally (Figs. 21.322 and 21.363); abdominal sternites 1-8 never bearing gills 26

25(24)

25'

26(25)

24

Ninth abdominal segment without operculum; abdominal sternites 1-8 sometimes bearing fasciculate (clustered) gills (Fig. 21.314)

29

Terminal abdominal segment rounded posteriorly (Fig. 21.322); head capsule with groups of6 stemmata (ocelli), 5 lateral and 1 ventral, or stemmata (eyes) absent

26'

27(26)

Terminal abdominal segment bifid or slightly emarginate(notched) posteriorly and with lateral ridges (Fig. 21.332); head capsule with groups of 5 lateral stemmata Abdominal segments I-YII membranous ventrally; mandibles with

prostheca 27' 28(27')

27

ELMIDAE(p. 856)

LIMNICHIDAE"(p. 853)

Abdominal segments 1-VIl sclerotized ventrally; mandibles with or without prostheca 28 Opercular chamber containing 2 retractile hooks and 3 tufts of retractile gills (Fig. 21.262); mandibles with prostheca (Fig. 21.335) LUTROCHIDAE(p. 853)

'Riparian or inhabiting littoral region. ^ Includes intertidal species in North America. Keys to genera not given.

800

28'

Chapter 21

Aquatic Coleoptera

Opercular chamber without hooks or gills; mandibles without prostheca (Fig. 21.323)

DRYOPIDAE(p. 855)

29(25')

Abdomen without gills

30

29'

Abdomen with distinct tufts of gills, either restricted to anal region (as in Fig. 21.315) or present on segments 1-7 (Fig. 21.314)

34

Abdomen bearing prominent, spine-like urogomphi on terminal segment (Figs. 21.21 and 21.23)

31

30(29) 30' 31(30)

Urogomphi absent; 9th abdominal segment rounded Each urogomphus with two points (Fig. 21.21); spiracles raised on tubercles(Fig. 21.20);

marine intertidal

SALPINGIDAE^"''(p. 868)

31'

Each urogomphus with a single point(Fig. 21.23); spiracles not elevated

32(31') 32'

Epieranial suture lyre-shaped (Fig. 21.22) Epieranial suture Y-shaped (Fig. 21.24)

33(30')

Mouthparts prognathous; median epieranial suture absent

(Fig. 21.25)

33

32

ANTHICIDAE'-"(p. 868) MELYRIDAE'-^'' HETEROCERIDAE'-''(p. 853)

33'

Mouthparts hypognathous; median epieranial suture present (Fig. 21.26)

34(29')

Abdominal segments 1-7 each with 2 ventral tufts of filamentous gills (Fig. 21.314); submentum not divided; 9th abdominal segment without prehensile appendages bearing hooks EULICHADIDAE Stenocolus (p. 853)

34'

Abdominal segments 1-7 without gill tufts; submentum divided longitudinally into 3 parts; anal region of 9th abdominal segment with 2 curved prehensile appendages covered with short spines (Fig. 21.315) PTILODACTYLIDAE (p. 853)

TENEBRIONIDAE

(p. 868)

Adults

To aid in identification to the family level, we provide a separate set offigures here, some of which are duplicated in the generic level keys. Flabitus drawings are provided for most families for comparison, and the additional figures given at the end of each couplet may be used to confirm identifications. 1 Compound eyes divided into separate dorsal and ventral segments (Fig. 21.27); second antennal segment elongate and scoop-shaped (Fig. 21.83); (Figs. 21.27, 21.81, 21.82) GYRINIDAE (p. 809) 1' Compound eyes undivided (Fig. 21.44); antennae variable but without scoop-shaped second antennal segment(Fig. 21.29) 2 2(1') Head produced anteriorly as a rostrum (Fig. 21.30); tarsal formula 4-4-4 CURCULIONIDAE^(p. 869) 2' Head not produced as a rostrum (Fig. 21.39); tarsal formula variable 3 3(2') Elytra covering entire abdomen or exposing only part of last abdominal tergite (Fig. 21.41) 6 3' Elytra truncate, exposing at least 2 entire abdominal tergites (Figs. 21.31, 21.32, 21.33, 21.30) 4 4(3') Antennae with 11 (rarely 10) segments, terminal segment no longer than combined length of 2 preeeding segments (Figs. 21.29, 21.34, 21.35, 21.36)

'Riparian or inhabiting littoral region. ^ Includes intertidal species in North America. Keys to genera not given.

5

Chapter 21

Aquatic Coleoptera

801

appendage

Figure 21.15

hook

Figure 21.19

Figure 21.16

Figure 21.17

epioranial suture

bifid urogomphus

Figure 21.22

urogomphus Figure 21.23

Figure 21.21 Figure 21.20

fleshy lobes

Figure 21.18 Figure 21.24

Figure 21.25

_.

..

Figure 21.26

median epioranial

suture

Figure 21.15 Donacia sp. (Chrysomelidae) larva,

Figure 21.22 Anthicus sp.(Anthicidae) larva, dorsal

lateral aspect.

aspect of cranium.

Figure 21.16 Donacia sp.(Chrysomelidae) larva, ventral aspect of terminal abdominal segment. Figure 21.17 Gurculionidae larva, lateral aspect. Figure 21.18 Hydroscapha sp.(Hydroscaphidae) larva, dorsal aspect. Figure 21.19 Sphaerius sp.(Sphaeriusidae) larva,

Figure 21.23 Endeodes sp.(Melyridae) larva, abdominal apex.

Figure 21.24 Collops sp.(Melyridae) larva, dorsal aspect of cranium. Figure 21.25 Heteroceridae larva, dorsal aspect of cranium.

antenna.

Figure 21.26 Tenebrionidae larva, dorsal aspect of

Figure 21.20 Aegiaiites sp. (Saipingidae) larva, dorsal

cranium.

aspect.

Figure 21.21 Aegiaiites sp.(Saipingidae) larva. abdominal apex.

802

Chapter 21

Aquatic Coleoptera

sternopieural

notopleural suture

metasternum

Figure 21.27

abdominal sternite 1

coxa! plate

trochanter

metafemur

r Figure 21.28 elytron

Figure 21.30 Figure 21.38 protrusible vesicle

elytron terminal

segment

Figure 21.32

Figure 21.31

Figure 21.36 Figure 21.29

Figure 21.37

Figure 21.34 F'Sure 21.35

plate

Figure 21.39

Figure 21.33

Figure 21.41

Figure 21.40

Figure 21.27 Gyretes sp.(Gyrinidae) adult, lateral

Figure 21.35 Anchycteis sp. (Ptilodactylldae) adult,

aspect.

male and female antennae.

Figure 21.28 Laccophilus sp. (Dytiscldae) adult,

Figure 21.36 Optioservus sp.(Elmidae) adult,

ventral aspect.

antenna.

Figure 21.29 Cyphon sp. (Sclrtldae) adult, antenna. Figure 21.30 Tanysphyrus sp.(Gurculionidae) adult, dorsolateral aspect. Figure 21.31 Hydroscapha sp.(Hydroscaphidae) adult, dorsal aspect. Figure 21.32 Melyridae adult, dorsal aspect. Figure 21.33 Thinopinus sp.(Staphylinidae) adult, dorsal aspect. Figure 21.34 Eubrianax sp.(Psephenidae) adult, antenna.

Figure 21.37 Hydroscapha sp.(Hydroscaphidae) adult, antenna.

Figure 21.38 Hydroscapha sp.(Hydroscaphidae) adult, ventral aspect of abdomen and metathorax. Figure 21.39 Hydroscapha sp.(Hydroscaphidae) adult, lateral aspect. Figure 21.40 Endeodes sp.(Melyridae) adult, dorsal aspect.

Figure 21.41 Omophron sp.(Carabidae: Omophroninae) dorsal aspect.

Chapter 21 Aquatic Coleoptera

803

4'

Antennae with 8 segments, terminal segment as long as combined length of4 preceding segments(Fig. 21.37); minute beetles, less than 1.5 mm long; hind coxal plates widely separated; Figs. 21.31 and 21.39 HYDROSCAPHIDAE Hydroscapha (p. 835)

5(4)

Abdomen and thorax with membranous yellow or orange protrusible (extendible) vesicles

5'

Protrusible vesicles absent; Fig. 21.33; various marginal habitats including intertidal;

(Fig. 21.32; most apparent in living beetles); Fig. 21.40; intertidal Figs. 21.278-21.28 6(3) 6' 7(6)

MELYRIDAE'--''^ STAPHYLINIDAE'-^'^'(p. 847)

Hind coxae expanded as broad, flattened plates covering all or part of abdominal sternites 1-3 (Fig. 21.42)

7

Hind coxae sometimes extending posteriorly along midline (Figs. 21.28 and 21.43), but never as broad plates 8 Hind coxal plates completely covering 2 or 3 basal abdominal segments (Fig. 21.42) and concealing all but apices of hind femora; beetles more than 2 mm long; Fig. 21.44.... HALIPLIDAE (p. 812)

7'

Hind coxal plates exposing abdominal segments laterally; bases of hind femora exposed; highly convex beetles less than 1.5 mm long; Fig. 21.45 SPHAERIUSIDAE Sphaerius(p. 836)

8(6')

Hind coxae with medial portion extending posteriorly to divide 1 st abdominal sternite into lateral sclerites (Figs. 21.28 and 21.43); prothorax with distinct notopleural sutures 9

8'

Hind coxae not extending posteriorly to divide 1 st abdominal sternite (Figs. 21.64 and 21.65); notopleural sutures almost always absent Hind tarsi and usually tibiae flattened, streamlined, and bearing long, stiff swimming bristles (Fig. 21.28) Hind tibiae and tarsi cylindrical or subcylindrical in cross section, without long, stiff swimming bristles (as in Fig. 21.33)

9(8) 9'

13 11

10

10(9')

Hind coxae extending laterally to margins of elytra; elytra at most with very short hairs; Fig. 21.46 AMPHIZOIDAE Amphizoa (p. 809)

10'

Hind coxae not extending laterally as far as elytra (Fig. 21.43); elytra bearing several long,

slender, erect sensory hairs 11(9)

CARABIDAE'*^*''

Fore and middle tarsi with 5 segments, segment 4 similar in size to segment 3(Fig. 21.47); scutellum concealed (Figs. 21.132 and 21.156) or exposed (Fig. 21.49)

12

1r

Fore and middle tarsi with 4 segments(Fig. 21.156) or with segment 4 very small, concealed between lobes of segment 3(Fig. 21.48); Fig. 21.49; scutellum concealed except in Celina DYTISCIDAE (in part)(p. 815)

12(11)

Hind tarsi with 2 similar claws (Fig. 21.204); scutellum concealed (Fig. 21.50) NOTERIDAE(p. 833)

12'

Hind tarsi with single claw (Fig. 21.135); if 2 claws, scutellum exposed (Fig. 21.49) DYTISCIDAE(in part)(p. 815)

13(8')

Antennae with terminal segment as long as combined length of 3^ preceding segments (Fig. 21.37)

13' 14(13')

Antennae with terminal segment no longer than combined length of 2 preceding segments (Fig. 21.36); terminal segments may be fused into a globular or elongate club Antennae terminating in abrupt, globular (Figs. 21.51 and 21.52) or elongate (Fig. 21.53)club

'Riparian or inhabiting littoral region. ^ Includes intertidal species in North America. Keys to genera not given.

15

14 15

804

14'

Chapter 21 Aquatic Coleoptera

Antennae slender, elongate (Fig. 21.35) or very short, thick, with basal segment enlarged (Fig. 21.54)

22

15(13,14)Antennae elbowed, with 2nd segment attached medially on elongate 1st segment; antennal club consisting of several compactly fused segments; flat, shiny black, intertidal HISTERIDAE'-^ Neopachylophus(p. 845) 15' Antennae with 2nd segment attached apically on 1st (Fig. 21.54); antennal club consisting of 2-5 articulated segments (Fig. 21.53) 16 16(15') Antennae about as long as head (Fig. 21.45); club with 3-5 segments (Fig. 21.54); tarsal formula 4-4-4, 5-5-5, or 5-4-4 (rarely 5-4-4 or 4-5-5) 17 16' Antennae much longer than head, length approaching half that of body (Fig. 21.55); club with 2-3 loosely articulated segments (Fig. 21.56); tarsal formula 3-3-3;

Fig. 21.55 17(16) 17'

PTILIIDAEiA''(in part)(p. 845)

Abdomen with 5-6 visible sternites; antennal club with 3 segments Abdomen with 6-7 visible sternites; antennal club with 5 segments (Fig. 21.53);

Fig. 21.57

18

HYDRAENIDAE^ (p. 845)

18(17)

Pronotum projected forward, totally or partly covering head (Figs. 21.59 and 21.244); first tarsomere very small (tarsi pseudotetramerous)

19

18'

Pronotum not or only slightly projected forward, not or slightly covering head; tarsi composed of five tarsomeres

20

19(18)

Pronotum lateral margins strongly expanded and bearing longitudinal carinae on disc; elytra with longitudinal carinae and two longitudinal rows of punctures between two contiguous carinae; compound eyes completely or partially divided; total body length up to 5 mm. Fig. 21.244 EPIMETOPIDAE Epimetopus(p. 844)

19'

Pronotum lateral margins not strongly expanded, pronotal disc smooth; elytra lacking carinae, with rows of coarse punctures (Fig. 21.59); compound eyes entire; total body length < 2 mm;dorsum of body sometimes encrusted with sand particles (Fig. 21.60) GEORISSIDAE Georissus(p. 845)

20(18') 20'

Pronotum lacking strongly impressed longitudinal grooves 21 Pronotum bearing five well-developed longitudinal grooves (Fig. 21.243) HELOPHORIDAE Helophoms (p. 845)

21(20)

Pronotum distinctly narrower than elytral base; overall body form narrow and elongate (Fig. 21.245); antennae before club with four glabrous antennomeres; meso- and metatibiae lacking natatory setae; elytra with strongly impressed sculpture, often encrusted with dirt; total body length < 5 mm HYDROCHIDAE Hydrochus (p. 845)

21'

Pronotum not distinctly narrower than elytral base; antennae before club with five or six glabrous antennomeres, if only four glabrous antennomeres before club, meso- and metatibiae bearing long natatory setae; elytra generally smooth or with moderate sculpture; size variable HYDROPHILIDAE(p. 836)

22(14')

Tarsal formula 5-5-5

23

22'

Tarsal formula 5-5-4 or less

34

23(22)

Abdomen with 5-7 visible sternites

23'

Abdomen with at least 8 visible sternites

24(23)

Prosternum expanded anteriorly as prominent lobe beneath head, head usually contracted into thorax concealing antennae and eyes (Figs. 21.62)

25

Prosternum not markedly expanded anteriorly beneath head, antennae clearly visible (Fig. 21.63)

29

Antennae usually thick, with enlarged basal segment(Figs. 21.54 and 21.62), about as long as head

26

24' 25(24)

'Riparian or inhabiting littoral region. ^ Includes intertidal species in North America. ^ Keys to genera not given.

24

LAMPYRIDAE'-"(p. 868)

Chapter 21 Aquatic Coleoptera

805

epipleuron coxa!

metasternum

plate metacoxa -

abdominal sternite 1 trochanter

Figure 21.44

Figure 21.43 Figure 21.42

3rd

segment

4th

segment

Figure 21.45

Figure 21.47

Figure 21.46

Figure 21.48

I Figure 21.55 Figure 21.50

} club

Figure 21.57 Figure 21.49

globular

-club

club

2nd

segment basal

segment

Figure 21.51

Figure 21.54

Figure 21.52

Figure 21.53

Figure 21.56

Figure 21.42 Haliplus sp. {Haliplidae) adult, dorsal

Figure 21.50 Suphis sp.(Noteridae) adult, dorsal

aspect.

aspect.

Figure 21.43 Harpalus sp.(Carabidae) adult, ventral aspect of thoracic region (after Larson 1975). Figure 21.44 Peltodytes sp. (Haliplidae) adult, lateral

antenna.

aspect.

antenna.

Figure 21.45 Sphaerius sp. (Sphaeriusidae) adult, dorsal aspect Figure 21.46 Amphizoa sp.(Amphizoidae) adult. dorsal aspect. Figure 21.47 Coptotomus sp.(Dytiscidae) adult, tarsus of foreleg. Figure 21.48 Hydroporus sp.(Dytiscidae) adult, tarsus of foreleg. Figure 21.49 Cybister sp.(Dytiscidae) adult, dorsal aspect.

Figure 21.51

Neopachylophus sp.(Histeridae) adult,

Figure 21.52 Emphyastes sp.(Curculionidae) adult, Figure 21.53 Hydraena sp.(Hydraenidae) adult, antenna.

Figure 21.54 Helichus sp.(Dryopidae) adult, antenna. Figure 21.55 Actidium sp. (Ptiliidae) adult, dorsal aspect.

Figure 21.56 Actidium sp. (Ptiliidae) adult, antenna. Figure 21.57 Ochthebius sp.(Hydraenidae) adult. dorsal aspect.

806

Chapter 21

hind coxa

Aquatic Coleoptera

Intercoxai process

elytron

Figure 21.58

gular sutures

Figure 21.60

Figure 21.59

sternopleural

mandible

suture'

antenna

prosternum pronotu

elytron

-trochantin

mesopleuron mesocoxa

Intercoxai process mesosternum

elytron

Figure 21.62 Figure 21.61

Figure 21.63

Figure 21.66

Figure 21.65

Figure 21.67

Figure 21.64 Figure 21.68

Figure 21.69

Figure 21.58 Geor/ssus sp.(Georissidae) adult, ventral aspect of abdomen and metathorax. Figure 21.59 Geor/ssus sp.(Georissidae) adult, dorsal

Figure 21.64 Pelonomus sp.(Dryopidae) adult, ventral aspect. Figure 21.65 Postelichus sp.(Dryopidae) adult, lateral

aspect.

aspect.

Figure 21.60 Geor/ssus sp.(Georissidae) adult, dorsal aspect with camouflage of sand grains. Figure 21.61 Hyc/roc/iara sp.(Hydrophilidae) adult. ventral aspect. Figure 21.62 Posfe/Zchus sp.(Dryopidae) adult. ventral aspect, head and sternum. Figure 21.63 Lutrochus sp.(Lutrochidae) adult, dorsal

Figure 21.66 coxa. Figure 21.67 Figure 21.68 Figure 21.69 aspect.

aspect.

Ancyronyx sp.(Elmldae) adult, hind Stenelmis sp.(Elmldae) adult, coxa. Ora sp. (Sclrtldae) adult, hind legs. Prinocyphon sp.(Sclrtldae) adult, dorsal

Chapter 21 Aquatic Coleoptera

807

Figure 21.70 Figure 21.71

Figure 21.78

Figure 21.72

Figure 21.75

Figure 21.73

Figure 21.74

Figure 21.77

Figure 21.76

Figure 21.70 Acneus sp.(Psephenidae) adult, dorsal aspect.

Figure 21.71

Ectopria sp.(Psephenidae) adult, dorsal

aspect.

Figure 21.72 Eubrianax sp.(Psephenidae) adult, frontal aspect of head. Figure 21.73 Psephenus sp.(Psephenidae) adult, dorsal aspect. Figure 21.74 Anchycteis sp. (Ptilodactylidae) adult, frontal aspect of head.

Figure 21.75 Anchycteis sp.(Ptilodactylidae) adult, dorsal aspect. Figure 21.76 Heterocerldae adult, dorsal aspect. Figure 21.77 Aegilites sp.(Salpingidae) adult, dorsal aspect.

Figure 21.78 Donacia sp.(Chrysomelidae) adult, dorsal aspect.

808

25' 26(25) 26' 27(26) 27' 28(25')

28'

Chapter 21

Aquatic Coleoptera

Antennae filiform or serrate (Figs. 21.29, 21.35, 21.36), much longer than head

28 Antennae with 10 or fewer segments; hind coxae contiguous; Fig. 21.63 27 Antennae with 11 segments; hind coxae separated (Fig. 21.64); Fig. 21.64 DRYOPIDAE(in part)(p. 855) Antennomeres 1 and 2 combined representing less than 1/3 of total antennal length LIMNICHIDAE"(p. 853) Antennomeres 1 and 2 combined representing at least 1/3 of total antennal length LUTROCHIDAE Lutrochus(p. 853) Anterior coxae transverse with trochantin visible (Fig. 21.62); antennae (if visible) very short, thick, with enlarged basal segment (Fig. 21.54); Fig. 21.65 DRYOPIDAE (in part)(p. 855) Anterior coxae round, trochantin concealed (Figs. 21.66 and 21.67), antennae slender, filiform (Fig. 21.36); body shape as in Figs. 21.369, 21.371, 21.373

ELMIDAE (in part)(p. 856)

29(24')

Tarsi with 4th segment deeply bilobed (Fig. 21.68); Fig. 21.69

SCIRTIDAE''"(p. 849)

29'

Tarsi usually filiform, 4th segment not bilobed

30

30(29')

Antennae as in Figs. 21.34, 21.35, 21.70, never concealed

31

30'

Antennae filiform or clavate as in Figs. 21.29, 21.36, partly concealed within prosternum; body shape as in Figs. 21.369, 21.371, 21.373

ELMIDAE (in part)(p. 856)

31(30)

Abdomen with 6 or 7 sternites; maxillary palp with 2nd segment longer than next 2 combined; Figs. 21.70 and 21.71 PSEPHENIDAE (in part)(p. 849)

31'

Abdomen with 5 sternites; maxillary palp with 2nd segment much shorter than next 2 combined

32(31')

Head with antennae inserted close together between eyes constricting clypeus from frons(Fig. 21.72); Fig. 21.73

32

PSEPHENIDAE(in part)(p. 849)

32'

Head with antennae inserted below eyes, not constricting clypeus(Fig. 21.74); Fig. 21.75

33(32')

Mandibles prominent, acutely margined above, rectangularly flexed at tip; head not retracted, moderately deflexed; 14-22 mm long (California) EULICHADIDAE Stenocolus(p. 853)

33'

Mandibles not prominent, arcuate at tip, not acutely margined above; head strongly deflexed (Fig. 21.318); less than 12 mm long (widespread) PTILODACTYLIDAE(p. 853)

34(22')

Tarsal formula 4-4-4 or 3-3-3; hind coxae contiguous or nearly so (as in Fig. 21.43)

33

37

34'

Tarsal formula 5-5-4; hind coxae usually separated

35

35(34')

Hind coxae separated by less than coxal width; basal 2 abdominal sternites separated by suture

36

35'.

Hind coxae separated by much more than coxal width; basal 2 abdominal segments fused; Fig. 21.77;

marine intertidal 36(35)

SALPINGIDAE''^-''(p. 868)

Eyes emarginate (notched) anteriorly; procoxal cavities enclosed

behind by prothorax 'Riparian or inhabiting littoral region. ^ Includes intertidal species in North America. '* Keys to genera not given.

TENEBRIONIDAE'-^-'^(p. 868)

Chapter 21

36'

ANTHICIDAE*'^(p. 868)

37(34)

Tarsal formula 4-4-4; beetles larger than 2 mm

37'

Tarsal formula 3-3-3; minute beetles less than 2 mm long;

38

Fig. 21.55

PTILIIDAE'-^-^(in part)(p. 845)

Antennae thickened apically, shorter than head and thorax; mandibles long, projecting horizontally before head;

Fig. 21.76 38'

809

Eyes oval or round; procoxal cavities enclosed behind by mesothorax

(as in Fig. 21.62)

38(37)

Aquatic Coleoptera

HETEROCERIDAE'-'^(p. 853)

Antennae thickened apically, longer than head and thorax (Fig. 21.78); mandibles small, directed ventrally;

Fig. 21.78

FAMILY DESCRIPTIONS AND KEYS TO THE

GENERA OF AQUATIC COLEOPTERA

Amphizoidae (Trout-Stream Beetles) The family Amphizoidae contains a single genus: Amphizoa LeConte. Presently,three species are recog nized, all from northwestern North America where the larvae and adults live in mountain and foothill

streams. Kavanaugh (1986) and Philips and Xie (2001) provide keys to the adults of North American species. They may be especially abundant on drift wood and trash floating in frothy eddies,along under cut banks among roots, or among accumulations of submerged pine needles. Although often rare in col lections, large numbers can be collected once their habitat is recognized. Fggs of amphizoids have been found in cracks on the undersurface of driftwood, although oviposition sites may occur more typically in high humidity or splash zones in partly submerged brush piles. Hatching takes place during middle to late August, and the larvae usually reach the second instar by winter. Larvae(Fig. 21.2)are predaceous and seem to restrict their diet to Plecoptera nymphs. They usually are found crawling on twigs or other wood; mature larvae are almost invariably entirely out of, but near, the water. Such larvae readily enter the water to seize prey, but quickly return to a twig or other support to

CHRYSOMELIDAE'*(p. 868)

its apex. This posture permits the larva to respire while afloat and enables it to quickly capture any prey that comes into reach or to grasp any solid object it touches. Mature larvae have been observed crawling from the water in late July and early August and have been found in protective cases lodged in debris-filled crev ices between logs. Newly emerged adults frequently are mud-covered, which suggests that pupation may occur in muddy creek banks. Adult amphizoids (Fig. 21.46) are poor swim mers usually found crawling on twigs or other sub merged plant material along undercut stream banks and under stones along stream margins. They are predaceous and,like the larvae, show a preference for Plecoptera nymphs. Adult amphizoids have been observed surfacing briefly, then carrying a bubble of air beneath and surrounding the elytral apices while submerged. The highly oxygenated habitat may enable the air bubble to serve as a physical gill, greatly extending the time submerged.

Gyrinidae (Whirligig Beetles) The family Gyrinidae contains approximately 1000 species. Nearly 60 species and subspecies are recorded for the United States and Canada. Whirligig

surface (spiracles of the eighth abdominal tergite at the surface), and the thorax and head are folded

beetles (Fig. 21.27) are a familiar sight on freshwater ponds, lake margins, open flowing streams, quiet stream margins, bog pools, swamps, and roadside ditches where they may form large aggregations or schools in late summer and autumn. These aggrega tions may contain a single species to as many as a dozen or more. Pond-inhabiting gyrinids often fly

under the abdomen so that the mandibles lie beneath

to large streams and lakes to overwinter. Modern

eat the victim.

When dislodged into relatively quiet water, amphizoid larvae assume a characteristic posture with the abdomen at and horizontal to the water's

'Riparian or inhabiting littoral region. ^ Includes intertidal species in North America. Keys to genera not given.

810

Chapter 21 Aquatic Coleoptera

species-level keys for all North American genera are available {Dineutus: Gustafson and Miller 2015, Gyretes: Babin and Alarie 2004, Gyrinus: Oygur and Wolfe 1991). Copulation occurs on the water surface, and the female lays her eggs on stems of emergent vegetation a few centimeters below the surface of the water.

After hatching in 1-2 weeks,larvae (Fig. 21.79) pass through three instars. They crawl about on sub merged objects, using their characteristic apical abdominal hooks (Fig. 21.80) and feed on small aquatic organisms. They can swim in an undulating fashion by using the abdominal gills, possibly as an

escape mechanism. Pupation takes place on shore above water level.

Adults (except for the rare and unusual Spanglerogyrus) are unique in having eyes com pletely separated into two portions (Figs. 21.27, 21.81-21.82). The lower portion remains completely submerged surveying the aquatic habitat; the upper portion views the above water habitat. Divided vision, chemical defense, and quick swimming movements allow them to avoid predators from above or below. They are predominantly scaven gers, feeding upon live or dead insects trapped or floating on the water surface.

Gyrinidae Larvae^

1

Head suborbicular with collum (neck) narrow and distinct (Fig. 21.84); mandible without retinaculum (tooth) on inner margin; nasale with median produced lobe, which may or may not be emarginate, and with a lower tooth on each side (Fig. 21.84); Fig. 21.79 Dineutus

1'

Head elongate, with collum not distinct, nearly as wide as remainder of head (Figs. 21.85-21.86); mandible with retinaculum

2(1')

Nasale with 2-4 teeth in a transverse row (Fig. 21.85)

Gyrinus

2'

Nasale without teeth (Fig. 21.86)(based upon early instar larvae from Missouri)

Gyretes

or without

2

Adults 1

Dorsal and ventral compound eyes in contact on lateral margin of head, separated only by a narrow ridge (Fig. 21.81); meso- and metatarsal segments as long as or longer than broad; length less than 3 mm

Spunglerogyrus

Dorsal and ventral compound eyes divided, upper eyes inset from lateral margin of head for a distance of at least half the width of an eye (Fig. 21.82); meso- and metatarsal segments 2, 3, and 4 much broader than long; total length 3-15 mm

2

2(1')

Lateral margins of pronotum and elytron pubescent (Fig. 21.27); elytron without striae; apical abdominal sternites with median longitudinal row oflong hairs; scutellum concealed; total length 3-5 mm Gyretes

2'

Pronotum and elytron entirely glabrous(without pubescence); apical abdominal sternites without longitudinal row of hairs

3(2')

Larger species, 9-15 mm in length; elytron smooth or with indistinct striae; scutellum concealed

3'

3

Smaller species, 4-7 mm in length; elytron with 11 distinct striae; scutellum visible (as in Fig. 21.81)

'The larva of Spanglerogyrus is unknown.

Dineutus

Gyrinus

Chapter 21

Aquatic Coleoptera

811

terminal hooks

Figure 21.81

lateral

gill

Figure 21.80

Figure 21.82

Figure 21.79

Figure 21.83

nasale

collum

Figure 21.84

U=J Figure 21.85

Figure 21.79 Dineutus sp.(Gyrinidae) larva, dorsal aspect.

Figure 21.80 Dineutus sp.(Gyrinidae) larva, ventral aspect of last abdominal segments. Figure 21.81 Spangierogyrus sp.(Gyrinidae) adult, lateral aspect of head and pronotum. Figure 21.82 Dineutus sp.(Gyrinidae) adult, lateral aspect of head and pronotum.

Figure 21.86

Figure 21.83 Gyrinus sp.(Gyrinidae) adult, antenna. Figure 21.84 Dineutus sp. (Gyrinidae) larva, dorsal aspect of head (after Sanderson 1982b). Figure 21.85 Gyrinus sp. (Gyrinidae) larva, dorsal aspect of head (after Sanderson 1982b). Figure 21.86 Gyrates sp.(Gyrinidae) larva, dorsal aspect of head (after Sanderson 1982b).

812

Chapter 21

Aquatic Coleoptera

Haliplidae (Crawling Water Beetles) Haliplidae is a relatively small family with about 240 species worldwide and with more than 70 species representing four genera from the United States and Canada. Females of the genus Haliplus have been observed to cut a hole with their mandibles in the

side of a filament of the aquatic macrophytes

Ceratophyllum and Nitella and deposit several eggs within the plant cell. Eggs of the genus Peltodytes are deposited on the leaves and stems of aquatic plants where hatching occurs within 8-14 days. Haliplid larvae (Figs. 21.87 and 21.89) pass through three instars and are herbivorous. Pupation occurs from 20 to 25 days after hatching in a spherical pupal chamber constructed by the larva in dry mud. The mature larva remains in the pupal chamber from 4 to 6 days before transformation.

Numerous investigators have discussed the feed ing habits of adult haliplids (Figs. 21.93 and 21.95). Early studies reported that the adults were carnivo rous but later studies have shown them to be herbivo

rous. The greatly expanded hind coxal plates of adult Haliplidae (Figs. 21.42 and 21.44) are unique among aquatic Coleoptera. The air store maintained by these plates is taken in by way of the tip of the abdomen, retained under the elytra, and acts as both a supple mental air store and in hydrostatic functions. One species, Brychius hungerfordi, is federally listed in the United States as 'Rare and Endangered'. It is only known from a few riffles in Michigan and Ontario (Roughley 1991; Strand and Spangler 1994; Keller et al. 1998).

Haliplidae Larvae

1

1'

2(1')

2'

Body segments each with 2 or more erect, segmented, hollow, spine-tipped filaments, each filament half as long as body (Fig. 21.87); forelegs chelate, 4th segment produced apically and edged with a solid row of small teeth so that 5th segment and claw can be closed on it (Fig. 21.88)

Body spines, except in 1st instar, never stalked or much longer than length of 1st body segment(Fig. 21.89); apical abdominal segment produced posteriorly in a forked or unforked horn; forelegs, if chelate, with 4th segment less produced and without solid row of small teeth (Fig. 21.90) Third antennal segment shorter than 2nd; forelegs moderately chelate, but 3rd instead of 4th segment produced, edged with 2 blunt teeth; apical abdominal segment unforked, strongly curved ventrally (Fig. 21.91); body without conspicuous spines Third antennal segment 2-3 times as long as 2nd; body with or without conspicuous spines

3(2')

Foreleg with 3rd segment produced and edged by 2 blunt teeth; apical abdominal segment unforked (except in 1st instar); body with conspicuous spines only on lateral margins

3'

Foreleg weakly to moderately chelate, 4th segment more or less produced, usually bearing 2-3 spines(Fig. 21.90); body with or without conspicuous spines

Peltodytes

2

Brychius 3

Apteraliplus

Haliplus

Adults 1

Pronotum rounded with distinct black blotch on each side of middle

near posterior margin (Fig. 21.93); last segment of both labial and maxillary palpi cone-shaped, as long as or longer than next to last; hind coxal plates large, only last abdominal sternite completely exposed; elytron with fine sutural stria in at least apical half

Peltodytes

Chapter 21 Aquatic Coleoptera

813

5th segment

Figure 21.88

Figure 21.87

4th segment

'Co

Figure 21.90

apical abdominal

segment

groove

Figure 21.91 projections

Figure 21.92

or spines

Figure 21.95

Figure 21.96 prosternum

Figure 21.89

Figure 21.94

Figure 21.93

Figure 21.97

Figure 21.87 Peltodytes sp. (Haliplidae) larva, lateral

Figure 21.93 Peltodytes sp. (Haliplidae) adult, dorsal

aspect.

aspect.

Figure 21.88 Peltodytes sp. (Haliplidae) larva, foreleg. Figure 21.89 Haliplus sp. (Haliplidae) larva, dorsal

Figure 21.94 Brychius sp. (Haliplidae) adult, dorsal aspect of head and pronotum. Figure 21.95 Haliplus sp. (Haliplidae) adult, dorsal

aspect. Figure 21.90 Figure 21.91

Haliplus sp. (Haliplidae) larva, foreleg. Brychius sp. (Haliplidae) larva, lateral

aspect.

Figure 21.96 Apteraliplus sp. (Haliplidae) adult, prosternum.

aspect of apex; of abdomen. Figure 21.92 Peltodytes sp. (Haliplidae) larva.

Figure 21.97 Haliplus sp. (Haliplidae) adult,

mandible.

prosternum.

814

Chapter 21 Aquatic Coleoptera

notch lateral

branch

frontal

projection

Figure 21.100

Figure 21.99

Figure 21.98

-cercus

Figure 21.101

lateral tracheal extension

Figure 21.107

Figure 21.104

A M

Figure 21.103

Figure 21.111

b;

Figure 21.108

Figure 21.109

Figure 21.105

Figure 21.106

Figure 21.98 Derovatellus sp.(Dytiscidae) larva, dorsal aspect of frontal projection of head (after Spanglerand Folkerts 1973). Figure 21.99 Pachydrus sp. (Dytiscidae) larva, dorsal aspect of frontal projection of head (after Spangler and Folkerts 1973). Figure 21.100 Hydroporus sp.(Dytiscidae) larva, dorsal aspect of frontal projection of head. Figure 21.101 Cybister sp.(Dytiscidae) larva, dorsal aspect.

Figure 21.102 Hydrotrupes sp.(Dytiscidae) larva, A: dorsal aspect; B: ventral aspect (after Larson et al. 2000). Figure 21.103 Hydrotrupes sp.(Dytiscidae) larva, lateral aspect (after Larson et al. 2000). Figure 21.104 Celina sp.(Dytiscidae) larva, apex of abdomen (after Spangler 1973).

Figure 21.110

Figure 21.105 Celina sp.(Dytiscidae) larva, dorsal aspect (after Spangler 1973). Figure 21.106 Pachydrus sp.(Dytiscidae) larva. A: dorsal aspect; B: ventral aspect (after Spangler and Folkerts 1973). Figure 21.107 Laccornis sp.(Dytiscidae) larva, terminal abdominal segments (after Larson etal. 2000). Figure 21.108 Hydroporinae sp.(Dytiscidae) larva. A: antenna dorsal; B: antenna ventral (after Larson et al. 2000). Figure 21.109 Oreodytes sp.(Dytiscidae) larva, head, dorsal aspect (after Larson et al. 2000). Figure 21.110 Desmopachria sp.(Dytiscidae) larva, head, dorsal aspect (after Larson et al. 2000). Figure 21.111 Hydroporinae (Dytiscidae) larva, maxilla, ventral aspect. A: with cardo; B: without cardo (after Larson et al. 2000).

Chapter 21 Aquatic Coleoptera

r

Pronotum immaculate or with median blotch anteriorly (Fig. 21.95), posteriorly, or both or with 2 indistinct blotches (Fig. 21.94); last segment of both labial and maxillary palpi sublate, shorter than next to last; hind coxal plates smaller, leaving last 3 abdominal sternites exposed; elytron without fine sutural stria

2(1')

T

3(2')

3'

815

2

Pronotum with sides of basal two-thirds nearly parallel(Fig. 21.94); epipleuron broad, extending almost to tip of elytron, which is never truncate; metasternum reaching epipleuron Pronotum with sides widest at base, convergent anteriorly (Fig. 21.95); epipleuron evenly narrowed, usually ending near base of last abdominal sternite, never reaching elytral apex; episternum completely separating metasternum from epipleuron Median part of prosternum and base of prosternal process forming a plateau-like elevation, at least in part angularly separated from sides of prosternum (Fig. 21.97) Prosternum evenly rounded from side to side, process raised above base (Fig. 21.96); tiny, length 1.5-2.5 mm; California north to Washington

Dytiscidae (Predaceous Diving Beetles) Dytiscidae are among the best adapted insects for aquatic existence and the most diverse of the water beetles with >4,300 described species, including more than 500 that occur in North America.

Despite the size and importance of this group among water beetles, surprisingly little is known of the life history of North American dytiscids. Mating occurs from early spring through autumn. Oviposition sites of dytiscids appear to correlate well with structural modifications of the ovipositor. Species having a long, flexible ovipositor (some Acilius) place their eggs loosely, usually 30-50 in a mass, above water in moist soil among grass roots or under organic debris. Most species possessing a cutting ovipositor (some Agabus, Coptotomus, Cybister, Dytiscus, Hydaticus, Ilybius, Laccophilus, and Thermonectus) insert their eggs into parts of living plants. A third group places its eggs on plant surfaces or inserts them, at most, halfway into the plant tissues(some Agabus, Colymbetes,Hydropoms, and Rhanlus). The larval stage (Figs. 21.112, 21.113, 21.114, 21.116, 21.121, 21.124, 21.125) of the Dytiscidae consists of three instars and requires from several weeks to several months, depending mainly upon season and the availability of food. Most dytiscid larvae rise to the surface and take in air through the large terminal spiracles. Cuticular respiration, how ever, appears to be common among first instar lar vae of many species. Agabus, Ilybius, and some Hydropoms have extensive networks of tracheae

Brychius

3

Haliplus Apteraliplus

near the ventral cuticle and are believed to exchange gases through this structure. Larvae of the genus Coptotomus possess lateral gills and can remain beneath the surface continuously (Fig. 21.113). All mature dytiscid larvae leave the water to prepare a pupal cell on land near the water's edge. As the common name for the group implies, larval dytiscids are predaceous. Their selection of food appears to be governed by their ability to catch and overcome their prey. Although most adult dytiscids are active predators, many also are scavengers.

Adult dytiscids (Figs. 21.131, 21.137, 21.144, 21.147,21.155,21.172,21.178,21.179,21.180,21.181,

21.187) readily leave the water and fly. Dytiscids splash directly into the water because the extreme

modification of the legs for swimming makes them useless for alighting on surfaces. Impact usually car ries them through the surface film,but frequently they return to the surface after a few seconds to fill the

subelytral air chamber. Presumably, the tracheal sys tem undergoes a reversal ofthe process in preparation for flight. There has been substantial progress over the last

decade in our understanding of diving beetle evolu tion and classification. Numerous phytogenies have been published based on varied data sources that have

finally begun to solidify the higher-level classification of the family. See Miller and Bergsten (2016) for a complete review of dytiscid classification and taxon omy at the global scale, including excellent keys to genus (adults only)for the world fauna.

816

Chapter 21 Aquatic Coleoptera

Dytiscidae Mature Larvae^

1

Head, anteriorly, with a frontal projection or nasale (Figs. 21.98-21.100); mandibles, in lateral aspect, distinctly curved upward; urogomphus 2-segmented HYDROPORINAE ...2

r

2(1)

Head, anteriorly, without a frontal projection (Figs. 21.101- 21.102); mandibles, in lateral aspect, not distinctly curved upward; urogomphus 1-or 2-segmented Stemmata (simple eyes) absent; subterranean, Texas

2'

Stemmata present, if absent, surface dwelling

3(2)

Last abdominal segment tapered to a point; first segment of urogomphus about three times as long as the second segment Last abdominal segment truncate; first segment of urogomphus about ten times as long as the second segment

y 4(2')

3

4 Haideopoms

Ereboporus

Last abdominal segment with recurved extension of lateral tracheal

trunks reaching beyond apex of abdomen (Fig. 21.104); urogomphus very short, >1/2 length of last abdominal segment(Fig. 21.105) 4'

25

Lateral tracheal trunks not extending beyond apex of last abdominal segment, terminating on apex; urogomphus of variable length

Celina

5

5(4')

Abdominal segment 6 sclerotized ventrally (Fig. 21.106B)

5'

Abdominal segment 6 membranous ventrally (as in Fig. 21.102B)

10

6(5)

Urogomphus longer than abdominal segment 8; head with frontal projection without distinct lateral notches (Fig. 21.100)

Desmopachria

6'

6

Urogomphus shorter than abdominal segment 8; head with frontal projection with lateral branches (Figs. 21.98 and 21.99)

7

Frontal projection of head elongate, constricted near base and broadly spatulate at apex, a lateral branch arising from each side near base (Figs. 21.98 and 21.99)

8

7'

Frontal projection broadly triangular, not constricted near base and with or without a lateral branch on each side, although a notch may be present on each side (Fig. 21.100); Fig. 2L106B

9

8(7)

Frontal projection with lateral branch with 2 spines on outer margin (Fig. 21.98); Florida

7(6')

8' 9(7')

9'

10(5') 10'

Frontal projection with lateral branch with 3 spines on outer margin; Texas Larva greatly widened in middle, greatest width approximately 0.25 of total length; leg with femur without swimming hairs along ventral margin; head with frontal projection without lateral branch Larva not greatly widened at middle (Fig. 21.106A); leg with femur with swimming hairs along ventral margin; head with frontal projection with lateral branch (Fig. 21.99) Stemmata absent or if present individual stemma reduced in size and subequal to maximum width of basal antennomere Stemmata present, larger and well-defined, at least 2x width of basal antennomere

Derovatellus

Vatellus

Hydrovatus

Pachydrus Hydrocolus 11

^ The larvae of Bidessonotus, Comaldessus, Crinodessus, Hydrodytes, Lioporeus, Neohidessus,Psychopompoms,and Stygoporus are unknown or undescribed. Anodocheilus, recently described from Neotropical material, is not included. In addition, some groups of genera, such as Nehrioporus/StictotarsuslBoreonectes, LiodessiislNeoclypeodytes and several genera of Agabini are known only from a few larval descrip tions and may be incompletely separated. The reader is referred to Larson et al.(2000)and Michat et al.(2017)for a more detailed discussion of taxonomy and identification of immature stages.

Chapter 21

11(IO') 1r

12(11') 12' 13(12) 13' 14(13')

Aquatic Coleoptera

Urogomphus very short, about one-half length of last abdominal segment; last abdominal segment not constricted at insertion of urogomphi (Fig. 21.107) Urogomphus longer, at least as long as last abdominal segment; last abdominal segment constricted at insertion of urogomphi Urogomphus with more than 8 setae (with secondary setae) Urogomphus with only 8 setae (it may appear as 7 as the seta that is proximad to the insertion of the urogomphus on the last abdominal segment is minute) Size extremely small(body length up to 0.5 mm); Florida Size greater than 0.5 mm

817

Laccornis 12 13

19 BIDESSINI(in part) Brachyvatus 14

Antennomere III with an anteroventral spinula (Fig. 21.108B)and with

(Fig. 21.108A)or without a lateral pore

15

14'

Antennomere without anteroventral spinula and lateral pore

15(14) 15'

Maxilla with cardo absent(Fig. 21.11 IB); antennomere III without lateral pore 16 Maxilla with cardo present(Fig. 21.1 llA); antennomere III with lateral pore (Fig. 21.108A).... 18

16(15)

Legs without swimming hairs; head capsule distinctly narrowed at level of occipital line (Fig. 21.109)

16'

17(16')

Legs with swimming hairs; head capsule, at most, slightly constricted at level of occipital line

Hygrotus (in part)

Oreodytes 17

Head capsule broader, less than 1.3 times as long as broad

Nebriopoms

17'

Head capsule longer, more than 1.4 times as long as broad

StictotarsusIBoreonectes^

18(15')

Legs with swimming hairs

18'

Legs without swimming hairs

19(12') 19' 20(19)

Legs with swimming hairs 20 Legs without swimming hairs 21 Antennomere III with anteroventral spinula and lateral pore (Figs. 21.108A and 21.108B) Neoporus (in part) Antennomere III without anteroventral spinula and lateral pore Hygrotus (in part) Smaller specimens; urogomphus with 3 long, more basal setae originating about equally from each other BIDESSINI (in part)...23 Larger specimens; urogomphus with 3 long, more basal setae more grouped together, not originating equally from one another .... HYDROPORINI(in part)... 22 Urogomphus shorter, less than 2.4 times as long as last abdominal segment Hydroporus Urogomphus longer, more than 2.6 times as long as last abdominal segment SanfiUppodytes Urogomphus more than 3 times as long as last abdominal segment Liodessus Urogomphus less than 2 times as long as last abdominal 24 Labial palp with segments distinctly unequal; basal segment distinctly

20' 21(19') 21' 22(21') 22' 23(21) 23' 24(23') 24' 25(1') 25' 26(25')

longer than second segment Labial palp with segments subequal; basal segment equal to or shorter than second segment (Fig. 21.112)

Aeo/>on/s (in part) Heterosternuta

Uvarus Neoclypeodytes

Abdominal segments 1 to 6 with lateral gills; lateral margins of segment 8 and urogomphus with fringe of long setae (Fig. 21.113) COPTOTOMINAE...Co/»rorom//s Abdominal segments 1 to 6 without lateral gills; lateral margins of segment 8 and urogomphus with or without fringe of long setae 26 Legs(femur and tibia) and lateral margins of last two abdominal segments (segments 7 and 8) with fringes of long setae or swimming hairs

DYTISCINAE...27

' Some North American species ofStictotams were recently moved into a new genus, Boreonectes, by Angus(2010)based largely on karyotypic data. Boreonectes currently has no known morphological synapomorphies and is difficult to diagnose, and may not be monophyletic (see Miller and Bergsten 2016 for a fuller discussion).

818

Chapter 21 Aquatic Coleoptera

26'

Legs and lateral margins of segments 7 and 8 without fringes of

27(26)

Head with anterior margin indented and appearing trilobed (Fig. 21.115); urogomphus very short and vestigial (Fig. 21.114) CYQ\STKWl...CybisterlMegadytes^ Head with anterior margin entire; urogomphus long and apparent 28 Head with posterior margin deeply indented medially; anterior margin of labium, between bases of labial palpi, modified anteriorly as the ligula 29 Head with posterior margin, at most, slightly indented medially; anterior margin of labium, between bases of labial, not modified, ligula absent 32

swimming hairs or long setae

27' 28(27')

28' 29(28)

29'

30(29') 30'

31(30') 31'

32(28') 32' 33(26') 33' 34(33')

34'

35(34)

33

Ligula short and simple but apically with 4 spines; head with stemmata of uniform size, not greatly enlarged Ligula long and simple or bifid and with less than 4 apical spines; head with some stemmata greatly enlarged (Fig. 21.121)

ERETINI... Freto

AC1L11NL..30 Ligula simple apically Graphodems Ligula bifid or least slightly sinuate apically 31 Ligula deeply sinuate apically and with short, spine-like setae (Fig. 21.122) Acilius Ligula shallowly sinuate apically and with elongate, spine-like setae (Fig. 21.123) .... Thermonectus Urogomphus laterally without fringe of swimming hairs; head laterally with temporal spines HYDATICINL..//j5 mm; Texas Vatellus Body short and broad (Fig. 21.144); length/width e/at«s

44(42')

Labial palpus very short, apical palpomere subquadrate (Fig. 21.185); western North America; Figure 21.187 AGABlNAE...HYDROTRUPINI.../7j14 mm

Rhantus^

Noteridae (Burrowing Water Beetles)

without appearing to remove any tissue. Noterid lar-

ThefamilyNoteridaeisdistributedwidely throughout the tropical regions of both hemispheres with only a few genera and species reaching the temperate zone. The family is represented in North America north of Mexico by five genera and about 15 species. Recent morphological(Miller 2009)and molecular(Baca et al. 2017) studies have clarified the relationships among noterid genera and significantly reorganized the internal classification of the family. With regard to the North American fauna, the genus Pronoterus is now considered a synonym of Suphisellus. The Palaearctic species Noterus crassicornis (Muller 1776) has been investigated thoroughly and its burrowing habits are the source of the family com-

also readily attack dead chironomid larvae. The morphology of the mandible suggests an omnivorous are predaceous. I" shallow water, larvae of N. crassicornis renew ^^eir air supplies by bringing the apex of the abdomen ^^e surface. However, m the typical method of obtaining air, the apical spine of the abdomen is used to pierce plant tissue to tap intercellular air in the manner of chrysomelid beetles of the genus Donacia Pupation of N. crassicornis is similar to that of The larva constructs a cocoon from small pieces of vegetable material mixed with mud particles ^oots of various species ofaquatic plants. Larvae ^^ew or pierce the plant root at the point ofattachment

mon name. Observations of noterid larvae in eastern

the cocoon. Air escaping from the lacerated tissue is

North America indicate that most, if not all, of these

^^ught in the cocoon as it is being constructed and

species do not share the strict burrowing habits of their European relatives. In fact, they lack the chitinous point at the apex of the abdomen with which A. crassicornis is presumed to pierce plant roots for the purpose of obtaining intercellular air. Food habits of the larvae (Figs. 21.199 and 21.200) are unknown. Larval N. crassicornis have been observed working their mandibles upon the surfaee of plant roots

within when the larva closes the distal end. Whether North Ameriean larvae of Noteridae behave ^ ^™har manner to N. crassicornis is unknown. The length of the pupal period is unknown but presumably more than a few weeks. Adults (Figs. 21.205 ^nd 21.206) are often confused with dytiscids but they ^^sily separated on the basis of leg structure and features of the ventral surface.

Noteridae

La/vae"

1 1'

Third antenna) segment not longer than 4th; mandible with stout preapical tooth Third antenna)segment more than twiee as long as 4th; mandible not strongly toothed

2(1')

Body globular (Fig. 21.199); 3rd antenna) segment about 12 times as long as 4th;

2'

Body cylindriform, not globular (Fig. 21.200); 3rd antenna) segment about 3 times longer than 4th; mandible simple

mandible serrulate

Suphisellus 2 Suphis

Hydrocanthus

'Two North American species previously included in the genus Rhantus were recently split apart into new or reinstated genera: R. sinuatus was moved to the genus Nartus and R. calidus was moved to the genus Meridiorhantus(see Balke et al. 2017). "The larvae of Notomicrus and Mesonoterus are undescribed.

834

Chapter 21

Aquatic Coleoptera

Figure 21.199

Figure 21.205

Figure 21.200 ,Incisor lobe

molar lobe

Figure 21.201

Figure 21.202

hook

'claws

Figure 21.203

Figure 21.204

Figure 21.199 Suphis sp.(Noteridae) larva, dorsal aspect (after Spangler and Folkerts 1973). Figure 21.200 Hydrocanthus sp.(Noteridae) larva, dorsal aspect. Figure 21.201 Noterus sp.(Noteridae) larva, foreleg (not In key)(after Bertrand 1972). Figure 21.202 Noterus sp.(Noteridae) larva, mandible (not in key)(after Bertrand 1972).

Figure 21.206

Figure 21.203 Hydrocanthus sp.(Noteridae) adult, foreleg. Figure 21.204 Notomicrus sp.(Noteridae) adult, foreleg. Figure 21.205 Suphis sp.(Noteridae) adult, dorsal aspect.

Figure 21.206 dorsal aspect.

Hydrocanthus sp.(Noteridae) adult,

Chapter 21

Aquatic Coleoptera

835

Adults 1

Apex of foretibia with curved hook or spur (Fig. 21.203); length usually over 2.0 mm

r

Apex of foretibia without curved hook or spur (Fig. 21.204); length usually

2(1)

Fore tibial hooks strong, curved, and conspicuous(Fig. 21.203); hind femur with angular cilia; prosternal process truncate posteriorly, or if rounded in the male, body form is very broad, almost hemispherical

less than 1.5 mm

2'

Notomicms

3'

3

Fore tibial hooks weak and inconspicuous; hind femur usually without angular cilia; prosternal process rounded posteriorly in both sexes

3(2)

2

Body form very broad, almost hemispherical (Fig. 21.205); foretibia with setal comb greatly reduced, absent, on outer margin and apex; color opaque black with irregular reddish marks on each elytron

5

Suphis

Body form elongate, not hemispherical (Fig. 21.206); foretibia with conspicuous setal comb extending distally along outer margin to apex; elytron uniformly black, reddish brown, or yellowish brown, without

markings or with an oblique yellowish crossbar just behind the middle, never with reddish marks

4(3')

4

Length usually less than 3 mm;apical segment of maxillary palpus emarginate at apex; prosternum with series of stiff setae anteromedial to forecoxae, prosternal process not broader than long, apex at least

twice its breadth between the anterior coxae 4'

Suphisellus (in part)

Length usually over 4 mm;apical segment of maxillary palpus truncate at apex; prosternum without series of stiff setae; prosternal process broader than long, apex very broad, at least 2.5-3 times its breadth between the anterior coxae;

Fig. 21.206 5(2')

Hydrocanthus

Foretibia elongate; body more attenuate posteriorly; metatibial spur smooth, not serrate; prosternum without stiff setae; length 2.7 mm or more Mesonotems Foretibia broader, triangular; body more oval; metatibial spur serrate; prosternum with few stiff setae anteromedial to forecoxae;

length 2.6 mm or less

Hydroscaphidae (Skiff Beetles) The Hydroscaphidae is a small family of nearly worldwide distribution.Only the genus Hydroscapha

Suphisellus (in part)

hydroscaphids frequently are very abundant and hundreds may be collected in a rather short time. Although adults (Fig. 21.31) may be found under

LeConte is known from western North America.

stones as much as a meter below the surface of

Although previously only a single species (H.

fast-flowing streams, larvae (Fig. 21.18) and adults occur most commonly on algae over which a thin film of water is flowing. Often the water is so shal low that the adults are only partially submerged. Hydroscapha occurs over a wide range of tempera tures, from hot springs(46°C) to mountain streams that freeze nightly throughout summer. Only one, remarkably large egg develops at a time, occupying a fourth of the female's abdomen. Larvae (Fig. 21.18) and adults (Fig. 21.31) remain in and feed upon algae. Pupation occurs at the edge of the water film or even among algae over which the water film is flowing. Pupae lie partially within the

natans LeConte) was known from the United States, a second species (H. redfordi) was recently described from hot springs in Idaho (Maier et al. 2010). This work also suggests that other isolated

North American populations may represent addi tional undescribed species. A molecular phytogeny of the entire family along with a world species checklist was recently published by Short et al. (2015). Hydroscaphids are rare in collections probably because most collecting is done with nets having too coarse a mesh to retain them, in addition to being often localized in distribution. Once located.

last larval exuviae.

836

Chapter 21

Aquatic Coleoptera

In larval H.natans, only the first pair ofthoracic spiracles and those of the first and eighth abdominal segments (Fig. 21.31) are functional. All three pairs are balloon-like, more than twice as long as wide. Presumably gas exchange takes place across the walls of these "balloons" providing a water-air interface available for respiration that is about an order of magnitude greater per milligram wet body weight than that across the spiracles of terrestrial

intertidal zones. Smetana (1978) provides detailed species-level keys to all species of Sphaeridiinae in

insects.

Hydrophilidae, see Short and Fikacek (2013). Eggs are deposited underwater in silk cases that may contain more than 100 eggs. Larvae go through three instars rapidly in one to several months. Even though a few species possess lateral gills allowing them to occupy deeper habitats, most larvae must obtain oxygen at the waterline through terminal abdominal spiracles. The larvae are poor swimmers and tend to lie in wait for prey. The genera Hydrophilus, Tropisternus and Hydrobius often consume their prey

Adult Hydroscapha apparently breathe by means of an air bubble carried beneath the elytra (physical gills, see Chapter 4). A fringe of cilia forms a plastron on the hind wings, and setae on the dorsum covered by the elytra serve to retain this bubble.

Sphaeriusldae (Minute Bog Beetles) Sphaeriusidae consists of a single genus {Sphaerius) with 19 species worldwide. Three species exist in North America distributed from Texas to

Washington. Because of systematic conflicts with Sphaeriidae (fingernail clams), the family and genus have undergone several name changes in the past sev eral decades. Microsporidae and Microsporus are more recent synonyms.

Much of their biology apparently is similar to Hydroscapha. Females produce one egg at a time. Both adults (Fig. 21.45) and larvae are found together along the margins of streams and bogs, usually under objects or in leaf litter where they feed on algae. Larval Sphaerius have balloon shaped spiracular gills on all eight abdominal segments rather than just the first and eighth segments of Hydroscapha. Adult Sphaerius have an air reserve beneath the elytra but lack the plastron of Hydroscapha.

North

America

and

remains an

authoritative

resource for the subfamily. The higher-level classifi cation of the family has undergone substantial changes in the last decade as new morphological and molecular data have shed light on our under standing of hydrophilid evolution. For a detailed review

of

the

current

classification

of

the

out of water. When mature, larvae leave the water to

construct pupal chambers in moist soil, under rocks, and in organic debris. Some species may pupate in emergent vegetation and floating algal mats some distance from shore.

Some adult hydrophilids are good swimmers but not as active as many of the Dytiscidae, with other groups of water scavenger beetles prefer to cling to debris or detritus and rarely if ever swim in the open water. They must return to the surface to renew their air supply. Typically they break the surface film with the antennae and side of the head; this allows gas

exchange along the plastron and air passage on the ventral surface of the thorax. Some gas exchange, providing an auxiliary oxygen source, may occur via the plastron while submerged. Adults are active flyers and have been shown capable of leaving the water many times. Mass emergence and flight periods are not uncommon, and large numbers may be attracted

Hydrophilidae (Water Scavenger Beetles)

to lights.

With about 3,000 species, including more than 250 in North America, the Hydrophilidae ranks sec ond in abundance among water beetles only to Dytiscidae, and these two families most often come to mind when one considers aquatic Coleoptera. The common name is not completely accurate because most hydrophilid larvae are predators, most adults are omnivores, consuming both living and dead materials, and many taxa of the subfamily Sphaeridiinae are not aquatic. The Sphaeridiinae are not included in the key, although species of Cercyon and Phaenonotum may be collected near water,and two species of Cercyon occur in California

Hydrophilidae are most common in small pools and ponds with emergent vegetation and few preda tors, but occur in most aquatic habitats including stream and river margins, seepage areas, and salt marshes. Adults can be collected throughout most of the year. With the exception of a few of the more common genera, larvae are rarely collected, perhaps because of the relatively short larval cycle, compared with most other families of aquatic Coleoptera.

Hydrochara rickseckeri is on the "Special Animals List" in the State of California due to its uncertain

and potentially threatened conservation status(Short et a/. 2017b).

Chapter 21 Aquatic Coleoptera

837

Hydrophilidae larvae"

1 r 2(1')

Abdominal segments with very long and thin lateral projections (Fig. 21.210) Abdominal segments without lateral projections, or if projections present (Figs. 21.226, 21.227, 21.229), not nearly so long and prominent Legs completely invisible from above and very reduced, 3-segmented, without claws; ligula present

2'

Legs sometimes very small and not visible from above, but always complete, 5-segmented, with claws (Fig. 21.234); ligula present or absent

3(2')

Mesothorax, metathorax, and 1 st abdominal segment each with distinctly branched, setiferous, lateral gills (Fig. 21.225); abdominal segments 2 through 6 each with 4 moderately long, setiferous, lateral gills (Fig. 21.226); southeastern United States Abdominal gills absent or, if present, with only a single lateral gill on each side of abdominal segments (Fig. 21.229)

y 4(3')

Berosus 2 Chaetarthria

3

Demllus 4

Mandibles strongly asymmetrical, either in shape or in number of teeth (as in Figs. 21.219 and 21.233)

5

4'

Mandibles symmetrical or nearly so in both shape and in number of teeth (as in Figs. 21.214, 21.239, 21.242)

9

5(4)

Frontal sutures parallel (Fig. 21.213) and not uniting to form an epicranial suture; ligula absent

5'

Frontal sutures not parallel(as in Fig. 21.242); epicranial suture present or not; ligula present

Laccobius 6

6(5')

First antennomere slightly shorter to as long as second antennomere

6'

First antennomere much longer than second (Fig. 21.228)

7

7(6')

Head subspherical; frontal sulci U-shaped; mandibles not symmetrical (Figs. 21.228 and 21.232)

8

7'

8(7)

Enochrus

Head subquadrangular or subrectangular (Figs. 21.230 and 21.242), frontal sulci V-shaped; mandibles symmetrical or not(Figs. 21.230 and 21.239), pronotum entirely sclerotized

Tropisternus

Mandibles robust, right mandible with a blunt tooth (Fig. 21.233), left mandible with a notch; abdominal segments with pairs of lateral

lobes with setae (as in Fig. 21.235); eastern North America

Hydrophilus (Dibolocelus)

8'

Right mandible thinner than left mandible, right mandible with bifid tooth (Fig. 21.232); abdominal segments without lateral setae Hydrophilus {Hydrophilus)

9(4')

Abdominal segments with broad, lateral projections (Figs. 21.227 and 21.229)

10

9' 10(9)

Abdominal segments without fleshy projections(as in Figs. 21.238, 21.241) Thoracic segments with broad, lateral fleshy projections(Fig. 21.227) and abdominal segments with slightly shorter fleshy projections; body usually tiny, less than 5 mm Thoracic segments with no or only very small lateral projections, much smaller than the lateral abdominal gills, which are well developed and pubescent (Fig. 21.229); body length usually larger, greater than 5 mm in mature larvae

11

10'

11(9') 11' 12(11)

Crenitis

Hydrochara

Posterior margin of tergite YIII weakly to strongly trifid 12 Posterior margin of tergite VIII concave or convex, never trifid 13 Clypeus with middle tooth (of five) more closely positioned to the two teeth to its right... Crenitulus

The larvae of the terrestrial subfamily Sphaeridiinae are not included in the key, as is the larvae of Hemiosus, known only in the Nearctic region from southern Arizona.

clypeus

'suture

Figure 21.213

Figure 21.208

Figure 21.215

Figure 21.217

Cjjp Figure 21.211

biforous

spiracle

Figure 21.221

Figure 21.209

Figure 21.207

Figure 21.223

Figure 21.216

Figure 21.214

Figure 21.224

Figure 21.219

Figure 21.210

Figure 21.220 Figure 21.222

Figure 21.207 Helophorus sp.(Helophoridae) larva, dorsal aspect. Figure 21.208 Epimetopus sp.(Eplmetopidae) larva, abdominal segments 8 and 9. Figure 21.209 Hydrophilidae larva, spiracle. Figure 21.210 Berosus sp.(Hydrophilidae) larva, dorsal aspect. Figure 21.211

Berosus sp.(Hydrophilidae) larva, dorsal aspect of mandibles. Figure 21.212 Berosus sp.(Hydrophilidae) larva, clypeus. Figure 21.213 Laccobius sp.(Hydrophilidae) larva, dorsal aspect of head. Figure 21.214 Anacaena sp.(Hydrophilidae) larva, dorsal aspect of mandibles. Figure 21.215 Anacaena sp.(Hydrophilidae) larva, clypeus. Figure 21.216 Paracymus sp.(Hydrophilidae) larva, dorsal aspect of right mandible. 838

Figure 21.218

Figure 21.225

Figure 21.217 Paracymus sp.(Hydrophilidae) larva, clypeus. Figure 21.218 Enochrus sp.(Hydrophilidae) larva, dorsal aspect. Figure 21.219 Enochrus sp.(Hydrophilidae) larva, dorsal aspect of mandibles. Figure 21.220 Ametor sp.(Hydrophilidae) larva, ventral aspect of head (after Bertrand 1972). Figure 21.221 Ametor sp.(Hydrophilidae) larva, clypeus (after Bertrand 1972). Figure 21.222 Helochares sp.(Hydrophilidae) larva, dorsal aspect. Figure 21.223 Helochares sp.(Hydrophilidae) larva. clypeus. Figure 21.224 Cymbiodyta sp.(Hydrophilidae) larva, clypeus (after Richmond 1920). Figure 21.225 Derallus sp. (Hydrophilidae) larva, dorsal aspect.

Chapter 21

12'

Clypeus with middle tooth (of five) clearly and strongly separated from each group of lateral teeth (Fig. 21.215)

13(11') 13'

First antennomere slightly shorter to as long as second antennomere First antennomere much longer than second (as in Fig. 21.228); southeastern United States Clypeus with deep medial emargination, without teeth;

14(13)

Aquatic Coleoptera

Anacaena 14 Hydrobiomorpha

southeastern United States

Helobata

14' 15(14') 15'

Clypeus without deep medial emargination, with two or more teeth Clypeus with 3 or 4 teeth Clypeus with 5 or more teeth {Sperchopsis may appear as having four teeth as the middle tooth is small but present as in Fig. 21.240)

16(15) 16' 17(15') 17' 18(17')

Frontal sulci Y-shaped; clypeus symmetrical; western North America Frontal sulci U-shaped or lyriform; clypeus slightly asymmetrical (Fig. 21.217) Clypeus with 5 distinct teeth though middle tooth may be small(Fig. 21.240) Clypeus with 6 or more distinct teeth; mandible with 2 inner teeth Clypeus with 6 distinct teeth, placed in 2 groups, 2 on the left and 4 on the right (Fig. 21.223); Fig. 21.222

15 16 17

18'

Clypeus with more than 6 teeth, those toward right not as clearly defined and with several smaller teeth (Fig. 21.224)

19(18')

Clypeus with tooth on left side smaller than tooth on right side (Fig. 21.224)

19'

Clypeus with left and right side teeth equal in size; eastern United States

20(17)

Middle tooth on clypeus smaller than the others (Figs. 21.239 and 21.240); prosternum entire; eastern United States; Fig. 21.238

20'

839

Ametor Paracymus 20 18 Helochares 19

Cymbiodyta

Helocombus^^ Sperchopsis

All teeth on clypeus subequal, outer left tooth usually a little distant from

the rest(Fig. 21.242); prosternum with a mesal fracture; Fig. 21.241

Hydrobius!Limnohydrobius^^

Adults

1

First segment of hind tarsi longer than 2nd; labrum not well sclerotized, usually retracted under clypeus; second labial palpomere with subapical tufts of numerous setae

r

2(1')

2'

SPHAERIDIINAE, not covered here

First segment of hind tarsi shorter than 2nd (except Cymbiodyta & Helocombus); labrum well sclerotized and exposed, not concealed by clypeus(except Helobata); second labial palpomere with scattered setae 2 Meso- and metaventral keels fused to form a single structure, often ending posteriorly in a pronounced spine (Fig. 21.246); body size large to very large(5-50 mm) F1YDROPHILINAE...HYDROPF1ILINI...3 Sternal structure not as above, never ending in large spine; body size variable, but never exceeding 15 mm

7

3(2)

Prosternum with median carina deeply emarginated or completely divided into two lobes for reception of anterior portion of mesosternal keel

4

3'

Prosternum with median carina not emarginated or divided

6

4(3)

Smaller, less than 15 mm in length; anterior margin of clypeus straight; labrum usually partly or entirely covered with fine setae Larger, greater than 20 mm in length; anterior margin of clypeus broadly emarginate; labrum without dense, fine setae Prosternal process closed in front, hood-shaped (Fig. 21.253); widespread

4'

5(4')

Tropisternus

5 Hydrophilus {s. str.)

The monotypic genus Helocombus is currently considered valid but unpublished data suggests it may be only be variant of Cymbiodyta; likely to be synonymized in the near future. Recent phylogenetic studies(Short et at. 2017a)found that the genus Hydroblus was not monophyletic and transferred two of the three North American species to the newly elevated Limnohydrobius.The only North American species remaining in Hydroblus is the widespread H.fuscipes.

right mandible

Figure 21.226 lateral gill

Figure

Figure 21.227

labium Jtiaxillary palp palpifer

stipes—If mandib e

cardo

Figure 21.235

Figure 21.229 Figure 21.231

Figure 21.230 Figure 21.237

Figure 21.232

Figure 21.236

Figure 21.240

femur

Figure 21.233 trochanter

tibiotarsus

claw

Figure 21.234

f suture

OD

Figure 21.239 Figure 21.238

Figure 21.241

Figure 21.226 Derallus sp.(Hydrophiiidae) larva, dorsal aspect of 2nd abdominal segment. Figure 21.227 Crenitus sp.(Hydrophiiidae) larva, dorsal aspect of 2nd abdominal segment (after Bertrand 1972). Figure 21.228 Hydrophilus (s. str.) sp.(Hydrophiiidae) larva, dorsal aspect of head. Figure 21.229 Hydrochara sp.(Hydrophiiidae) larva, dorsal aspect. Figure 21.230 Hydrochara sp.(Hydrophiiidae) larva, dorsal aspect of head. Figure 21.231 Hydrochara sp.(Hydrophiiidae) larva, ventral aspect of head. Figure 21.232 Hydrophilus (s. str.) sp.(Hydrophiiidae) larva, mandibles.

Figure 21.233 Hydrophilius (subgenus Diboioceius) sp.(Hydrophiiidae) larva, mandibles. 840

Figure 21.242

Figure 21.234 Hydrochara sp.(Hydrophiiidae) larva, leg. Figure 21.235 Tropisternus sp.(Hydrophiiidae) larva, dorsal aspect. Figure 21.236 Tropisternus sp.(Hydrophiiidae) larva, dorsal aspect of last abdominal segments. Figure 21.237 Hydrobiomorpha sp.(Hydrophiiidae) larva, clypeus (after Bertrand 1972). Figure 21.238 Sperchopsis sp.(Hydrophiiidae) larva, dorsal aspect. Figure 21.239 Sperchopsis sp.(Hydrophiiidae) larva, dorsal aspect of head. Figure 21.240 Sperchopsis sp.(Hydrophiiidae) larva, clypeus. Figure 21.241 Hydrobius sp.(Hydrophiiidae) larva, dorsal aspect. Figure 21.242 Hydrobius sp.(Hydrophiiidae) larva, dorsal aspect of head.

Chapter 21 Aquatic Coleoptera

5'

Prosternal process not closed in front, bifurcate (Fig. 21.256); eastern North America

6(3')

Anterior margin of clypeus broadly emarginated (Fig. 21.248), exposing thin membrane; southeastern United States

6' 7(2') 7' 8(7)

8'

9(8')

841

Hydrophilm (subgenus Dibolocelus) Hydrobiomorpha

Anterior margin of clypeus straight (Figs. 21.249-21.251); widespread Hydrochara Scutellum much longer than wide (Fig. 21.255); middle and hind tibia with fringe oflong swimming hairs on dorsal face (Fig. 21.255)....HYDROPHILINAE...BEROSINI...8 Scutellum as long as wide or rather longer than wide; middle and hind tibia without fringe of long swimming hairs 10 Dorsum entirely black. Body form subglobular (Fig. 21.254), laterally compressed; metaventrite with medial carina; metanepistenum partially or completely concealed by the elytral margins; eastern United States, particularly the southeast Dorsum at least partially (usually mostly) yellow or brown with scattered black markings; body form slightly convex, not laterally compressed; metaventrite with medial hollow more or less developed; metanepistenum apparent

Demllus

9

Mesoventrite with laminar elevation; abdominal ventrite I may have a median carina and/or lateral excavations but not side carinae;

pubescence on hind femur long, less dense; protarsus of males with four tarsomeres; size variable (1.5-9 mm).(Fig. 21.255); very common and widespread 9'

Berosus

Mesoventrite with elevated plate, usually medially hollowed; abdominal ventrite I with a medial and two side carinae;

pubescence on hind femur short, dense; protarsus of males with five tarsomeres; size small( ) ; ) ) ) ) ))) ) ) ))) )

00

00

05

Family Genus

Matus(4)

Lioporeus(2)

Liodessus(9)

Laccornis(9)

Laccophilus(14)

Laccomimus

llybius(14)

Hygrotus(43)

Continued

pumilio

Species

divers

climbers

lentic—^vascular

and depositional;

divers

hydrophytes

lentic—vascular

Swimmers;

Lotic—

depositional;

hydrophytes

lentic—vascular

Swimmers; climbers

Lotic—erosional

hydrophytes

lentic—^vascular

Swimmers; Lotic—erosional and depositional; climbers

Swimmers; climbers

Lentic—organic litter, moss

hydrophytes

Swimmers; divers;

depositional;

Lotic—

Lentic—littoral

hydrophytes

lentic—vascular

Swimmers;

Lotic—

depositional;

hydrophytes

lentic—vascular

Swimmers; climbers

Lotic—

Habit

depositional;

Habitat

COLEOPTERA

'Emphasis on trophic relationships

**SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic

*Data listed are for adults unless noted

Order

of species in parentheses)

Taxa (number

Table 21A

North

(piercers)

Predators

East, South

East of Great Plains

Predators

Widespread

Widespread

Widespread

Florida

Widespread

Widespread

Distribution

American

(piercers)

(piercers)

Predators

(piercers)

Predators

(piercers)

Predators

piercers

Predators-

(piercers)

Predators

(piercers)

Predators

Trophic Relationships

10

SB

UM

7.9

M

Ecological

5

5

(continued)

3463, 6804

3463, 6804

3463, 6804, 863

3417, 3460

3463, 5068, 6804, 5856

2933, 2934,

2614, 6804

2607, 2638, 3417, 3463, 6804, 4933

2618, 3417, 3463, 3085

NW MA** References***

Tolerance Values*

) )))) )

00

00

Family

felipi

Psychopomporus

Rhantus(8)

princeps

Pachydrus

Oreodytes(17)

Neosojtopterus(2)

Neodypeodytes(8)

Neobidessus U)

Nebrioporus(3) {=Deronectes, in part)

calidus

Meridiorhantus

Species fraternus

Genus

Megadytes

Continued

Swimmers;

Habit

climbers

climbers

divers

hydrophytes; depositional

lotic—

Swimmers;

Lentic—vascular

Subterranean

depositional

Swimmers;

Lentic—littoral; lotic—

lentic—littoral

Swimmers; climbers

Lotic—erosional

and depositional;

Swimmers; climbers

Lentic—vascular

hydrophytes

lentic—littoral

Swimmers; climbers

Lotic—erosional

and depositional;

hydrophytes

lentic—vascular

Swimmers;

Lotic—erosional

and depositional:

hydrophytes

lentic—^vascular

and depositional; climbers

Lotic—erosional

Habitat

**SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ***Emphasi$ on trophic relationships

*Data listed are for adults unless noted

Order

of species In parentheses)

Taxa (number

Table 21A

(piercers)

Predators

(piercers)

Predators

(piercers)

Predators

(piercers)

Predators

(piercers)

Predators

(piercers)

Predators

(piercers)

Predators

(piercers)

Predators

Trophic Relationships

Widespread

Southeast

Widespread (primarily West)

Northern

Southwest

West,

Florida to Texas

Widespread

Florida

Distribution

American

North SE

UM

M

Ecological

5

-

3417, 3463, 3464, 5529, 6623, 863

6804

3417,3463

3417, 3463

3463

3463, 6804

3463, 3085

6804

NW MA** References***

Tolerance Values*

) ) ) j > ) i ) ) ) ) ) ) ; ) ) ^ ) ))) ) ) ) ) ) )

o

90 00

Adults

Larvae

Water Beetles

- Burrowing

Noterldae (17)

Family

Uvarus(9)

Thermonectus(6)

Stygoporus

Stictotarsus(23) i=Deronectes, in part)

climbers

lotic—

climbers

hydrophytes Same as larvae

borrowers;

borrowers

Predators

(engulfers)

Generally swimmers; climbers;

COLEOPTERA

gatherers

collectors

Predators

(engulfers),

Generally

Generally

(piercers)

Predators

(piercers)

Predators

(piercers)

Predators

(piercers)

Predators

Trophic Relationships

lentic—vascular

hydrophytes

lentic—^vascular

Swimmers; climbers

Lotic—erosional

and depositional;

depositional

hydrophytes;

Swimmers; divers;

climbers

Swimmers;

Lentic—vascular

Shallow wells

hydrophytes

**SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ***Emphasis on trophic relationships

oregonensis

climbers

lentic—vascular

Swimmers;

Lotic—erosional

divers

Swimmers;

Habit

and depositional;

depositional

Lentic—littoral,

Habitat

lotic—

Species

Sanfilippodytes

Genus

(26)

Continued

*Data listed are for adults unless noted

Order

of species in parentheses)

Taxa(number

Table 21A

North

Widespread

Southwest

Oregon

Southwest

Widespread, but mostly

Widespread

Distribution

American SE

UM

M

Ecological

(continued)

283, 284, 2614, 3179, 3463, 3464, 4599, 6804, 5168

2618, 3463, 6804

3463, 6159

3463

2618

NW MA** References***

Tolerance Values*

Family

Adults

Larvae

Hydroscapha (2)

Suphisellus(7)

Suphis

Notomicrus(2)

Mesonoterus

inflatus

addendus

Species

and margins (including thermal springs)

Lotic—erosional

and margins (including thermal springs)

Lotic—erosional

hydrophytes

lentic—^vascular

depositional;

Lotic—

hydrophytes

Lentic—vascular

lentic—littoral

depositional,

Lotic—

hydrophytes

Lentic—vascular

hydrophytes

Lentic—^vascular

Habitat

North

West(especially Scrapers (bluegreen algae) Southwest) Clingers

Widespread

East, South

South

Florida

East, South

Distribution

American

Scrapers West(especially (bluegreen algae) Southwest)

Trophic Relationships

Clingers

Climbers

Climbers

Burrowers

Climbers

Climbers

Habit

**SE = Southeast, DM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ***Ennphasis on trophic relationships

*Data listed are for adults unless noted

Beetles

(2) - Skiff

Genus

Hydrocanthus(6)

Continued

Hydroscaphidae

Myxophaga

Order

of species in parentheses)

Taxa (number

Table 21A

-

6.9

SE -

UM

M

Ecological

7

2644, 2646,3179, 3409, 3463, 3464, 4599, 5367, 2339

6804

6804

5427, 6804

6804

NW MA** References***

Tolerance Values*

) ) ) ) > ) ; ))) ) V ) ; ) ) ) ) ))) ) ) ) ) )

Ki

06

06

00 U)

QC

Beetles

Featherwing

Ptiliidae (1)-

or Hister Beetles

- Clown Beetles

Histeridae (1)

Loving Beetles

- Minute Mud-

Georissidae (2)

Sphaeriusidae (3)- Minute Bog Beetles

Family Genus

Motschulskium

Neopachylophus

Georissus(2)

Adults

Larvae

Sphaerius(3)

Continued

sinuatocolle

suldfrons

Species

margins (semiaquatic)

Intertidal beach

Intertidal

wrack

Borrowers

Borrowers

(sand)

Borrowers

Borrowers; climbers

Lentic and lotic

Lotic margins

Borrowers; climbers

margins (semiaquatic)

Habit

Lentic and lotic

Habitat

COLEOPTERA

**SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ***Emphasis on trophic relationships

*Data listed are for adults unless noted

Polyphaga

Order

of species in parentheses)

Taxa (number

Table 21A

Scrapers (fungi)

Predators

Scrapers (algae)

Scrapers (algae)

Scrapers (algae)

Trophic Relationships

California coast

California coast

Northwest

Midwest,

Washington

Texas, Southern California,

Washington

Texas, Southern California,

Distribution

American

North

SE

UM

M

Ecological 8

8

(continued)

1732, 6459

2340

3282, 6459

3282

701,3463

NW MA** References***

Tolerance Values*

Adults

Larvae

Beetles

Hydrophilidae (183)-Water Scavenger

Beetles

(26) - Water Scavenger

Hydrochidae

Beetles

(43)- Water Scavenger

Helophoridae

Shore Beetles

(4)- Hooded

Epimetopidae

Family Genus

Anacaena (3)

Ametor(2)

Hydrochus(26)

Helophorus(43)

Epimetopus(4)

Continued

Species

gatherers

swimmers

American

depositional (detritus, fine sediments)

Mexico

East and West

Coasts, Illinois, Indiana, New

Burrowers

(silt)

West

Widespread

Widespread

Southwest

Distribution

Lentic—littoral;

Clingers

Generally collectors—

Generally divers;

Generally predators (engulfers)

herbivores

Shredders—

herbivores

Shredders—

Trophic Relationships

North

lotic—

depositional

lotic—

Lentic—littoral;

Same as larvae

depositional

lotic—

Generally climbers

hydrophytes;

Climbers

Climbers

Habit

Lentic—vascular

especially margins

erosional,

Lentic and lotic

especially margins

erosional,

Lentic and lotic

Habitat

**SE = Southeast, DM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ***Emphasis on trophic relationships

*Data listed are for adults unless noted

Order

of species in parentheses)

Taxa (number

Table 21A

7.9

SE

-

UM

-

M

5

5

-

NW

Ecological

3388, 3389, 3463

5586

367, 2737,3179, 3463, 3464, 4599, 5949, 6139, 6205

3463

1671,3388, 3389, 5089,6328

6804

MA** References***

Tolerance Values*

) ) ) ) > ) ) ))) ) ) ) ; ) ) ) ) )) ) ) ) > ) )

4^

00

00

00 00 Ul

Family

Lentic—littoral

Same

Enochrus-Adults

Piercers—

COLEOPTERA

**SE = Sootheast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ***Emphasis on trophic relationships

gatherers Same

herbivores

Collector—

climbers

depositional

Burrowers-

Swimmers; divers;

Lentic—littoral;

sprawlers

gravel)

depositional lotic—

Borrowers

(sand and

Lentic—littoral;

gravel)

depositional

lotic—

Borrowers (sand and

Lentic—littoral; lotic—

depositional

lotic—

Enochrus-larvae

Enochrus(25)

Derallus

Cymbiodyta (23)

Crenitulus(1)

Crenitis(M)

altus

Lentic—littoral;

Chaetarthria (14)

Climbers

Collectors—

gatherers

borrowers

shredders

gatherers;

collectors—

herbivores;

Piercers—

Clingers;

I otic—

Marine intertidal

Berosus(24)

Cercyon

Habit

climbers

Habitat

depositional

Species Swimmers: divers;

Genus

Trophic Relationships

Lentic—littoral;

Continued

"Data listed are for adults unless noted

Order

parentheses)

Taxa (number of species in

Table 21A

North

Widespread

Coasts

East and Golf

Widespread

Coast

East and Pacific

Widespread

Pacific Coast

Widespread

American Distribution

8.5

SE

UM

M

5009

(continued)

280, 3388, 3389,

3463

3463

3463

4618

280, 5009, 6804

Ecological NW MA** References***

Tolerance Values*

Family Genus

Adults

Larvae

Laccobius(24)

Adults

Larvae

Hydrophilus(3)

Hydrochara (9)

Climbers;

climbers

depositional divers; climbers

depositional

**SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ***Emphasis on trophic reiationships

Swimmers;

Lentic—littoral; lotic—

lotic—

Swimmers; divers;

clingers; sprawlers Lentic—littoral;

Lentic—littoral

herbivores

Piercers—

herbivores

gatherers; piercers—

Collectors—

(engulfers)

Predators

fusdpes

Hydrobius

climbers

depositional

lotic—

Swimmers; divers;

casta

Hydrobiomorpha Lentic—littoral;

East

Widespread

Widespread

Widespread

South

Southwest

exilis

Hemiosus

Southeast

Distribution East, Southwest

Widespread

Relationships

bifidus

Habit

North

American

Helocombus

larvalis

Habitat

Trophic

Helochares(3)

Helobata

Continued

*Data listed are for adults unless noted

Order

of species in parentheses)

Taxa (number

Table 21A

SB

UM

1.9

M

Ecological

280, 4618

280, 4040, 6160, 6622, 6838

3817, 6804

282, 3388, 3389

6804

NW MA** References***

Tolerance Values*

) ) ) ))) ))) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) \

as

00 00

Swimmers; divers;

Adults

borrowers

Aleochara

(53)*

Marine beaches

rocky coasts Clingers

COLEOPTERA

Predators (fly eggs)

herbivores

littoral, marine Intertidal and

shredders—

and lentic—

gatherers;

collectors—

climbers;

beaches

Predators

herbivores

gatherers; piercers—

Collectors—

freshwater lotic

Rove Beetles

West Coast

-

-

8

5

5

5

NW

10

5

-

MA**

Ecological

863

280, 2718, 5009, 6806,6328,2187,

5585

3463, 6328

References***

6459

(continued)

910, 3463,4118, 4995, 5287,4341,

5217, 5218, 5219,

-

-

-

M

5220

9.8

6.5

-

UM

Predators

(engulfers);

Generally shorelines and

Staphylinidae

Widespread

East, South

-

SE

Tolerance Values*

(engulfers)

Generally clingers and

climbers

Climbers

depositional

lotic—

Lentic—littoral;

(wood)

depositional

lotic—

SE = Southeast UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic Emphasis on trophic relationships Not all species in the family or genus are aquatic

***

*★

gravel)

depositional Clingers

(sand and

lotic—

Lentic—littoral;

Borrowers

Lentic—littoral;

Larvae

Tropisternus(14)

Sperchopsis tesselata

Distribution East

Habit

Widespread (primarily South)

Habitat

North American

Limnohydrobius(2)

Species

Trophic Relationships

Paracymus(15)

Genus

(478)**** -

Family

Continued

Data listed are for adults unless noted

Order

of species in parentheses)

Taxa(number

Table 21A

Family Genus

Diglotta (3)

Diaulota (6)

Lotic and

Borrowers

Clingers

Clingers

Clingers

Clingers

Clingers

Clingers

Clingers

Clingers

Habit

**SE = Southeast, DM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ***Emphasis on trophic relationships ****Not all species in the family or genus are aquatic

Beaches, marine (sand)

(rock crevices)

Marine coasts

(leaf litter)

lentic—littoral

Carpelimus(79)

(rock crevices)

Marine coasts

(under beach wrack)

Marine coasts

lentic—littoral

Lotic and

stones)

wrack and

(under beach

Marine coasts

(rock crevices)

Marine coasts

Habitat

Marine coasts

catalinae

depressa

Species

Cafius(13)

Bryothinusa

Bryobiota (2)

Bledius(90)****

Biaraxis

Amblopusa (2)

Continued

*Data listed are for adults unless noted

Order

of species In parentheses)

Taxa (number

Table 21A

Predators

Predators

Predators (fly eggs)

Predators

gatherers (decaying algae)

Collectors—

South

Trophic Relationships

West Coast, New Jersey

California)

(Alaska to Baja

West Coast

Widespread

Gulf Coasts

East, West, and

California Coast

California)

Columbia to

(British

West Coast

Widespread

Florida islands

(Alaska to California)

West Coast

Distribution

American

North

Tolerance Values*

) ) ) )))))) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) )

oe oc 00

so

so

Family Genus

algophila

blumbea

Halobrecta

Heyterota

West Coast

Marine beaches

Borrowers

**SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ***Emphasis on trophic relationships

(sand)

Predators

herbivores

Borrowers

nudus

Philonthus

(Alaska to California)

West Coast

California

Oregon Marine beaches

northern

Coasts,

East and West

(Alaska to California)

West Coast

Florida

(Alaska to California)

Alaska to

borealis

Parambbpusa

COLEOPTERA

West Coast

(Alaska to California)

Clingers

Lentic—littoral

Pontomalota (2)

American

Distribution

Marine coasts

Shredders—

Predators

crustaceans)

insects and

Predators(on

Trophic Relationships

Widespread

Clingers

Clingers

Clingers

Clingers

Clingers

Clingers

Habit

North

Clingers

(rock crevices)

Marine coasts

(rock crevices)

Marine coasts

(on seaweed)

Marine coasts

(under beach wrack)

Marine coasts

(under beach wrack)

Marine coasts

(under litter)

Marine coasts

Habitat

Microbledius(4)

Micralymma (2)

Liparocephalus(2)

crassus

Species

Hadrotes

Giulianium (3)

Continued

*Data listed are for adults unless noted

Order

of species in parentheses)

Taxa (number

Table 21A

Toierance Values*

(continued)

Ecological

3 )))) 3 > ) ) )) )) 3 ) ) ) ))) ) )) ) ) )

Flower Beetles

Melyridae (3) - Soft-Winged

Family

Genus

Endeodes(3)

Treopalpus

Thinusa (2)

Thinopinus

Thinobius(24)

Tarphiota (2)

Marine beaches

Clingers

Borrowers?

Borrowers

Climbers

Clingers

**SE = Southeast, DM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ***Emphasis on trophic relationships ****Not all species in the family or genus are aquatic

lithotarinus

(sand)

Marine beaches

(sand)

Marine beaches

lentic—littoral

Lotic and

(sand and wrack)

Marine beaches

surface

Skaters

(eject oils)

Lotic and lentic—littoral

Stenus(167)

California to

California

Washington to

West Coast

(Alaska to California)

Predators

(engulfers)

Columbia

British

West coast

Predators

Widespread

(Alaska to California)

West Coast

Widespread

Widespread

Widespread

Distribution

American

North

(engulfers of Amphipoda)

Predators

Predators

Simuliidae)

Predators

(engulfers of

Trophic Relationships

Climbers

Clingers

Habit

(runners)

Lentic—littoral

Habitat

Lotic—erosional

pictus

Species

Psephidonus (12)****

(3)

Psamathobledius

Continued

*Data listed are for adults unless noted

Order

of species in parentheses)

Taxa (number

Table 21A

SE

UM

Wl

Ecological

6459

523, 3463, 3843,

1183,4995,4996

1347, 5943

NW MA** References***

Tolerance Values*

) ) ) ))) ) )) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) )

00 vc o

00 vo

Beetles

(9)- Darkling

Tenebrionidae

Beetles

Waisted Bark

- Narrow-

Salpingidae (3)

Family Genus

Epantlus

Phaleria (8)

Aegialites(3)

Adults

Larvae

Continued

obscurus

Species

beaches

Intertidal

beaches

Intertidal

Borrowers

Borrowers

COLEOPTERA

North

coasts

Pacific coast

gatherers Scrapers;

gatherers

collectors—

Pacific, Gulf, and Atlantic

collectors—

Pacific Coast

Pacific Coast

Pacific Coast

Distribution

American

Scrapers;

herbivores

shredders—

Predators

(engulfers, especially mites);

Clingers (rock crevice dwellers)

Marine rocky coasts

beaches

Predators

(engulfers)

Clingers (rock crevice dwellers),

Marine beaches

(engulfers)

Predators

Trophic Relationships

and intertidal

beaches

Clingers (rock crevice dwellers),

Marine beaches

and intertidal

Habit

Habitat

''*SE = Southeast, DM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ***Emphasis on trophic relationships

*Data listed are for adults unless noted

Order

of species in parentheses)

Taxa(number

Table 21A

SE

UM

M

Ecological

(continued)

1732, 6459

1732, 6459

2

3463, 5613, 5817, 6131,4776

NW MA** References***

Tolerance Values*

Adults

Limnebius(13)

Hydraena (29)

(7)

Gymnochthebius

American

swimmers

margins);

**Data listed are for larvae

**SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ***Emphasis on trophic relationships

(especially margins)

Clingers; climbers

Lotic—erosional; lentic—littoral

(especially margins)

Clingers; climbers

Lotic—erosional;

climbers

Clingers;

crevices)

rock

cobbles,

lentic—littoral

Lotic—erosional

(especially margins)

Lotic—erosional

hydrophytes Generally clingers (logs,

(primarily semiaquatic)

vascular

(runners)

lentic—

emergent

(or stream

Generally clingers;

Lotic—erosional

gatherers

collectors-

Scrapers;

(engulfers)

Widespread

Widespread

South

Northeast,

Distribution

Ecological NW MA** References***

4617, 4618, 4619

4619

201, 4617, 4618,

4618, 4619, 4621

Larvae

Predators

Trophic Relationships

Moss Beetles

Habit

2450, 3179, 3463,

Habitat

3464, 4599, 4617,

Species

Tolerance Values*

Hydraenldae

Genus

North

(94)- Minute

Family

Continued

*Data listed are for adults unless noted

Order

of species in parentheses)

Taxa(number

Table 21A

) ) ) ) > ) ) )) ) ))) ) ) ) ) ) ) ) ) ) ) ) ))

so bO

00

00 vo

Eubriinae

Adults

Dicranopselaphus

Acneus(4)

Psephenus(7)

Eubrianax

COLEOPTERA

*'^Data listed are for larvae

Clingers

Clingers

Scrapers

Scrapers

Scrapers

Clingers

: Mid-Atlantic

Lotic—erosional

Lotic—erosional

Lotic—erosional

Scrapers

Nonfeeding

Scrapers

Scrapers

Trophic Relationships

Clingers

clingers

oviposit) Lotic—erosional

Ovipositing females are

water to

Clingers

(Females enter

erosional

**SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = ***Emphasis on trophic relationships

variegatus

edwardsii

Lotic and

Larvae

lentic—

erosional

Clingers Clingers

Marine intertidal

Clingers

Habit

Generally lotic

species)

intertidal

and margins(8

Lotic—erosional

Habitat

Pennies

vandykei

Species

and lentic—

Neochthebius

Ochthebius(44)

Genus

Psephenidae (16)-Water

Family

Continued

*Data listed are for adults unless noted

Order

Taxa (number of species in parentheses)

Table 21A

Widespread

Francisco Bay)

(south to San

West Coast

West

East (1 sp.).

West

Pacific Coast

Widespread

Distribution

American

North

25*+

-

SE

4*+

UM

3,5*+

M

Ecological

4*+

4*+

5*+

736

3010

(continued)

732, 1077, 1363, 4226, 4229, 5300, 6426, 3975, 4228,

1044,4227

5518

5420, 2958

2643, 3179, 3463, 3464, 4230, 4599,

743, 745, 2637,

4619

2450,4617,4618,

NW MA** References***

Tolerance Values*

D ))>))) )))) )) 3 J ; ) 3 ) ) ) 3 3 3 3 3 3

J

■1^

)

00

Lotic, cave

Stygoparnus

)

)

)

)

*"^Data listed are for larvae

)

)

)

)

)

)

)

)

)

)

)

)

3.2

4*+

M

)

5

4**

NW

)

5

5*+

MA**

Ecological

5422

2411, 6081

736, 6804

2411, 6081

2634

4599, 5417

3179, 3463, 3464,

671, 672, 1077, 5300, 6804

References***

)

6800

)

)

3179, 3463,3464,

5

5**

UM

4599, 5829, 6044,

Texas

Comal Springs,

Southwest

5.4

4.3*+

SE

Scirtldae (50)

)

Clingers

Clingers

Texas to

Florida, Illinois

Widespread

Northeast

Southwest,

East

Distribution

Climbers

)

North American

Clingers

**SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ***Emphasis on trophic relationships

)

herbivores

shredders—

scrapers;

Generally

herbivores

shredders—

Generally

Scrapers

Trophic Relationships

Tolerance Values*

- Marsh Beetles

springs

Lotic—erosional

hydrophytes

vascular

Postelichus (3) comalensis

Lentic—

Pelonomus

emergent

Lotic—erosional

Helichus (7)

hydrophytes (emergent zone)

Lentic—vascular

Climbers

climbers

lentic—littoral;

Adults (all entries) lotic—erosional

Clingers;

Clingers

Habit

Generally

erosional

lentic—

Lotic and

Habitat

Borrowers

obscurus

Species

Terrestrial

Dryops (2)

Ectopria (3)

Genus

Lan/ae

Water Beetles

Dryopidae (14) - Long-Toed

Family

Continued

*Data listed are for adults unless noted

Order

of species in parentheses)

Taxa (number

Table 21A

)

vo Ul

00

Adults

Larvae (all entries)

Family Genus

Terrestrial

Scirtes(6)

Climbers

Shredders—

piercers—

hydrophytes

herbivores

herbivores;

vascular

Lentic—floating

COLEOPTERA

**SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ***Emphasis on trophic relationships ****The genus Cyphon is no longer valid and has been divided into 4 genera (see Page 849 for further details).

robustus

Widespread

Northeast

East

Sarabandus

Florida, Texas Tree holes

Ora (3))

Northeast

East

Widespread

Widespread

Distribution

American

North

Prionocyphon (2)

Microcara

Tree holes; seeps

Sacodes(3) {=Flavohelodes)

bogs); tree holes

marshes and

litter; including

(sediments, leaf

Lentic—littoral

(possibly a few semiaquatic)

herbivores

herbivores; piercers—

hydrophytes

gatherers; shredders—

vascular

emergent

Generally scrapers; collectors—

Generally climbers; sprawlers

Trophic Relationships

Generally

Habit

lentic—

Habitat

Tree holes; seeps (including mineral springs)

explanata

Species

Elodes(7)

Cyphon (27)*

Continued

*Data listed are for adults unless noted

Order

parentheses)

of species in

Taxa (number

Table 21A

Ecological

(continued)

399, 3285

2188, 3463

3463

3463

NW MA** References***

Tolerance Values*

3 D ) >))) ) J )) )))) ) ))) ) ) 3 ) ) ))

GC

Adults

Larvae

- Riffle Beetles

Elmidae (102)

Family

variegatus

wawona

Ancyronyx

Atractelmis

Species

dispar

Genus

Ampumlxis

Continued

hydrophytes

Lotic—erosional

Clingers

Clingers; sprawlers

Mid-Atlantic

Lotic—erosional

and depositional (detritus)

borrowers

Clingers;

scrapers

gatherers;

collectors—

Generally

herbivores

shredders—

hydrophytes Lotic—erosional

gatherers; scrapers;

collectors—

few climbers

lentic vascular

and lentic—

erosional; few

Generally

Clingers;

herbivores

shredders—

lentic vascular

Generally lotic

gatherers; scrapers;

erosional; few

Generally collectors—

Clingers; few climbers

Trophic Relationships

Generally lotic

Habit

and lentic—

Habitat

**SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = ***Emphasis on trophic relationships

*Data listed are for adults unless noted

Order

of species In parentheses)

Taxa (number

Table 21A

Washington

California to

East, Southeast

West

Distribution

American

North

6.9

.

SE

6

.

UM

4

.

M

Ecological

4

5422

5544

201,202, 737, 739, 743, 746, 747, 793,3179, 3463, 3464, 4073, 4335,4599, 5272, 6454, 6062, 6328, 5416, 6296

NW MA** References***

Tolerance Values*

) ) ) ))) ) ) ) ) ) ) ) > ))) ) ) ))) ) ) ) )

0\

00 ve

Family Genus

marroni

Huieechius

Lara (2)

ferrugineus

Hexacylloepus

Heterlimnius(2)

Heterelmis(5)

dietrichi

Bryelmis Barr (Coleoptera: Elmidae: Elminae), is a new genus of riffle beetle from the Pacific Northwest(See reference 6913).

COLEOPTERA

*★ **

(wood)

(continued)

(in wood)

including semiaquatic)

5697

167, 168, 762, 4

West(montane) detritivores burrowers

6328, 6338

Clingers;

SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic Emphasis on trophic relationships

***

**

Shredders—

4

Lotic—erosional

.

735, 762

1077, 5544

762

(wood debris,

.

6

Ecological References***

740

.

4

5

5

4

4

NW MA**

Arizona

Southwest

West

Southwest

3.2

.

.

M

Clingers

Clingers

Clingers

Clingers

6

.

.

UM

Tolerance Values*

Lotic—erosional

and wood)

(cobbles, gravel,

Lotic—erosional

gravel)

(cobbles and

Lotic—erosional

(cobbles and wood)

Lotic—erosional

submerged roots)

(wood debris and

Clingers; climbers

Lotic—erosional

and depositional

(submerged macrophytes) Southeast

6.4

Widespread

Clingers; climbers

Lentic and

.

SE

lotic— erosional

West

West

Distribution

.

Dubiraphia (11)

Gonielmis

North American

Arizona

Clingers

Clingers

Habit

Trophic Relationships

Clingers

submerged roots)

(cobbles and

Lotic—erosional

liverworts

Lotic—on

Habitat

Lotic—erosional

addenda

Species

Cylloepus(2)

Cleptelmis

Bryelmis(3)****

Continued

*Data listed are for adults unless noted

Order

of species in parentheses)

Taxa (number

Table 21A

3 D ) > 3 )} 3 3 ) 3 ))3 J ) ) 3 ) ) ) 3 ) ) 3 3 3

)

)

00

00

Family Genus

Oulimnius(2)

Ordobrevia

Lotic—erosional

gravel)

(cobbles and

Lotic—erosional

gravel)

(cobbles and

Lotic—erosional

)

)

)

)

I

)

1

)

)

)

)

)

Clingers

Clingers

)

)

)

gatherers(adults)

(sediments and detritus)

Scrapers (larvae); collectors—

Clingers

Clingers

Clingers

Clingers

gatherers

Collectors—

Trophic Relationships

and depositional

Lotic—erosional

gravel)

(cobbles and

Lotic—erosional

gravel)

(cobbles and

Lotic—erosional

(cobbles and gravel)

borrowers

Clingers; climbers;

Lotic—erosional

Clingers

Clingers

Habit

and depositional

(wood debris)

and depositional

Lotic—erosional

gravel)

(cobbles and

Lotic—erosional

Habitat

"*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ""♦Emphasis on trophic relationships

nubifera

caesa

Neoelmis

Optioservus(15)

boeseli

glabratus

Species

Neocylloepus

Narpus(3)

Microcylloepus(6)

Macronychus

Macrelmis(3)

Continued

"Data listed are for adults unless noted

Order

of species in parentheses)

Taxa (number

Table 21A

North

)

)

East, Southeast

West

Widespread

Southwest

Texas, Arizona

West

Widespread

East

Southwest

Distribution

American

)

1.8

2.7

2.1

4.7

SB

)

'

■

4

3

4

UM

"

)

'

2.75

■

2.9

M

Ecological

4

4

4

4

2

>

"

4

)

736

2334

)

)

1077, 3271, 5389, 5390, 6453, 6328,

288, 762, 979,

734, 736

762, 6328

5415

1371,5544

NW MA** References***

Tolerance Values*

)

vo

vo

00

Xenelmis

Clingers

Lotic—erosional (coarse sediments and

Generally clingers; borrowers

American

California, Oregon

West

Arizona

Widespread

California

Texas

Distribution

5.4

SE

UM

M

NW

MA**

Ecological

6328

742

3010

736, 1077, 1752, 5388, 5389, 5390,

References***

Anchycteis

Terrestrial (near

Adults

Borrowers

COLEOPTERA

**SE = Southeast, DM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ***Emphasis on trophic relationships

and depositional

Lotic—erosional

leaf litter)

lotic margins in

Generally lotic— erosional and depositional

Larvae (all entries)

(chewers)

herbivores

Shredders—

herbivores

detritlvores and

shredders—

(continued)

3463, 3505, 6804

2175,2176, 3461,

gravel)

(cobbles and

Lotic—erosional

Generally

Clingers

Lotic—erosional

Clingers

gatherers

adults climbers

semiaquatic, lotic margins

detritus)

collectors—

climbers;

adults

Scrapers;

Trophic Relationships

2868

velutina

Habit

Lotic—erosional; Cllngers;

Habitat

Tolerance Values*

Ptilodactylidae

Zaitzevia (2)

sandersoni

nigra

Rhizelmis

Stenelmis(33)

davicornus

Species

Phanocerus

Genus

North

(2)- ToeWinged Beetles

Family

Continued

*Data listed are for adults unless noted

Order

of species In parentheses)

Taxa (number

Table 21A

On

streamside

vegetation

lotic margins in leaf litter)

Lutrochidae (3)

Beetles

- Travertine

Adults

Larvae

Throsdnus(3)

Borrowers

*'^Data listed are for larvae

**SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ***Emphasis on trophic relationships

marine intertidal

Beach zone

marine intertidal

gatherers?

Borrowers

Collectors—

Beach zone

borrowers

Loving Beetles

Limnichidae (3)

Generally clingers or

large rocks

debris near

Terrestrial (near

Clingers on organic

Clingers

Habit

Adults

and depositional

Lotic—erosional

Habitat

Lotic—erosional

scutellaris

bicolor

Species

Larvae

Stenocolus

Anchytarsus

Genus

- Minute Marsh-

Beetles

- Forest Stream

Eulichadidae (1)

Family

Continued

*Data listed are for adults unless noted

Order

Taxa (number of species in parentheses)

Table 21A

American

Texas, southern California

Collectors—

California

Texas, southern

California

Northern

East

Distribution

gatherers?

gatherers?

collectors—

Generally

May not feed

detritivores

Shredders—

(rotting wood)

detritivores

Shredders—

Trophic Relationships

North

3,8*-^

SE .

UM .

M

Ecological .

.

743, 1747, 3463, 3464, 5414, 5418

3463

3463

743, 1747, 3463, 3464, 5414, 5419

5421

NW MA** References***

Tolerance Values*

) ) ) ) ) ) ) )) ) ) )) ) ) ) ) ) ) ))) ) )) )

VO o

vo

hydrophyte

Agasides

vascular

zones)

Adults

from

in floating and submerged

COLEOPTERA

**SE = Southeast, DM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ***Emphasis on trophic relationships ****Not all species in the family or genus are aquatic

biological control of Alternanthera

Introduced for

herbivores

Shredders—

On

surfaces)

leaf

Sprawlers (on floating

herbivores

Shredders—

Shreddersherbivores

(macroalgae)

herbivores

(wood):—

detritivores

Shredders—

(macroalgae)

herbivores

(wood);—

detritivores

Shredders—

Trophic Relationships

Alternanthera

floating leaves)

surface of

hydrophytes (generally

Lentic—vascular

(below surface

tissues)

Clingers (may obtain air directly

Lentic—vascular

Larvae

hydrophytes

hydrophytes

Leaf Beetles

Generally clingers; sprawlers

Clingers

Clingers

Habit

Generally

(especially mineral springs)

Lotic—erosional,

(especially mineral springs)

Lotic—erosional,

Habitat

lentic—^vascular

hygrophila

Species

Chrysomelidae

Adults

Larvae

Lutrochus(3)

Genus

(113)**** -

Family

Continued

*Data listed are for adults unless noted

Order

of species In parentheses)

Taxa (number

Table 21A

North

South

East, Southwest

East, Southwest

Distribution

American

SE

UM

2.9

2.9

M

Ecological

5035

(continued)

229, 230,231,

3179, 3463, 3464,

736, 744

736, 744

NW MA** References***

Tolerance Values*

) )) > ) )) ) ))))) 3 ) ) )))) ) ) ) )) 3

VO

Family

American

harlsii

phellandrii

Poedlocera

Prasocuris

**SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ***Emphasis on trophic relationships ****Not all species in the family or genus are aquatic

hydrophytes

vascular

variety of

On a wide

Sdrpus

On Carex and

hydrophytes

vascular

variety of

On a wide

Northeast

Northeast

Widespread

Mountains

Plateumaris(17)

East of Rocky

On

biological control of Lythrum Potamogeton

North, East

East

Widespread

Widespread

Distribution

Neohaemonia (4)

Introduced for

Trophic Relationships

On Lythrum

or Caltha

On Ranunculus

hydrophytes

vascular

variety of

On a wide

hydrophytes

vascular

Habit

North

Neogalerucella (2) {=Galerucella, in part)

Hydrothassa (4)

nymphaeae

On a wide

Donada (32)

Galerucella

Amaranthaceae

variety of

On

Habitat

Disonycha

Species

(36)****

Continued

*Data listed are for adults unless noted

Order

of species in parentheses)

Taxa (number

Table 21A

NW MA**

Tolerance Values*

Ecological

229

230

228, 630, 2680

6264

630, 2680, 2681, 3758, 3944

References***

))) ) ) ) ) )) ) ) )) ) ) ) ') ) ) ) ) ) ) ) ) )

o

©

Generally clingers;

Habit

Shredders—

Widespread

East

Distribution

SE

UM

M

Ecological NW MA** References*** 1

Adults

Larvae

(265)**" -

Lentic—littoral

Bagous(33)

COLEOPTERA

**SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA ; '"Emphasis on trophic relationships "**Not all species in the family or genus are aquatic

Sprawlers; clingers

Generally clingers and sprawlers

(in stems)

burrowers

climbers; sprawlers (on leaf surfaces).

■ Mid-Atlantic

hydrophytes)

vascular

(floating

Lentic—littoral

(emergent and floating vascular hydrophytes)

Auleutes

(12)""

Same as larvae

(most semiaquatic)

hydrophytes

lentic—vascular

Generally

herbivores

Shredders—

(chewers and miners)

herbivores

Widespread

3944

(continued)

217

Nymphaeaceae

On

Habitat

True Weevils

Species

North American

1520, 3463, 3464, 3944, 5878, 5965,

Pyrrhalta {&)****

Genus

Trophic Relationships

Tolerance Values*

Curcuiionidae

Family

Continued

'Data listed are for adults unless noted

Order

of species In parentheses)

Taxa(number

Table 21A

))) ) ) ) ) ) ) )) ) ))) ) ) ) J ) ) )) ) ) )

1

VO

o ■c>.

)

Family

salvinae

mimrticus

marinus

fuciola

lecontei

angulicollis

Cyrtobagous

Dryotribus

Elassoptes

Emphyastes

Euhrychiopsis

Gononotus

Species electus

Genus

Brachybamus

Continued

(decaying Fucus)

Marine beaches

Potamogeton)

and

)

)

)

)

)

)

)

)

)

)

)

)

(under debris)

Clingers

)

)

herbivores

hydrophytes. Myriophyllum

Shredders—

Climbers:

clingers

Lentic—littoral

(vascular

macroalgae)

under

Shredders—

detritivores

Shredders—

detritivores

Shredders—

molesta

control S.

Introduced to

)

Trophic Relationships

detritivores

Clingers

Clingers

Clingers

Sprawlers; clingers

Habit

marine (sand.

Beach zone

on wood

Marine beaches

on wood

Marine beaches

On Salvinia

Eleocharis)

hydrophytes. especially

vascular

(emergent

Lentic—littoral

Habitat

**SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ***Emphasis on trophic relationships

*Data listed are for adults unless noted

Order

of species in parentheses)

Taxa (number

Table 21A

)

Florida

)

East, Midwest

Pacific Coast

Columbia

British

California to

Florida coasts

South Carolina,

East

Distribution

American

North

)

SE

1

UM

M

)

)

NW MA**

Tolerance Values*

Ecological

)

175

)

)

1203, 2460, 4339

1200, 1202, 1201,

176

175

4406

5878

References***

)

SO o 'M

Family

Neochetina (2)

Neobagoidus

Marine beaches

herbivores

herbivores

Eichhomla)

hydrophytes:

vascular

Shredders-

Climbers;

clingers

Lentic—littoral

(emergent

Lachnanthes)

hydrophytes:

Shredders-

Climbers;

clingers

Lentic—littoral

Clingers

Clingers

Clingers

(vascular

(on wood)

Marine beaches

(on wood)

Marine beaches

(on wood)

COLEOPTERA

**SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ***Emphasis on trophic relationships ****Not all species in the family or genus are aquatic

carlsoni

liUoralis

Macrorhyncolus

Mesites(2)

lineahs

Lentic—littoral

Lixus(69)**

Macrancylus

hydrophytes (emergent and floating zones)

(emergent and floating vascular hydrophytes)

Lentic—vascular

rice)

hydrophytes:

and submerged foliage of rice)

herbivores (roots

vascular

Shredders—

Climbers;

clingers

Trophic Relationships

Lentic—littoral

Habit

(submerged

Habitat

Listronotus

Species

i+Lixellus) (81)****

Lissorhoptrus(6)

Continued

*Data listed are for adults unless noted

Order

of species in parentheses)

Taxa (number

Table 21A

Florida to Texas

Florida

coasts

East and Gulf

California Coast

coasts

East and Gulf

Widespread

Widespread

East

Distribution

American

North

Ecological

4406

4406

(continued)

3944, 175

3193, 6064

NW MA** References***

Tolerance Values*

) ) ) ) ) ) ) ) ) ) ) ) ) )) ) ) )) ) ) ) ) ) ) )

vo

Family Genus

stratiotes

hydrophytes:

herbivores

Pelenomus(13)

Paralichus

Onychylis(6)

herbivores

herbivores

**SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ***Emphasis on trophic relationships ****Not all species In the family or genus are aquatic

Shredders—

Climbers;

clingers

Lentic—littoral

(emergent and floating vascular hydrophytes: Polygonum)

(under wood and litter)

Marine beaches

Nuphar)

Pontederia and

Shredders—

Climbers;

clingers

Lentic—littoral

(emergent and floating vascular hydrophytes:

hydrophytes: Scirpus and probably others)

vascular

Climbers:

clingers

Lentic—littoral

(emergent

hydrophytes: Typha)

vascular

Shredders—

Lentic—littoral

(emergent

Pistia)

Introduced to control P.

Trophic Relationships

Lentic—littoral

Habit

(vascular

Habitat

Notiodes

ninyops

affinis

Species

(12)****

Notaris(2)

Neohydronomus

Continued

*Data listed are for adults unless noted

Order

of species in parentheses)

Taxa (number

Table 21A

1 ") ) > ))) 1 1 ))))) r) 1

o

East, West, Northern

Florida

East

Widespread

Widespread

Southern

Distribution

American

North SE

1

UM

M

Ecological

175

175

4406

1 )

NW MA** References***

Tolerance Values*

vo

o -4

Family

Lentic—littoral

COLEOPTERA

**SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ***Emphasis on trophic relationships ****Not all species in the family or genus are aquatic

duckweeds)

Lentic—littoral

(floating vascular hydrophytes:

Tanysphyrus

Azola)

(floating vascular hydrophytes:

Climbers

(Lemna)

herbivores

Shredders—

herbivores

Shredders—

herbivores

hydrophytes: Polygonum)

Shredders—

Climbers;

clingers

Shredders

herbivores

Shredders—

Trophic Relationships

Lentic—littoral

Borrowers

Clingers

Habit

(vascular

(in marine timber)

Marine beaches

hydrophytes: Lepidium, Ptilmnium)

(vascular

Lentic—littoral

(emergent and floating vascular hydrophytes: Myriophyllum)

Lentic—littoral

Habitat

(2)*.

(3)****

Steremnius

Stenopelmus rufinasus

spadix

Pselactus

Rhinoncus(7)

setosus

Pnigodes

Species leucogaster

Genus

Phytobius

Continued

*Data listed are for adults unless noted

Order

of species in parentheses)

Taxa (number

Table 21A

Widespread

East, West

Widespread

Widespread

East Coast

South, West

Widespread

Distribution

American

North SE

UM

M

Ecological

5878

2675

5878

1202

(continued)

NW MA** References***

Tolerance Values*

) ) J ) J ) ) ) ) )J ) ) ) ))) > ))) J 3 3 ) )

Family Genus

Tournotaris(2)

Thalasselephas

Continued

testaceus

Species

American

herbivores

hydrophytes: Typha)

vascular

Shredders—

Lentic—littoral

(emergent

North, West

California

Distribution Southern

Trophic Relationships

Marine beach

Habit

zone (sand and rock crevices)

Habitat

North SE

UM

M

Ecological 176

NW MA** References***

Tolerance Values*

**SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ***Emphasis on trophic relationships

*Data listed are for adults unless noted

The following curculionid genera are associated with littoral or wetland plants, at least in part: Amalorrhynchus, Barilepton, Barnius, Cyllndridia, Dirabius, Eudodminius, Haplostethops, Microcholus, Notiodes, Peracalles, Tyloderma.

The following list includes chrysomelid genera that feed on littoral and wetland plants: Donadella, Hippuriphila, Lysathia, Neocrepidodera, Phaedon, Plateumaris, Pseudolampsis.

Other staphylinid genera inhabiting the banks of streams or lake margins (periaquatic): Abdiunguis, Acidota, Acylophorus, Adota, Anaquedius, Arpedium, Artochia, Atanygnathus, Bamona, Beeria, Boreaphilus, Brachygluya, Brathinus, Coryphiomorphus, Coryphium, Cylindrarctus, Derops, Dianous, Diochus, Empelus, Erichsonius, Eucnecosum, Geodromicus, Gnathoryphium, Gyrohypnus, Gyronycha, Gnypeta, Haida, Hemiquedius, Hesperolinus, Eloloboreaphilus, Hydrosmecta, Hydrosmectina, Ischnosoma, Kalissus, Lathrobium, Lesteva, Lithocharodes, Microedus, Micropeplus, Mycetoporus, Myllaena, Myrmecopora, Neobisnius, Nisaxis, Nitidotahdnus, Nordenskioldia, Ocdephelinus, Ochthephilus, Olophrum, Orobanus, Orus, Paederus, Phlaeopterus, Phiionthus, Pselaptrichus, Riechenbachia, Rybaxis, Scopaeus, Staphylinus, Stenistodermus, Subhaida, Tachyporus, Tachyusa, Tasgius, Tetrascapha, Thinodromus, Tychobythinus, Unamis, Vellica, Vicelva.

Tetraleucus.

Other carabid genera associated with lentic and lotic margins: Acupalpus, Anchonoderus, Anisodactylus, Antrichis, Ardistomis, Asphidum, Aspidoglossa, Badister, Bembidion, Brachinus, Broscodera, Calybe, Chlaenius, Clivina, Diplochaetus, Dereylinus, Diplocheila, Diplous, Euphorticus, Evolenes, Geopinus, Lachnocreptis, Lachnophorus, Lophoglossus, Loxandrus, Omophron, Oodes, Oodinus, Oxycrepis, Paratachys, Patrobus, Pelophila, Pericompsus, Phyrpeus, Platidiolus, Pogonodaptus, Platypatrobus, Pterostichus, Scarites, Schizogenius,Semiandistomis, Stenocrepis, Stenolophus, Tachys,

Order

of species in parentheses)

Taxa (number

Table 21A

) ))) > ) ) ) ) ) ) ))) 1 ) ) > ) ))) > )))

vo o 00

AQUATIC HYMENOPTERA Andrew M. R. Bennett

Ccmadifm National Collection ofInsects, Arachnids and Nematodes

Ottawa, Ontario, Canada INTRODUCTION

In an earlier review of North American aquatic Hymenoptera (Hagen 1956), only species with adults known or suspected to either dive or crawl beneath the water surface to parasitize or to obtain their hosts were included. The advantage ofthis definition is that there is little ambiguity regarding what constitutes an

aquatic species. The disadvantage is that the utility of the list is limited to use by those who actually witness

aquatic behavior(only observed in 12 North American species in 11 genera). Alternatively, the 3rd edition of the current volume followed the definition of Burghele

(1959) who included all species parasitizing aquatic stages of insects. This broader definition was more inclusive, but suffered from lack ofprecision regarding what constitutes an aquatic stage of a host because of

subjectivity regarding the exact time when parasitism began. For example, should parasitoids of pre-pupal Dytiscidae that crawl out of the water to pupate be included? In order to remove the ambiguity regarding host aquatic stages, an even broader definition,

following Hedqvist (1967) has been adopted in this edition. This includes all Hymenoptera that are

parasitoids of aquatic invertebrates, regardless of the location of the parasitized host stage. This definition may be faulted for being too broad (by including some species that parasitize terrestrial life stages of aquatic species), but it is likely the most useful to those interested in collecting and/or rearing Hymenoptera around water. The 11 genera that have witnessed accounts of adult aquatic behavior in North America are indicated in the table of species.

All aquatic Hymenoptera parasitize immature host stages except for the pompilid Anoplius depresslpes Banks (Vespoidea). This wasp attacks adult pisaurid spiders (genus Dolomedes) that run over and dive under the surface of the water and may stay

submerged for some time. Since A. depressipes can crawl into the water and run on the bottom, it may

sting the spiders underwater before transporting them to nests made in the bank (Evans 1949; Evans and Yoshimoto 1962; Roble 1985).

Aquatic Hymenoptera in North America that parasitize immature host stages belong to the superfamilies Ichneumonoidea, Platygastroidea, Diaprioidea, Chalcidoidea, and Cynipoidea. Most are internal parasitoids of aquatic immatures that are usually found in plant tissues. The few external parasitoid species oviposit on larvae, pre-pupae,and pupae that are either in terrestrial cocoons, or plant mines. Within the Ichneumonoidea, aquatic braconids are known from the subfamilies Alysiinae, Braconinae, Opiinae, and Microgastrinae (although the latter has no aquatic species in North America). The Alysiinae

and Opiinae are all endoparasitoids ofcyclorrhaphous Diptera(Wharton et al. 1997)and several genera have been associated with Hydrellia spp. (Ephydridae) (Berg 1949). A species of Bracon (Braconinae) has been reared as an ectoparasitoid from stem-boring Lepidoptera in water reeds(Frohne 1939). Among aquatic ichneumonids, five subfamilies have aquatic species: Agriotypinae, Campopleginae, Cryptinae, Cremastinae, and Phygadueontinae (the latter previously part of Cryptinae). The Agriotypi nae (Agriotypus) comprises 16 Eurasian species that crawl underwater to parasitize pre-pupae and pupae of caddisflies in fast-running streams (Aoyagi and Ishii 1991; Elliot 1982; Bennett 2001). Aquatic campoplegines are not known from North America. Within North America, Apsilops hirtifrons(Ashmead) (Cryptinae) has been witnessed entering the water to parasitize stem-mining Lepidoptera larvae in the Eastern and Midwestern United States (Cushman

1933). Tanychela pilosa Dasch (Cremastinae) parasit izes crambid larvae that form silk retreats on the bot

tom of streams in the West(Resh and Jamieson 1988; Jamieson and Resh 1998) and therefore the females must enter the water to oviposit on hosts. There are

909

910

Chapter 22 Aquatic Hymenoptera

also species in several phygadeuontine genera that parasitize larvae and pre-pupae of host stages that occur in and around the water's edge (e.g., Sulcarius spp. on limnephilid pre-pupae; Medophron spp. on dytiscid pre-pupae and some Bathythrix spp. on gyri-

parent first larval instar(Fig. 22.6) hatches. The larva feeds by sucking in yolk spheres from the host egg, which can be seen in the gut of the second instar (Fig. 22.7). As the yolk cells are digested, the gut of the third instar becomes paler (Fig. 22.8). The gregarious

nid pre-pupae)(Townes 1970). The Platygastridae (Platygastroidea) are parasitoids of insect eggs. Tiphodytes gerriphagus(Marchal) is a widespread species that has been observed crawl

larvae do not attack each other and face in different

ing on submerged vegetation to locate its host gerrid eggs (Matheson and Crosby 1912). There are also several other platygastrid genera that parasitize eggs of tabanids, nepids and a species of Thoronella has been found phoretic as adults on an aeschnid(Carlow 1992), which implies that it parasitizes aeschnid eggs. Within the Diapriidae (Diaprioidea), Trichopria columbiana (Ashmead) parasitizes Hydrellia spp. (Ephydridae), and is a good swimmer (Berg 1949). Other species of Trichopria have been associated with sciomyzid and psephenid pupae. Species oftwo genera of Cynipoidea, both eucoiline figitids, are associated with aquatic habitats. Hexacola hexatoma (Hartig) and Kleidotoma parydrae Beardsley have both been reared from Parydra (Ephydridae) (Deonier and Regensburg 1978; Meyers and Deonier 1993). The remainder of aquatic North American Hymenoptera belong to the Chalcidoidea (Chalcididae, Eulophidae, Mymaridae,Pteromalidae, and Trichogrammatidae). Species of Chalcis (Chalcididae) parasitize eggs of Stratiomyidae laid on emergent vegetation around the water's edge (Boucek and Halstead 1997).

The tiny mymarid wasps (fairyflies) are all inter nal parasitoids of insect eggs. Species of Anagrus, Anaphes, Caraphractus, and Polynema have been reared from eggs of aquatic Coleoptera, Hemiptera, or Odonata. Specimens of the genus Ptilomymar are generally found around water and have been collected in

aquatic emergence traps in the Philippines (Freitag 2004; Huber, pers. comm.), which implies that P. magnificum Yoshimoto from northeastern North America is also aquatic, although no host is known. The biology of Caraphractus cinctus Walker, which is probably the most completely known of any aquatic hymenopteran, is summarized in Figures 22.1-22.10. After mating underwater, on the surface film, or on emergent plants, this species oviposits underwater in dytiscid beetle eggs, either exposed or in plant tissues(Fig. 22.1). The number ofeggs depos ited depends upon the size of the host egg, but up to 55 progeny have been obtained from a single dytiscid egg. In such cases the adults are often micropterous (with reduced wings). After the egg (Fig. 22.2) is deposited, it enlarges (Figs. 22.3-22.5) and the trans

directions if they are in the same host egg (Figs. 22.8 and 22.9). Opaque white areas, which probably repre sent excretory products, can be seen in the terminal larval instar (Fig. 22.9). Wastes are stored as a single mass in the pupal stage(Fig. 22.10). The adults emerge underwater, respire cutaneously, and swim with their wings (Jackson 1958c, 1961b). The female of C. cinc tus probes each egg of Agabus sp. with her ovipositor and usually rejects eggs already parasitized. Arrhenotokous parthenogenesis occurs as in most parasitic Hymenoptera, where an unmated female will produce only male progeny and a mated female can deposit both fertilized and unfertilized eggs. When C cinctus oviposits in small host eggs like Agabus bipustulatus, the proportion of fertilized eggs usually increases when hosts are offered in quick succession; the number of eggs deposited in each host is then reduced to two or one with a resulting sex ratio ofabout 17% males. In host eggs offered to a female at long

intervals, three eggs are deposited, and the resulting sex ratio is usually one male and two females. Under high competition between female parasitoids for a few host eggs, about 47'/o male progeny results(Jackson 1966). Jackson (1961a) was able to obtain four or five generations of C. cinctus from A. bipustulatus reared in an unheated room near a window in Scotland, but by the end of October all full-grown larvae (pre-pupae) entered a diapause state. These diapausing pre-pupae pupated in the spring and emerged as adults from March to May. Interestingly, dytiscid eggs do not diapause but develop throughout the year when the temperature is suitable. Jackson (1961a) experimen tally varied the photoperiod and found that 9 hours of complete darkness induced diapause in C. cinctus lar vae (which were sensitive to extremely low light inten sity); she concluded that in ponds, direct development ofthe parasitoid would occur with long daylight hours combined with faint light at night (twilight) or with a few hours ofdarkness. Temperature was not the decid ing factor in the induction of diapause. Aquatic Trichogrammatidae are all internal parasitoids of insect eggs. Although some species of the large genus Trichogramma can swim with their legs, remain underwater up to 5 days, and have been reported to mate within the submerged host egg, most "aquatic" species will undoubtedly be reared from terrestrial eggs of aquatic insect species. Thus far, no Trichogramma species have been reported as

Chapter 22 Aquatic Hymenoptera

911

Figure 22.7 Figure 22.6

1mm

Figure 22.1

Figure 22.2 Figure 22.3 Figure 22.5 0.1S mm

Figure 22.4

0.6 mm

Figure 22.10 Figure 22.8

Figure 22.9

Figure 22.1 Female Caraphractus cinctus Walker (Mymarldae) ovipositing underwater in an egg of Agabus bipustulatus (L.)(Dytiscidae) laid in a sphagnum leaf. One other host egg is shown attached to the leaf with gelatinous cement (from Jackson 1958b). Figure 22.2 Caraphractus cinctus ovarian eggs (from Jackson 1961b). Figure 22.3 Caraphractus cinctus egg dissected from host 20 minutes after laying (from Jackson 1961b). Figure 22.4 Caraphractus cinctus egg (laid by same female as egg in Fig. 22.3) dissected from another host about 72 hours after laying (from Jackson 1961b). Figure 22.5 Caraphractus cinctus egg of another female about 72 hours after laying; more advanced, showing reduced size and developing embryo. Figure 22.6 Caraphractus cinctus first larval instar dissected from host about 70 hours after laying

(from Jackson 1961b); HG, hindgut; M, mouth; MG, midgut; MU, muscle cells extending from body wall to esophagus; O, esophagus; X, probably sex cells. Figure 22.7 Caraphractus cinctus second larval instar (from Jackson 1961b). Figure 22.8 Egg of Agabus bipustuiatus with twothird stage larvae of Caraphractus cinctus, about 9 days after laying (from Jackson 1961b). Figure 22.9 Two full-grown parasitic larvae of Caraphractus cinctus in host egg. A, discolored spot around the oviposition puncture in shell of Agabus bipustuiatus (from Jackson 1961b). Figure 22.10 Eggs of Agabus bipustuiatus, on leaf of Juncus sp., containing two newly formed pupae of Caraphractus cinctus; the one above is a male, the one below a female (from Jackson 1961b).

912

Chapter 22 Aquatic Hymenoptera

parasitoids of submerged host eggs. However, Lathromeroidea gerriphaga Pinto is known to use its

KEY TO THE FAMILIES OF "AQUATIC"

wings to swim if accidentally submerged, but does not swim underwater in search of eggs (Henriquez and Spence 1993; Pinto 2006). Prestwichia is a genus that has often been reared from aquatic insect eggs in Europe (Henriksen 1922; Jackson 1956b). Martin (1927) claims it was found in New York but this record has not been substantiated. A species has

All known hymenopteran families that contain species that enter the water as adults are included in the key, as well as families having species that parasit ize aquatic hosts, regardless of whether the parasit ized host stage is terrestrial or aquatic. Because many of the hymenopteran larvae that parasitize aquatic insects cannot be separated satisfactorily on a mor phological basis, a key to the larvae is not presented. Rather, some general descriptive information for the larvae ofeach family is given below,followed by a key

recently been discovered in Florida (Pinto 1997)and it is probably associated with aquatic insect eggs, but no host record is currently known for this species (see Table 22A). Within the Eulophidae, four genera are associ ated with eggs and pupae of aquatic beetles or odonates (Table 22A), although it is not known if any enter the water in search of hosts. In the Pteromali-

dae, six genera have been reared from aquatic hosts but only Urolepis rufipes (Ashmead) has been wit nessed entering water to find hosts (Howarth and Polhemus 1991; Gibson 2000).

HYMENOPTERA

to adults. The reader is cautioned that larvae of most

genera of Hymenoptera are unknown and thus the descriptions below may not apply to all Hymenoptera larvae, whether aquatic or terrestrial. For additional larval descriptions and figures, the reader is referred to Parker (1924), Hagen (1964), and Short (1978) as well as the references to larvae listed in the section on

Additional Taxonomic References following the key. Larvae

EXTERNAL MORPHOLOGY

Generally, Hymenoptera that parasitize aquatic insects show little or no external morphological adap tations in the larva compared with those parasitizing terrestrial hosts. Most aquatic Hymenoptera have endoparasitoid larvae and since the larvae ofendoparasitoids are already modified to live in a liquid envi ronment (host hemolymph), no modifications would be expected. The only truly aquatic hymenopterans with ectoparasitoid larvae are in the ichneumonid genus Agriotypus which form a pre-pupal ribbon-like respiratory filament that extrudes from the host caddisfly pupal case and remains functional throughout the pupal stage (Bennett 2001). Two hypothesized aquatic adaptations in adult Hymenoptera associated with water are:(1) modified tarsi and claws for holding on to the substrate under water, and (2)relatively dense body setae or sculpture that could help in the formation of a plastron of air around the body. Several figitid species in the endemic Hawaiian genus Aspidogyrus have elongated tarsi and divided claws(unique in eucoiline figitids) that enable them to move over rock surfaces in swiftly moving streams, thus facilitating the parasitization of their dipteran hosts (larval canacaeids and ephydrids) (Beardsley 1992). The ichneumonid genera Tanychela and Agriotypus similarly possess relatively elongate tarsal claws(Dasch 1979; Bennett 2001). Dense body hair is found in several taxa including Tanychela, Agriotypus, and Apsilops(Townes and Townes 1962) and in the pteromalid genus Urolepis(Gibson 2000).

1. Vespoidea (Pompilidae)(Description based on terrestrial Pepsis sp.)(Evans 1959) First and final instar: body segmented, elongate, grub-like, head differentiated from body, caudal appendage absent; mandibles large, tridentate, maxil lary and labial palpi projecting as tubercles; spiracles present.

2. Platygastroidea (Platygastridae - previously Scelionidae) (Description based on Martin 1927 for Tiphodytes) First instar: body unsegmented, but divided into three regions by constrictions, long caudal appendage present; mandibles large and sharply pointed, palpi reduced; spiracles absent. Final instar: body segmented, elongate/ovoid without constrictions or a caudal appendage; mandi bles slender and thread-like with broad base, palpi reduced; spiracles present. 3. Diaprioidea (previously Proctotrupoidea) (Diapriidae)(Description based on O'Neill 1973 for Trichopria) First instar: body segmented, head differentiated from body, posterior segment generally modified into a short,forked appendage; mandibles small and indis tinct, pair of projecting papillae present dorsal to mouth; spiracles absent.

Final instar: body segmented, head not differenti ated from body, caudal appendage absent; mandibles distinct, with or without denticles, papillae absent dorsal to mouth; three pairs of spiracles present.

Chapter 22 Aquatic Hymenoptera

4. Ichneumonoidea (Braconidae and Ichneumonidae) (Description based on Short 1952, 1978; Hagen 1964). Body segmented, elongate, head generally differ entiated from body, caudal appendage present or absent; mandibles present and generally outlined by strongly sclerotized rods, especially in later instars, palpi disc-like, spiracles present or absent. First instar: most Ichneumonidae have a caudal

appendage, whereas most Braconidae do not; spira cles are generally present in ectoparasitoids and absent in endoparasitoids. Final instar: generally without a caudal append age (long, paired appendage present in Agriotypus) (Bennett 2001); endoparasitoids generally have smooth mandibles and disc-like antennae, ectopara sitoids generally have denticulate mandibles and papillate antennae; spiracles almost always present but apparently absent in Agriotypus. 5. Chalcidoidea

a) Chalcididae {Chalets) Description based on Brachymeria podagrica (Fabricius), a terrestrial pupal parasitoid of Sarcophaga (Parker 1924), thus Chalets spp. parasitizing eggs of Stratiomyidae may differ. First instar: body segmented, elongate, head con stricted from body and posterior segment forming an elongate caudal appendage; hook-like mandibles present, palpi disc-like; spiracles absent. Final instar: body segmented and ovoid, head only slightly separated from body, caudal appendage absent; triangular denticulate mandibles present, palpi disc-like; nine pairs of spiracles present.

b) Eulophidae (Descriptions based on Parker 1924; Hagen 1964; Jackson 1964) First instar: body segmented, elongate to ovoid, head constricted from the body, posterior segment narrowed into a caudal appendage {Aprostoeetus) or segment not narrowed (e.g., terrestrial genus Meltttobta), posterior segment bearing two spine-like pro cesses in some taxa(e.g.,some Tetrasttehus); mandibles hook-like, palpi reduced; four pairs of spiracles {Aprostoeetus), five pairs {Meltttobta), or spiracles absent{Mestoeharts and some species of Tetrasttehus). Final instar: body segmented, elongate to ovoid, head constricted from body,posterior segment generally not narrowed except in Aprostoeetus (spine-like pro cesses absent in Tetrasttehus); mandibles hook-like, palpi disc-like except in Mestoeharts, maxillary palpi twosegmented; nine pairs of spiracles present or spiracles absent {Mestoeharts and some species of Tetrasttehus). c) Mymaridae (larvae too diverse for general family description except that palpi are always indistinct and spiracles always absent)

913

Camphractus (Description based on C. ctnetus) (Jackson 1961b) First instar: (Fig. 22.6) body unsegmented, elon gate, cylindrical, head not separated from body, cau dal appendage absent; mandibles absent. Final instar:(Fig. 22.9) as first instar except body sac-like.

Anagrus (Description based on terrestrial Anagrus spp.)(Bakkendorf 1934; Sahad 1984) First instar: body unsegmented, sac-like, head barely or not separated from body by constriction, four needle-like caudal processes may be present; mandibles minute.

Final instar: body weakly segmented, more elon gate than first instar, caudal process absent; hook-like mandibles present. Anaphes (Description based on the terrestrial Anaphes vtetus Huber)(Nenon et al. 1995) First instar: body unsegmented, with medial con striction, head not clearly differentiated, last segment narrowed into a long appendage; mouth bearing one small tooth.

Final instar: body sac-like, with weak segmenta tion between head, thorax and abdomen, caudal

appendage absent; hook-like mandibles present. Polynema (Description based on the terrestrial Polynema strtatteorne Girault(Balduf 1928) First instar: body weakly segmented, spindleshaped, tapering strongly anteriorly and posteriorly, head constricted from body, posterior segment tapered to a long, pointed caudal appendage with a sharp basal tooth; mandibles indistinct. Final instar: body lacking segments or very weakly segmented, sac-like, head not differentiated from body, caudal appendage absent; hook-like mandibles present.

e) Pteromalidae (Description based on Parker 1924)

First instar: body segmented, elongate to ovoid, head generally separated from body, caudal append age generally absent; mandibles hook-like, palpi reduced; four or five pairs of spiracles present or spir acles absent.

Final instar: body segmented, ovoid to cylindrical without a caudal appendage, mandibles present(gen

erally triangular), palpi reduced; 9 to 10 pairs of spir acles present.

f) Trichogrammatidae (Description based on Parker 1924 for terrestrial Trtehogramma) First instar: body unsegmented, ovoid, sac-like, without differentiated head or caudal appendage; mandibles minute or absent, palpi reduced; spiracles absent.

914

Chapter 22 Aquatic Hymenoptera

Final instar: body ovoid, weakly segmented, head not differentiated from body, caudal appendage absent; mandibles hook-like, palpi reduced; spiracles absent. 6. Cynipoidea (Figitidae)(Description based on Hagen 1964) First instar: body segmented, elongate with several long, fleshy thoracic appendages, head differentiated from body, long, caudal appendage present; mandibles small, palpi reduced; spiracles absent. Final instar: body segmented, ovoid, caudal appendage absent; mandibles bidentate (Hexacola) or with a series of denticles (Kleidotoma), palpi reduced; spiracles present.

Adults

1.

r.

2(1).

Wing venation reduced; front wings usually with no enclosed cells, but, if present, with less than 5 enclosed cells; wings usually with long marginal fringe; hind wings veinless or with one vein (enclosed cells absent); rarely wingless (Figs. 22.11-22.13,22.16) Wing venation well developed; front wings with more than 5 enclosed cells and without long marginal fringe, at least on anterior margins; hind wings with more than 2 veins and with at least one enclosed cell (Figs. 22.19-22.22) Hind femur enlarged and with toothed or denticulate ventral margin

2

9

CHALCIDIDAE

2'.

Hind femur not enlarged and without toothed ventral margin

3

3(2).

Tarsi 4-or 5-segmented

4

3'.

Tarsi 3-segmented (Fig. 22.11); parasites of insect eggs

4(3).

Tarsi 4-segmented (Fig. 22.13)

5

4'.

Tarsi 5-segmented (Figs. 22.16, 22.20)

6

5(4).

Marginal vein short, terminating within the first third of wing's length; stigmal vein absent(Fig. 22.12); body not metallic; parasites of insect eggs

5'. 6(4').

Marginal vein long, extending beyond one-half of wing's length; stigmal vein present(Fig. 22.13); body with metallic reflections or highly colored Antennae inserted on shelf at middle of face (Fig. 22.14)

6'.

Antennae not arising from shelf at middle of face (Fig. 22.16B)

7(6').

Antennae with 2 distinctly smaller segments (anelli) between pedicel and first funicle segment; together the anelli thinner and usually shorter than first funicle segment(Fig. 22.15)

7'. 8(7'). 8'. 9(T).

9'. 10(9). 10'.

TRICHOGRAMMATIDAE

Antennae without 2 distinctly smaller segments (anelli) between pedicel and first funicle segment(Fig. 22.17)

MYMARIDAE EULOPHIDAE DIAPRIIDAE 7

PTEROMALIDAE 8

Scutellum with 2 pits at base and an elevated pit("cup")on disk; forewing without stigmal vein; abdomen compressed

FIGITIDAE

Scutellum without an elevated cup (Fig. 22.18)forewing with stigmal vein (Figs. 22.13, 22.16A); abdomen depressed; parasites of eggs

PLATYGASTRIDAE

Hind wings without an anal lobe (Figs. 22.20, 22.21); trochanters 2-segmented (Fig. 22.20); antennae with more than 15 segments

10

Hind wings with an anal lobe (Fig. 22.19); trochanters 1-segmented; antennae with less than 15 segments and curled apically; parasites of spiders POMPILIDAE Forewing with 2 recurrent veins(second recurrent vein is a crossvein in apical lower half of wing disk)(Fig. 22.20) ICHNEUMONIDAE Forewing with one recurrent vein (no crossvein in apical lower half of wing) (Figs. 22.21-22.22)

BRACONIDAE

Chapter 22 Aquatic Hymenoptera

915

marginal fringe

antenna inserted on shelf

u ■ ) ) ) ) ) ) ) ) ) ) ) ) )

Maruini

Moth Flies -

Lentic—littoral

Telmatoscopus) Clogmia

Paramormia

holes); lotic— depositional

Pericoma &

holes); lotic— depositional

(including tree

Lentic—littoral

holes); lotic— depositional

(including tree

Lentic—littoral

(including tree

Pericomaina (47)

Borrowers

Borrowers

Borrowers

gatherers

Collectors—

gatherers

Collectors—

gatherers

Collectors—

gatherers

Scrapers;

madicolous)

Clingers

gatherers

collectors—

collectors—

Lotic—erosional

borrowers

Generally

Generally

gatherers

abdominal

prolegs)

collectors—

Scrapers;

Trophic Relationships

Clingers (elongate

Habit

(margins,

(previously =

Maruina (3)

(detritus)

lentic—littoral

(aquatic mosses)

Lotic—erosional

Habitat

Psychodinae

Species

Generally lotic— depositional;

Nymphomyia (2) (=Palaeodipteron)

Genus

Psychodidae (73)-

Nymphomyiidae (2)

Family

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Order

of species in parentheses)

Taxa (number

Table 23B

Widespread

West

East

Distribution

American

North SE UM

5.6

M

4.0

1.0

10.0

NW

Ecological

1073,2975, 3484, 5282, 5473, 6116, 6117, 6328, 6331,6677, t

2688, 4846, 6677, 574

2937, 2975, 3179, 4040, 4599, 4844, 6677, 6492

245, 1297, 2845, 3131, 966, 1163, 2406, 3681, 3683, 5523

MA* References**

4.0

Tolerance Values

)) ) ) )) ) )))) ))) ) ) ) ) ) )))) ) ) )

o

vo

Phantom Crane Flies

Ptychopteridae (16) (=Liriopeidae) -

Psychodini (23)

Family Genus

Threticus

Psychoda

Thornburghiella

Stupkaiella

Pneumia

Continued

Species

Generally lotic— depositional (including springs); lentic— littoral (sediments. detritus)

holes)

(including tree

Lentic—littoral

beaches—marine

depositionai;

(detritus); lotic—

Lentic—littoral

beaches—marine

(detritus); lotic— depositional;

Lentic—littoral

littoral (detritus)

depositional (margins); lentic—

Lotic—

littoral (detritus)

depositional (margins); lentic—

Lotic—

depositional (margins); lentic— littoral (detritus)

Lotic—

Habitat

Habit

DIPTERA

burrowers

Generally

Borrowers

Borrowers

Borrowers

Borrowers

Borrowers

Borrowers

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Order

of species In parentheses)

Taxa (number

Table 23B

gatherers

collectors—

Generally obligate

gatherers

Collectors—

gatherers

Collectors—

gatherers

Collectors—

gatherers

Collectors—

gatherers

Collectors—

gatherers

Collectors—

Trophic Relationships

North

East

Widespread

Widespread

Distribution

American

9.9

SE

UM

3.7

M

7.0

10.0

NW

Tolerance Values

(continued)

1756

2937,2975, 3179, 4599, 6677, 6492,

6677

1073,3178, 4003, 5281, 5473, 6117,

MA* References**

Ecological

j J ) ) ) ) ) ) ) ) ) ) ) ) ) j ) )) ) ) ) ) ) ) )

Thaumaleidae (28)

(3)

Trichothaumalea

Lotic—madicolous

Lotic—madicolous

Thaumalea

verralli

Lotic—madicolous

Androprosopa (24)

Lotic—madicolous

Lotic—erosional

Protoplasa

Scrapers

Scrapers

Clingers

Clingers

North

East, West

Newfoundland

Only in

Widespread

Eastern

Western

Widespread

Widespread

Widespread

Distribution

American

5.0

SE

UM

M

1.0

7.0

NW

Tolerance Values

Ecological

5467, 6677

6677

2120, 2975, 5473, 5763,

212, 2120, 6677, 5470

47

1734,3195, 5114, 6707

3826

3828, 5103,

42, 2417, 2673,

42, 1757

42, 2417, 5103

MA* References**

) ) ) ) ) ) ) )) ) ) ))) ))) ) ) ) )) ) ) ) ) )

Scrapers

Scrapers

Clingers

Clingers

borrowers

Sprawlers—

borrowers

Sprawlers—

borrowers

Lotic—erosional

Protanyderus(3)

Sprawlers—

Lotic—erosional

(sediments)

herbivores

(Sphagnum)

hydrophytes (bogs)

Collectors— shredders—

Primitive Crane Elies

- Solitary Midges

Collectors—

gatherers

lentic—vascular

Borrowers

Collectors-

gatherers

gatherers;

Lotic—

Ptychoptera (10) (=Uriope)

Borrowers

Borrowers

Habit

Trophic Relationships

depositional;

Lotic—

depositional

hydrophytes (emergent zone)

lentic—^vascular

depositional;

Lotic—

Habitat

Bittacomorphella

fitchii

Species

(5)

Bittacomorpha (2)

Genus

Tanyderidae (4)-

Family

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Order

of species In parentheses)

Taxa(number

Table 23B

vo

Athericidae (4)

Amblypsilopus(16)

Achradocera (2)

Achalcus(5)

Suragina

condnna

Sprawlers— burrowers

Lentic—littoral

margins; lotic— margins

Lotic—margins

Predators

Generally sprawlers—

Generally lentic and lotic margins (semiaquatic)

DIPTERA

(piercers)

Generally predators (piercers)

burrowers

and depositional

Widespread

Widespread

West Coast

Texas, Mexico

(piercers)

Sprawlers—

Lotic—erosional

burrowers

Southwest

Predators

burrowers

Widespread

(piercers)

Sprawlers— Predators

North

American Distribution

Lotic—erosional

Habit

Trophic Relationships

and depositional

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Dolichopodidae(1127)Long-Legged Flies

(=Rhagionidae, in part) - Snipe Flies Atherix(3)

Continued

Family

Lower Brachycera

Order

of species in parentheses)

Taxa (number

Table 23B

9.7

2.1

SE

4.0

2.0

UM

3.1

M

4.0

2.0

2.0

(continued)

4892

4819

1143, 2650, 2975, 4040, 929, 1561, 6605, 6677,

4247, 4249, 6381, 6382, 4248, 6383

2793

2937, 2975, 3179, 4321, 6659, 4322, 4599, 5959, 6328, 6381, 2792, 3438, 5339, 6492,

4249, 5958

Ecological NW MA* References**

Tolerance Values

J ) ) ) ) ) ) ) )) ) ))) ) ) ) ) ) ) ) ) )) ) )

vo

Family Genus

borrowers

borrowers

Lentic—Iittorai

(margins),

DoUchopus(317)

Widespread

Widespread

Lotic—margins

Harmstonia (2)

Lentic—Iittorai

(margins)

Gymnopternus (76)

East

Widespread

Widespread Arizona

Lotic—margins

Widespread

Widespread

Widespread

Enlinia (8)

(piercers)

Predators

(piercers)

Erebomyia

estuaries; iotic— margins

Sprawiers—

Lotic—margins

(margins); Iotic— margins

Lentic—littoral

borrowers

(margins)

Diostracus(3)

Diaphorus(38)

Sprawiers—

Lentic—iittorai

Predators

gatherers?

Collectors—

(piercers)

Predators

Widespread

Widespread

borrowers

Predators

(piercers)

Distribution

Widespread

Sprawiers—

Lentic—Iittorai

borrowers

iakes; iotic— margins

(margins)

Sprawiers—

Beach zone—

beach zone)

Sprawiers—

Lentic—iittorai

Habit

(margins and

Habitat

North American

Chrysotus(107)

exalloptera

Species

Trophic Relationships

Chrysotimus(7)

Campsicnemus (22)

Asyndetus(23)

Argyra (46)

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atiantic ** Emphasis on trophic reiationships

Order

of species in parentheses)

Taxa (number

Table 23B

SE

UM

M

NW

Ecological

4892

4892

1560, 2650, 6115, 6677,

4892

6140, 6677

5522

1560, 2650,

4892

478, 995, 2650, 6115, 6677,

MA* References**

Tolerance Values

) ) ) ))) ))) ))))) ) ))) ) ) )) ))))

4^

Ul

vo

Family Genus

borrowers (in fine detritus)

Sprawlers— borrowers

Beach zonemarine

Beach zone—

Melanderia (3)

Nematoproctus (7)

Nanomyina

Micromorphus(7)

Lotic—seeps (algal mats)

Liancalus(5)

Lotic—margins

marine

Beach zone—

(margins)

Lentic—littoral

shore line)

marine (rocky

Lentic—littoral

(margins)

barbata

Predators

Sprawlers—

Lentic—littoral

(margins),

Predators

borrowers

Clingers

DIPTERA

(piercers)

Predators

(piercers)

Sprawlers—

(piercers)

Predators

(piercers) (especially midges)

borrowers

estuaries

(piercers)

Sprawlers— Predators

(piercers)

Predators

Trophic Relationships

Lentic—littoral

Sprawlers (damp moss)

Habit

(margins)

tree holes

depositional (moss); lentic—

Lotic—

Habitat

Lamprochromus

slossonae

Species

(3)

Keirosoma

Hypocharassus(2)

Hydrophorus(50)

Hydatostega (3)

Hercostomus(26)

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Order

of species in parentheses)

Taxa (number

Table 23B

American

East

East Coast

Widespread

West Coast

Widespread

Widespread

Florida

East Coast

Widespread

Widespread

Widespread

Distribution

SE

UM

M

NW

Tolerance Values

(continued)

5320, 6677

1145, 478, 5473, 6115

6677

2650, 5522,

6677

995, 1560, 5522, 6115,

6677

995, 1560, 5522, 6115,

MA* References**!

Ecological

) ) J )) ) ) ) ) ) J ) ) ) ) ) ) ) ) ) ) ) ) ) ) )

Family

Predators

Lentic—littoral

(margins)

Lotic—margins

Syntormon (20)

Widespread

Widespread

Tennessee

California, Texas,

North and

Widespread

East

Widespread

Widespread

West

West Coast

East, South

Widespread

Distribution

West

(piercers)

Predators

(piercers)

North American

Lentic—littoral

borrowers

Sprawlers—

Clingers (in algae on rocks)

Habit

Trophic Relationships

(margins)

(margins); lotic— margins

Sympycnus(28)

(3)

Sympycnidelphus

Scelius(15)

Rhaphium (85) Lentic—littoral

Lotic—margins

Peloropeodes(10) univittatus

mats)

Plagioneurus

Lotic—seeps (algal

Pelastoneurus

marine (intertidal, on rocks)

Beach zone—

(41)

Parasyntormon (20)

Paraphrosylus(6) {=Aphrosylus)

(margins); lotic— margins

Lentic—littoral

Habitat

Lotic—margins

Species

Nepalomyia (4)

Genus

Paradius(15)

Continued

*SE = Southeast, DM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Order

of species in parentheses)

Taxa (number

Table 23B

SE

UM

M

NW

Tolerance Values

Ecological

6115,4892

4892

4892

4892

1144, 1146,

2650, 5320, 6115, 6677

MA* References**

) ) ) )) » ))) ) ) ) ))) ) ) ) ) ) )) > ) ) ) )

a\

J

vo vo

Predators

(piercers) Generally predators (piercers)

Generally sprawlers—

Generally lotic— erosional and

Empididae (282)-

Dance Flies

Chelipoda (6)

Chelifera (12)

depositional

borrowers

Unknown

(bogs)

lentic—littoral

Sprawlers—

Lotic—

borrowers

depositional;

littoral

(detritus); lentic—

rocks)

borrowers

Predators

lakes

marine (Intertidal rocks)

Beach zone—

Sprawlers—

Thinophilus(25)

Thambemyia

Beach zone—

borealis

(margins)

Lentic—littoral

depositional

Lotic—

lotic—margins

(piercers)

Teuchophorus(4)

Telmaturgus pan/us

Predators

borrowers

(piercers)

Sprawlers—

Lentic—tree

holes, beaches;

Tachytrechus(38)

Predators

(piercers) (especially Dasyhelea)

borrowers

Habit

Sprawlers—

Habitat Lentic—tree holes

Species

Systenus(6)

Genus

Trophic Relationships

Clingers (in algae on

Family

Continued

DIPTERA

*SE = Southeast, DM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Order

of species In parentheses)

Taxa (number

Table 23B

North

Widespread

Widespread

Widespread

Coastal

Widespread

East

Widespread

East and South

Distribution

American

8.1

SE

6.0

UM

3.5

M

6.0

6.0

NW

Tolerance Values

6043

(continued)

2391,3626,

6328

679, 3390, 3624, 3627, 6677, 4602,

1243, 2937, 2975, 3179, 3216, 3217, 4599, 5747, 6677, 929, 1561, 5829

5522

5822

6115

6115

4892

6115, 6677,

1147, 3322,

6677

MA* References**

Ecological

J ) I I ) ) ) ) ) J ) )) > ) ) ) )) ) ) ) ) ) )

Family

Neoplasta (12)

Metachela (3)

and depositional

(16)

burrowers

and depositional

burrowers

banks)

just above on

(also in moss mats at water level, or

Sprawlers—

Lotic—erosional

and depositional

banks)

just above on

(also in moss mats at water level, or

Sprawlers—

borrowers

Sprawlers—

Clingers

Clingers

Habit

Lotic—erosional

(detritus)

Lotic—erosional

pullata

Lotic—erosional

(bogs)

lentic—littoral

Lotic—erosional;

Habitat

Hemerodromia

Heleodromia

(2)

Dolichocephala

Clinocera (42)

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Order

of species In parentheses)

Taxa (number

Table 23B

(piercers)

Predators

(piercers)

Predators

(piercers)

Predators

Relationships

Trophic

North

Widespread

Widespread (mostly West)

Widespread

Widespread

Widespread

Widespread

Distribution

American SE

UM

IVI

Ecological

6.0

6.0

3937

2391, 3390, 3624, 3629, 5309, 2382,

2391, 3390, 3624, 3625, 2382, 3937

3937

478, 1073, 1077, 2391, 2975, 3216, 3390, 6062, 6114, 6677, 3624, 410, 2382, 3936,

6113, 5469

679, 3390, 4356, 6114, 6677, 5473, 6328, 5469

NW MA* References**

Tolerance Values

» ) ) I ) ) ))) ) )) > ) ) ) ) ) ) )))) > ) ) )

vo 00

Pelecorhynchidae (7)

Oreoleptidae(1)

Family Genus Predators

Glutops(7)

Oreoteptis

Wiedemannia (6)

Sprawlers— borrowers

Lotic—

Clingers

depositional

Lotic—erosional

Clingers

Clingers

Lotic—erosional

Trichodinocera

Lotic—erosional

Clingers

Lotic—erosional

Roederiodes(6)

(6)

borrowers

Sprawlers—

(150)

DIPTERA

Predators

herbivores?

shredders—

(piercers):

Predators

(piercers)

Predators

(piercers)

Predators

(piercers)

Predators

Simuliidae)

(piercers)(pupal

Widespread

Northwest

West, North

Widespread

Widespread

Widespread

Lentic—littoral

borrowers

Rhamphomyia

Predators

(piercers)

Sprawlers—

Widespread

Oreothalia (5)

Lotic—margins?

West

Proclinopyga (5)

(piercers) (Simuliidae, Trichoptera)

American

Distribution

West

borrowers

Trophic Relationships

Southeast,

Sprawlers—

Lotic—erosional

Habit

(moss)

Habitat

Lotic—erosional,

torrenticola

Species

North

seeps

Oreogeton (8)

Continued

t

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Order

parentheses)

Taxa (number of species in

Table 23B

J

SE

UM

M

3,0

3.0

6.0

6.0

6.0

5.0

NW

Tolerance Values

Ecological

(continued)

5937, 5939

3302

6874

6115, 5471

6115, 5468

6677

478, 1011, 3390, 4278,

6115

5469

5469

5578

MA* References**

© o

Hoplitimyia (4)

Hedriodiscus(7)

Euparyphus(22)

Caloparyphus(12)

Anoplodonta

Allognosta (4)

nigrirostris

borrowers

Lentic—vascular

Sprawlers— borrowers

Lotic—erosional

»

Climbers

Sprawlers

and depositional (margins)

hydrophytes)

(vascular

depositionnal

Lotic—

and depositional (margins)

Lotic—erosional

lotic—erosional

hydrophytes (emergent zone);

Sprawlers

borrowers

littoral

Sprawlers—

Lotic—erosional

and depositional (margins); lentic—

littoral

Sprawlers—

West t

Widespread

Widespread

Widespread

West t

East

Distribution

American

North

SE

UM

M

7.0

7.0

8.0

NW

Tolerance Values

Ecological

3942, 5753

3942, 4920, 5466, 5473

5473

3942, 5466,

2937, 2974, 2975, 3179, 3942, 3943, 4599, 5180, 5181, 6677

MA* References**

i ) 1 ) > ) ) ) ) I ) I > )) )))

gatherers

Collectors—

Scrapers

scrapers

gatherers;

Collectors—

gatherers

Collectors—

gatherers

Collectors—

gatherers

Collectors—

gatherers

swimmers

Lotic—erosional

collectors—

Generally

Trophic Relationships

Generally sprawlers—

Habit

and depositional (margins); lentic—

littoral

Habitat

Generally lentic—

Species

Stratiomyidae (186)

Genus

- Soldier Flies

Family

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Order

of species In parentheses)

Taxa (number

Table 23B

o o

Family Genus

and depositional (margins) Mostly terrestrial,

i^Hermione)

Sargus(6)

Stratiomys(31) (=Stratiomyia)

borrowers

Lotic—erosional

Oxycera (7)

Collectors—

Sprawlers— borrowers

Lotic—erosional

and depositional (margins); lentic— littoral (including saline pools)

DIPTERA

filterers)

gatherers (and

Collectors—

scrapers

vascular

hydrophytes (emergent zone)

gatherers;

Collectors—

Scrapers

gatherers; scrapers?

Collectors—

gatherers

some lentic—

Climbers

Sprawlers—

hydrophytes (emergent zone)

Sprawlers

Lentic—vascular

pools, marshes); lotic—margins

Lentic—littoral, beaches (saline

Swimmers; sprawlers

borrowers

littoral

Collectors—

Sprawlers—

Lotic—erosional

and depositional (margins); lentic— gatherers

gatherers

borrowers

Collectors—

Sprawlers—

Trophic Relationships

Lotic—erosional

Habit

and depositional (margins)

Habitat

Odontomyia (30)

Species

{=Eulalia)

Nemotelus(38)

Myxosargus(4)

Labostigmina (20)

Continued

NW

Ecological

7.0

6205

478,2974, 3942, 5180, 5473, 1204,

6677

478, 2975, 3942, 5180,

MA* References**

Widespread

(continued)

1762, 2974, 3278, 3306, 3942, 5180, 6020, 6677

3943, 5180

M

Widespread

UM

202, 3942, 5180, 6677, 201, 6492

SE

Tolerance Values

East

Widespread

Widespread

Widespread

Widespread t

Distribution

American

North

J ) > )) ) ) )) J ) ) ) ) ) ) )

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Order

of species in parentheses)

Taxa(number

Table 23B

; j J

o o

Tabanidae (332)Horse Flies, Deer Flies

Family

Genus

Chrysops(83)

Chlorotabanus

Brennania (2)

Bolbodimyia

Atylotusi^A)

Apatolestes0^)

Agkistrocerus(2)

Continued

crepuscularis

atrata

Species

»

»

Predators

Predators

(piercers) borrowers

(piercers)

Predators

depositional

lotic—

nr

SE o

UM

M

q

8.0

NW

Tolerance Values

Ecological

4016, 1969, 5309, 5943

5936, 1163, 3671, 3954,

478, 780, 4683,

2191

4046

777

855, 5936, 6388, 1163, 5940, 5941

3458

782

5943

6677, 3394,

5936, 1163, 4016, 6388,

4599, 5073,

240, 855, 2937, 2975, 3179,

MA* References**

q

^ ) 1 ) 1 ) )) ) F ) ) ) ) ) )) 1

Widespread

Coast

Sooth, East

California coast

Southwest

Widespread

California coast; West

South

Distribution

Predators

Sprawlers—

and estuaries;

North American

(piercers)

(piercers)

Predators

(piercers)

Lentic—littoral;

borrowers

Predators

(piercers)

beaches—marine

Sprawlers—

Lentic—littoral

borrowers

(sediments)

Sprawlers—

Lentic—coastal dunes

Sprawlers

borrowers

Lotic—erosional

Sprawlers—

Lentic—littoral

borrowers

(sediments and mosses)

Sprawlers—

(piercers)

borrowers

marine

Predators

Beach zone—

lentic—littoral

depositional;

Lotic—

sediments, and detritus)

Generally predators (piercers)

Trophic Relationships

Sprawlers—

borrowers

lentic—littoral

(margin,

Generally sprawlers;

Habit

Generally lotic— depositional;

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Order

of species in parentheses)

Taxa (number

Table 23B

©

Table 23B

^

Family

borrowers

Tabanus(108)

Stenotabanus(6)

depositional

Predators

DIPTERA

Predators

(piercers)

borrowers

(piercers)

Sprawiers—

borrowers

Lentic—littoral; lotic—erosional,

Sprawiers—

marine

borrowers

Beach zone—

Sprawiers—

Lentic—littoral;

lotic—margins

depositional

borrowers

S/7Wus(11)

Sprawiers—

Lentic—littoral; lotic—

Predators

(piercers)

borrowers

(piercers)

Predators

Trophic Relationships

Sprawiers—

Merycomyia (2)

depositional

Sprawiers—

Lentic—littorai;

borrowers

lotic—

Sprawiers—

Lentic—littorai

Habit

(sediments)

Habitat

Lentic—tree holes

Species

Leucotabanus(2)

Hybomitra (55)

Haematopota (5)

Genus

Widespread

South

Central, West

East

Sooth

Widespread

Widespread

Distribution

American

North

9.7

SE

5.0

UM

M

5.0

NW

Ecological

(continued)

1163, 478, 1073, 1077, 4016, 3671, 5071, 5086, 5943, 1969, 2194, 2877, 5309, 574

2193

1163, 3393, 4016, 5933

2192, 5936

2191

5994

1163, 5936,

855

1163, 5180,

MA* References**

5.0

Tolerance Values

) > ) ) ) ) ))) ) ) ) ) ) ) ) ) ) ) ) )

Continued

, >•

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = ^ Northwest, MA = Mid-Atlantic "* Emphasis on trophic relationships

Order

of species in parentheses)

Taxa (number

0'

o o

Family Genus

anomala

marine intertidal

Beach zone—

marine intertidal

Beach zone—

Borrowers

Borrowers

gatherers

collectors—

North

NW

Ecological

2650, 6604

2650, 5945

2650, 6604, 6663, 6677, 3795, 6676

MA* References**

California

Coasts

East and West

Coast

Sootheast

Florida

y ) ) ) ))

6187

1451, 1612, 2668, 3802, 3852, 3853, 4692, 4693,

3792

5083

M

2650, 5082,

UM

West and

SE

Tolerance Values

Sooth Coasts

coast

California

United States

Coasts of

East and West

Distribution

American

) > ) r) ) J ) ) )) ) ) j

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = : Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Coelopina

Coelopa (4)

(seaweed wracks)

intertidal

Generally shredders;

Generally borrowers

beaches— marine

Scrapers (graze algae)

Scrapers (graze algae)

Scrapers (graze algae)

scrapers

Generally

Trophic Relationships

Borrowers

Generally

marine intertidal

Beach zone—

Borrowers

Borrowers

Borrowers

Flies

dianneae

Procanace

marine intertidal

Beach zone—

marine intertidal

Beach zone—

marine intertidal

Beach zone—

Coelopidae (5)- Kelp

aicen

Paracanace

Nocticanace(3)

Canaceoides(7)

marine intertidal

Beach zone— Borrowers

borrowers

intertldal

Generally

beaches— marine

Habit

Generally

Habitat

Beach Flies

Species

Canacidae (14) -

Canace(2)

Continued

Brachycera-Cyclorrhapha

Order

of species In parentheses)

Taxa (number

Table 23B

o o

Discomyzinae (34)

Shore Flies, Brine Flies

Ephydridae (464)-

Dryomyzidae (2)

Family

Genus

americanum

undnata

Clasiopella

Species

Clanoneurum

Ceropsilopa (7)

Oedoparena (2)

Continued

gatherers;

sprawlers

and vascular

gatherers; some shreders—

herbivores and,

borrowers

(some climbers,

planktonic)

miners

marine

Beach zone—

(mangrove)

emergent zone

Borrowers—

Marine—vascular

hydrophytes,

DIPTERA

(miners)

herbivores

Elorida

Coasts

West and East

West and Sooth Coasts

West Coast

Distribution

marine

Shredders—

predators (piercers)

Generally collectors—

Generally

North American

Beach zone—

Generally lentic— littoral; marine shores; lotic— margins

predators (engulfers)

scrapers;

(miners);

herbivores

shredders—

borrowers;

hydrophytes)

Generally collectors—

Generally

(barnacles)

barnacles

Generally lentic— littoral (margins

Predators

(engulfers)

Borrowers—

inside

marine intertidal

Habit

Trophic Relationships

Beach zone—

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Order

of species in parentheses)

Taxa (number

Table 23B

SE

UM

M

6.0

NW

Tolerance Values

Ecological

(continued)

3798

5460, 6677

2975, 3179, 4599, 5307, 5310, 5460, 6604, 6677, 1410, 1782, 1911, 5957, 6685, 1569, 1917, 1416, 5710, 3095

449, 1411, 2523, 2937,

3806

781, 1782, 3209, 5320,

MA* References**

Family Genus

Lentic—littoral

petroleum)

masses of

Collectors—

gatherers; shredders— herbivores

(miners)

"Planktonic" and burrowers

gatherers

decaying vegetation? Lentic—littoral

Collectors—

decaying vegetation

Burrowers—in

Collectors—

gatherers

Burrowers—in

larvae

Gulf area of

Widespread

Coasts

East and West

Widespread

Widespread

United States

California,

Florida

Collectors—

associated with

West and South Coasts

gatherers (organics

Sprawlers— burrowers, or large "planktonic"

carrion)

and pools of crude petroleum

Marine—shores

cressoni semilutea

Mimapsilopa

and waste oil

Lentic—pools of crude petroleum

marine

Beach zone—

collectors—

(detritus)

gatherers (scavengers of

Mollusca;

and margins

Lentic—littoral

East and South

Paratissa

Psilopa (8)

American

Distribution

Coasts

Parasites of

Trophic Relationships

Lentic—littoral

Borrowers

Habit

North

and margins

Habitat

Lentic—littoral

petrolei (petroleum flies)

Species

Leptopsilopa (3)

Helaeomyia

Guttipsilopa (2)

Discomyza (2)

Cressonomyia (3)

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Order

of species in parentheses)

Taxa (number

Table 23B

SE

UM

M

NW

Tolerance Values

Ecological

6677

478, 2975,

3796, 5310

5310, 5709

6677

6677

478, 5460,

MA* References**

) ) ) ) > ))) ) ) ) ) > )) ) ) ) ) ) > ) ) ) ))

o\

o o -a

Ephydrinae (145)

Family Genus

adfinis

Trimerinoides

herbivores; collectors—

borrowers or climbers

depositional (thermal springs);

Generally

gatherers; parasitic on

borrowers; swimmers

Marine—salt

marshes; lentlclittoral (alkaline lakes)

Cirrula (4)

(=Hydropyrus)

herbivores

(macroalgae); mats)

DIPTERA

gatherers

collectors—

Shredders—

Borrowers (in

floating algal

Widespread

West

North and

Widespread

Widespread

snails?

Collectors—

Sprawlers—

Callinapaea (2)

Lentic—littoral

gatherers; scrapers

marine—shores

and salt marshes

shredders—

Generally sprawlers—

Generally lentic— littoral; lotic—

West

North, West

(piercers)(spider egg masses)

Predators

hydrophytes (emergent zone)

Climbers

United States

(miners)?

hydrophytes Lentic—vascular

Eastern and southern

American Distribution

herbivores

Trophic Relationships Shredders—

Borrowers?

Habit

vascular

Lentic—littoral;

Habitat

North

Calocoenia (2)

Brachydeutera (4)

madizans

Speaes

Trimerina

Rhysophora (2)

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Order

of species in parentheses)

Taxa(number

Table 23B

SE

UM

M

(continued)

S460, 6671

38, 297S, 380S,

3092

1S20, 6163,

3073, S460, 6604, 6669, 2376, 3804, 3807, 6677,

1S69

1S20, 394, 378S, S320

Ecological NW MA* References*

Tolerance Values

3 )) ) ))) J 3 )) ) ))) ) ))) ) > )) ) ) J

Family

Shredders— herbivores

(macroalgae) Shredders— herbivores

Burrowers (in

floating algal mats)

Sprawlers— burrowers

Marine—salt marshes

Lentic—littoral

Limnellia (11) Lentic—littoral?

ponds

lentic—alkaline

Marine—shores

and salt marshes;

pools)

saline lakes and

(alkaline and

Lamproscatella

guttipenne

Lentic—littoral (in algal mats, including alkaline lakes); lotic— depositional (including thermal springs)

(12)

Haloscatella (5)

Eutaenionotum

Ephydra (13)

gatherers

collectors—

(macroalgae);

Collectors—

gatherers

burrowers

Habit

Trophic Relationships

Sprawlers—

Habitat

Dimecoenia (2)

Species Lentic—margins

Genus Coenia (2)

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Order

of species In parentheses)

Taxa (number

Table 23B

North

Widespread

Widespread

Widespread

North

Widespread

Coasts

East and West

Widespread

Distribution

American SE

UM

M

NW

Tolerance Values

Ecological

3786

5460

3788, 5310,

38, 486, 705, 707, 1097, 2975, 4707, 5310, 5460, 1569, 6671, 6677, 307

3805, 5460

5310, 5460

1916, 3784,

MA* References**

) ) ) ) ) ) ) ) ) ) ) ) >')) ) ) ) ) ) )) ) ) ) )

© cc

o vo

o

Family Genus

metallica

Rhinonapaea

Scatella (17)

Setacera (8)

alaskense

Species

Philotelma

Parydra (34)

Paracoenia (7)

Continued

climbers

margins

marshes; lotic—

shores and

Sprawlers-

hydrophytes and algal mats (including thermal springs); marine—

Sprawlers

Lentic—vascular

(algal mats), including saline pools; lotic— margins

Lentic—littoral

depositional

Lotic—

depositional and margins

lotic—

Borrowers

DIPTERA

gatherers; scrapers (graze algae)

Collectors—

scrapers (graze algae)

herbivores;

Shredders—

Scrapers

material)

gatherers (fecal

Lentic—littoral;

collectors—

Widespread

Widespread

North

West

Widespread

Widespread

Scrapers (graze bluegreen algae);

lentic—littoral

Sprawlers

Habit

North

American Distribution

Trophic Relationships

Lotic—alkaline thermal springs (in algal mats);

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Order

of species in parentheses)

Taxa (number

Table 23B

SE

UM

M

(continued)

1910, 3669, 5460, 6069, 6669, 6831, 1569, 5473

1099, 1413,

1910, 1914, 2975, 5310, 3789, 6677, 6830, 307

1413, 1417, 5310, 5460

6035, 307

705, 4645,

Ecological NW MA* References*

Tolerance Values

©

h-i

Gymnomyzinae (84)

Family Genus

lacteipennis

nitida

Diphuia

Species

Didasiopa

Athyroglossa (8)

Allotrichoma (11)

Thinoscatella (2)

Scatophila (24)

Continued

herbivores and

(some climbers,

planktonic)

Marine—shores

depositional

lotic—

Lentic—littoral;

material

hydrophytes, decaying organic

vascular

Lentic—littoral,

marine—shores

Lentic—temporary puddles(some near dung); lotic—margins;

Borrowers

snails)

puddles, and decaying

temporary

gatherers

Collectors-

Collectors—

gatherers(some in dung and decaying snails)

Borrowers (in

dung,

predators

gatherers; some shredders—

borrowers

margins

collectors—

Generally

Generally

Scrapers

Generally lentic—

Sprawlers

Habit

Trophic Relationships

littoral; marine— shores, lotic—

Marine—littoral

margins

marshes; lotic—

marine—salt

Lentic—littoral;

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Order

of species in parentheses)

Taxa (number

Table 23B

North

East Coast

Widespread

Widespread

Widespread

Coasts

East and West

Widespread

Distribution

American

SE

) ) ) ) ) ) ) ) ) ) ) ) > )))) ) ) )

o

UM

M

Ecological

3799

3794, 5460,

5310

5320

2251, 3301,

5206

575, 1411, 5310, 6677,

6677

1414, 5460,

MA* References**

) ))) )

NW

Tolerance Values

Family Genus

Lamprodasiopa (2)

Hydrochasma (3)

Hecamedoides

Hecamede (2)

(8)

Gymnoclasiopa

Glenanthe(5)

North

(semiaquatic)

Borrowers (in moss and

algae)

Lentic and iotic—

margins (primarily terrestrial)

marine—shores

lotic—margins;

Lentic—littoral:

Lentic—littoral?

collectors—

marine

gatherers (scavengers)

Borrowers

Probably

Beach zone—

DIPTERA

gatherers

Widespread

Widespread

Widespread

East Coast

Widespread

Coasts

Lentic—littoral

East and West

East, South

Widespread

Widespread

Distribution

American

Marine—salt

Collectors—

(engulfers of frog eggs)

Predators

gatherers

Collectors—

Trophic Relationships

marshes

hydrophytes (in frog eggs)

Borrowers

algae)

terrestrial)

Lentic—vascular

moss and

Gastrops(2)

Borrowers (in

margins (primarily

Habit

Lentic and lotic—

Habitat

Lentic—littoral

glaucelius

Species

Ditrichophora (7)

Discocerina (8)

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Order

of species In parentheses)

Taxa (number

Table 23B

SE

UM

M

NW

Ecological

(continued)

5460

6677

3797, 5460,

5460

578, 6180

5310

1413, 1922, 5310, 6677

MA* References**

Tolerance Values

) ) )) ) ) ) ) ) ) )

Family Genus

Tronamyia

iindsieyi

California

pools)

Southern

Lentic—littoral

Widespread

(saline lakes and

margins

marshes; lotic—

marine—salt

snails)

United States

snails)

Northern Collectors—

gatherers (decomposing

Borrowers (in

East Coast

decomposing

small mollusks

scavengers of

herbivores; predators or

Shredders—

Florida

Widespread

margins

Sprawiers

(engulfers) (midge larvae)

Widespread

Lotic and lentic—

Lentic—littoral;

helices

Platygymnopa

marine

Beach zone—

Polytrichophora

grandis

Placopsidella

marine

Beach zone—

depositional and margins

lotic—

Lentic—littoral;

(incl. alkaline lakes and ponds)

lentic—littoral

Marine—shores;

East and West

Distribution

Coasts

Predators

North American

Beaches—marine;

Borrowers

Habit

Trophic Relationships

lentic—saline

Habitat

(4)

bahamensis

sbssonae

Species

Paraglenanthe

Ochthera (13)

Mosillus(3)

Lipochaeta

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Order

of species in parentheses)

Taxa (number

Table 23B

SE

UM

M

NW

Tolerance Values

Ecological

5460

6670

3791, 3793

1413, 5459, 5460, 6677

5460

3808, 5308,

5308, 5460

MA* References**

J )) ) J I ) I ) ) ) ) ) )) )) ) ) ) ))) I ) )

H-l

o

o

Hydreliiinae (146)

Family

Notiphila (53)

Lemnaphila

Hydrellia (70)

Atissa (5)

Continued

scotlandae

sprawlers

hydrophytes);

(miners) (especially Potamogeton)

miners (of duckweeds)

Collectors—

gatherers and filterers

Borrowers

(attached to

roots by respiratory spine)

Lentic—littoral

hydrophytes, bogs); lotic— depositional

(miners of duckweeds)

herbivores

Shredders—

(detritus, vascular

duckweeds)

Borrowers—

Lentic—^vascular

hydrophytes (floating zone—

marine—shores

watercress);

hydrophytes,

(vascular

lotic—erosional

and depositional

herbivores

miners

Shredders—

Borrowers—

Lentic—vascular

hydrophytes;

lakes and pools)

marine; lentic— littoral (saline

Beach zone—

marine—shores

deposltional;

gatherers

borrowers;

lotic—

Generally collectors—

Generally

Generally lentic—

Habit

Trophic Relationships

littoral (vascular

Habitat

DIPTERA

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Order

of species in parentheses)

Taxa(number

Table 23B

North

Widespread

East

Widespread

Widespread

American Distribution SE

UM

M

(continued)

3424

6127, 6151, 6677, 5309,

452, 478, 804, 1419, 2523, 2724, 3787, 3944, 5460,

6677

4599, 5371,

452, 1412, 2247, 2975, 3944, 4599, 5460, 6604, 1520, 6677, 1415, 3094

5310, 5460

Ecological NW MA* References**

Tolerance Values

>> ) ) ) )) ) ))) ) ) )

J

■1^

)

o

llytheinae (55)

Family Genus

llythea (3)

)

>

)

)

)

)

)

)

)

)

)

Mid-Atlantic

(sediments)

depositional

lotic—

Lentic—littoral;

margins

Lotic and lentic—

Hyadina (9)

depositional

V

Sprawlers

Sprawlers

Sprawlers

)

Generally borrowers; sprawlers

littoral; lotic—

Sprawlers

Generally lentic—

emergent vascular hydrophyte beds)

(detritus at margins in

Lentic—littoral

Marine—shores

lentic—littoral

Marine—shores;

dung)

sediments near

(margins in

Lentic—littoral

)

gatherers

)

Collectors—

)

)

West, South

)

)

)

)

)

5310

1910, 5310

Widespread Scrapers (graze bluegreen algae)

5310

5310, 5460

5310, 6677

1910, 5310

)

Ecological

)

References**

North, East

Widespread

Widespread

Widespread

Tolerance values

Scrapers (graze bluegreen algae)

gatherers

collectors—

Generally

gatherers

Collectors-

gatherers

South Coasts

Distribution West and

Collectors-

North American

marine—shores

Sprawlers

Habit

Trophic Relationships

Lentic—littoral:

Habitat

Lentic—littoral

sallnum

nudus

Species

AKysfa (3)

Typopsilopa (4)

Schema

Ptilomyia (6)

Paralimna (5)

Oedenops

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = ** Emphasis on trophic relationships

Order

of species in parentheses)

Taxa (number

Table 23B

)

Ul

h-*

o

Heterocheilidae (1)

Helcomyzidae (1)

Family

Genus

Heterocheila

Hekomyza

Zeros(5)

Pseudohyadina

Philygria (6)

Pelina (10)

Nostima (10)

Lytogaster(8)

Continued

hannai

mirabilis

iongicornis

Species

Scrapers (graze bluegreen algae)

Sprawlers

Shredders;

(seaweed wracks) Burrowers

intertidal

Beach zone— marine intertidal

collectors—

burrowers

beaches—marine

DIPTERA

gatherers

collectors—

Shredders;

gatherers

Generally shredders;

Generally

gatherers

collectors—

Generally

marine intertidal

Burrowers

(seaweed wracks)

intertidal

Beach zone—

collectors—

burrowers

gatherers

Generally shredders:

Generally

gatherers

Collectors—

beaches—marine

Sprawlers

(decaying vascular plants)

detritivores

Shredders—

Scrapers (graze bluegreen algae)

Sprawlers

Sprawlers

Scrapers (graze bluegreen algae)

Sprawlers

Habit

Trophic Relationships

Generally

depositional (sediments)

Lotic—

hydrophytes

Lentic—vascular

margins

Lotic and lentic—

margins

Lotic and lentic—

margins

Lotic and lentic—

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW : ^ Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Order

of species in parentheses)

Taxa (number

Table 23B

North

Northwest

Pacific

Northwest

Pacific

East, South

East

Widespread

West, Midwest

Widespread

Widespread

Distribution

American SE

UM

M

NW

Tolerance Values

(continued)

3801, 3855

1613, 3936, 3854, 6188

5310

5310, 6449

5310

1910, 1913,

1915

5310

1910, 1912,

MA* References**

Ecological

J ) J > )J J J ):) ) )3 3 J J 3 3 ) ) ) 3 ) ) ) 3 3

Muscidae (271) (=Anthomyiidae, in part, by some authors) - House Flies, Stable Flies, Green Bottle Flies

Family

Lotic—erosional

Lotic—

Spilogona (135)

lentic—littoral

depositional;

Lentic—tree holes

Lotic—erosional

depositional (margins); marine shores (algae)

lotic—

Lentic—littoral;

(especially mosses)

Phaonia (81)

Lispoides

Lispe(25)

Limnophora (10)

depositional (small enriched ponds)

Lentic—

erosional

depositional,

littoral: lotic—

Generally lentic—

Habitat

Graphomya (9)

aequifrons

Species

Lotic—erosional?

Genus

Carlcea (10) {=Lispocephala)

Continued

Habit

Sprawlers?

Sprawlers

Borrowers

Sprawlers

Generally sprawlers

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Order

of species In parentheses)

Taxa(number

Table 23B

(piercers) on mosquito larvae

Predators

(piercers)

Predators

Tipulidae

Simuliidae,

(piercers) on Oligochaeta,

Predators

(piercers)

Predators

(piercers)

Predators

Generally predators (piercers)

Trophic Relationships

North

Ecological

1782, 2526, 5473, 5491

6677

2489, 2526, 2975, 6604, 1782, 5491,

6492

2223, 2526, 2975, 3720, 1782, 4016, 5070, 6677, 5491, 6750,

1782, 5491

6677

1782, 6604,

3095

2937, 2975, 3179, 4599, 1782, 5490, 5829, 6677,

MA* References**

Widespread

6.0

NW

1782, 2526, 5909, 6677

M

2489, 5473

UM

Widespread

7.0

SE

Widespread

Widespread

Widespread

Widespread

Widespread

Distribution

American

Tolerance Values

) ) ) J ) ) ) ) ) ) ) ) ) ) ))))) ) ))) ) ) )

a\

o

-4

o

Anthomyiidae, in part)Dung Flies

Cordiluridae

Scathophagidae (53) (=Scatophagldae, in part, Scopeumatidae,

Fletcherimyia (6) (=Blaesoxipha, in part)

Generally burrowers—

miners (in plant stems); sprawlers

Generally lentic— hydrophytes; lotic— depositional

miners (bases of pitcher plants)

vascular

Burrowers—

(pitcher plants)

Burrowers

Burrowers

Habit

Lentic—littoral

miners

Burrowers—

filter beds

detritus), trickling

(sediments and

Lentic—littoral

beds)

(trickling filter

Lotic—erosional

Habitat

Semiaquatic

cornuta

Species

Flesh Flies

Megaselia (spp.?)

Dohrniphora

Genus

Sarcophagidae (7)-

Phoridae (2) - Coffin Flies, Scuttle Flies

Family

Continued

DIPTERA

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Order

of species in parentheses)

Taxa(number

Table 23B

predators (engulfers)

(miners);

herbivores

shredders—

Generally

rotifers

predators on

invertebrates);

Collectors— gatherers (scavengers on trapped

gatherers

Collectors—

gatherers

Collectors—

gatherers: predators (engulfers) of Psychodidae

Collectors—

Trophic Relationships

North

Widespread

Widespread

Widespread

Distribution

American

SE

UM

M

NW

Tolerance Values

(continued)

2937, 2975, 3179, 4599, 1782, 6185, 6677, 3095

1831, 1832, 1940, 541, 1312, 1888

4504, 3095

1002, 1446

1446, 3178

MA* References**

Ecological

J ) J j > ) 3 ) 3 :) )3 3 > ))3 ) 3 3 3 3 3 ) ) 3 3

Marsh Flies

inside snails

depositional in snails

Sprawlers

stems)

Snail-Killing Flies,

(sewage beds in oxidation ponds)

Lentic—littoral

Generally burrowers,

dncta

miners (plant

snails

Generally predators (engulfers) or "parasites" of

gatherers

collectors—

Scrapers;

(engulfers)

Predators

roots)

Burrowers—

petioles of Nuphar, roots of Potamogeton)

stems and

hydrophytes (emergent zone)

(miners in

miners (plant

Lentic—vascular

Shredders— herbivores

Burrowers—

Lentic—vascular

Carex); predators (engulfers) of Ceratopogonidae

Juncus, and

(miners Scirpus,

herbivores

Shredders—

Trophic Relationships

hydrophytes (submerged and floating zones)

Generally lentic— littoral; lotic—

Spaziphora

Orthacheta (3)

confluens

miners (plant stems)

Lentic—vascular

hydrophytes (emergent zone)

Cordilura (44)

Hydromyza

Borrowers—

Lotic—

Habit

depositional

Habitat

Acanthocnema

Species

(4)

Genus

(=Tetanoceridae, Tetanoceratidae) -

Sciomyzidae (175)

Family

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Order

of species in parentheses)

Taxa (nunriber

Table 23B

Widespread

Widespread

Northeast

Widespread

East

American Distribution

North

SE UM

M

6.0

NW

Tolerance Values

Ecological

3095

1782,3212,

1782, 3034

4294

452, 3944, 4271, 4599, 3798, 5799, 6402, 6405

6185, 6267, 6677, 6868

1995, 1996, 2975, 4293,

MA* References**

) ) ) ) ) ) ) ) ) ) ) ) ) V )))) ) ) ))) r) )

00

Family Genus

americana

Colobaea

arcuata

mixta

Euthycera Hedria

Eigiva (2)

Dictyadum (2)

Dictya (25)

pubera

Species

Atrichomelina

Antichaeta (8)

Continued

Burrowers—

inside snails

Lentic—^vascular

hydrophytes (emergent zone); lotic—margins

Burrowers—

inside snails

Lentic—vascular

hydrophytes (emergent zone); lotic—margins

Burrowers— inside snails

hydrophytes (emergent zone)

DIPTERA

snails

(engulfers) or "parasites" of

Predators

snails

(engulfers) or "parasites" of

Predators

snails

(engulfers) or "parasites" of

Predators

gatherers (scavengers on decaying snail)

collectors—

snails; some

Predators

(engulfers) or "parasites" of

Burrowers—

inside snails

Lentic—vascular

Lentic—littoral

Lentic—littoral

snails

masses and

temporary ponds) snails

Predators

(engulfers) or "parasites" of snail eggs and

Burrowers—i

snail egg

Lentic—littoral

Habit

Trophic Relationships

(marshes and

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Order

of species In parentheses)

Taxa (number

Table 23B

North

North

East, Midwest

North

Midwest

North,

Widespread

States

Eastern United

Widespread

(continued)

453, 454, 1908

453, 3214, 4470, 6677

456, 6119

3215

1923,3210,

3179

453, 454, 455, 457, 456, 1837, 2937, 2975,

North, West

Ecological References**

Distribution

American

Tolerance Values

Family Genus

fingernail

temporary ponds)

clams

Borrowers—

inside

Lentic—littoral

(engulfers) or "parasites" in fingernail clams (Sphaeriidae)

Predators

snails

Predators

(engulfers) or "parasites" of

Borrowers—

snails

(engulfers) or "parasites" of

Predators

snails

(engulfers) or "parasites" of

Predators

Littorina

(engulfers) or "parasites" of snails; 1 sp. in

Predators

Trophic Relationships

inside snails

(marshes and

Renocera (4)

(bogs)— semiaquatic

Lentic—littoral

flats

marine—mud

Borrowers—

inside snails

Lentic—littoral

(margins);

(bogs)

Lentic—littoral?

decora

hydrophytes (emergent zone)

Lentic—littoral

Borrowers—

inside snails

Lentic—vascular

marshes

Borrowers— inside snails

marine—salt

Habit

Lentic—littoral;

Habitat

Pteromicra (14)

Poecilographa

Pherbellla (40)

ferrugineus lemenitis

Oidematops

Species

Pherbecta

Limnia (17)

Hoplodictya (5)

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Order

Taxa (number of species in parentheses)

Table 23B

North

Widespread

Midwest, East

Widespread

North

East

Widespread

Widespread

Distribution

American

North SE

UM

M

NW

Ecological

1921

453, 1909,

6677

2975, 5183,

2975, 6677

3213

656, 1907,

3211

5748

4291

MA* References**

Tolerance Values

) ) i ) ) ) ) ) j ) ) ) ) > ) ) ) ) ) ) ) > ))))

O

o

o

Flower Flies, Rattail Maggots

Syrphidae (125) -

Family Genus

Borrowers

Collectors—

Lentic—littoral

hydrophytes)

and vascular

(pond margins

DIPTERA

Borrowers

gatherers

Widespread

Widespread

Widespread

East, Southwest

Widespread

North

Widespread

Widespread

Southwest

Distribution

gatherers

Collectors—

Lentic—tree holes

Chalcosyrphus

North American

Collectors—

Chrysogaster(10)

Lentic—tree holes

Ceriana (5)

Borrowers

gatherers

Lentic—tree holes

(engulfers) or "parasites" of snails(and slugs)

Predators

snails

(engulfers) or "parasites" of

Predators

(engulfers) or "parasites" of snails (and slogs)

Predators

Trophic Relationships

(27)

Lentic—tree holes

Borrowers—

inside snails

Lentic—vascular

hydrophytes (emergent zone)

Borrowers—

inside snails

Lentic—littoral

(including temporary ponds)

Borrowers—

inside snails

hydrophytes (emergent zone)

Habit

Lentic—vascular

Habitat

Cailicera (4)

canadensis

macropus

Species

Blera (16)

Trypetoptera

Tetanocera (30)

Sepedon (16)

Sepedomerus

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Order

of species in parentheses)

Taxa (number

Table 23B

SE UM

M

10.0

NW

Tolerance Values

(continued)

2445, 2975, 3439, 6677

1782

1782, 5161

2445, 2937, 2975, 3179, 1782, 4599, 5069, 6677, 3678, 6189

453, 454, 2975, 5179, 3218, 6045, 6677, 1919, 1920

453, 4292

454

MA* References**

Ecological

3 )))3 D ) ) 3 )) ) 1 )) ) ) )) ) ))) ) ) )

©

Family Genus

Lentic—littoral Collectors—

Spilomyia (11)

Sericomyia (11)

Lentic—littoral

Palpada (10)

Lentic—^tree holes

Collectors—

gatherers

Lentic—littoral

gatherers

Collectors—

(bog mat pools)

(sediments and detritus)

Lentic—tree holes

Orthonevra (16)

marshes)

Predators—

engulfers

Lentic—littoral

(margins,

Neoasda (10)

gatherers

Collectors—

gatherers

Collectors—

gatherers

Lentic—tree holes

Borrowers

Borrowers

Myolepta (7)

(detritus and organic sediments)

substrates)

Collectors—

gatherers

Borrowers

(especially low 02 organic

gatherers

Collectors—

Trophic Relationships

Lentic—littoral

Borrowers

Habit

(sediments and detritus): lotic— depositional

(sediments and detritus)

Lentic—littoral

Habitat

Lentic—^tree holes

aeneus

Species

Mallota (9)

Helophilus(10)

Eristalis(18)

Eristalinus

Continued

"SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ** Emphasis on trophic relationships

Order

of species in parentheses)

Taxa (number

Table 23B

Widespread

Widespread

South

Widespread

Widespread

Widespread

Widespread

Widespread

Widespread

Widespread

Distribution

American

North

10.0

SE

UM

0.0

M

NW

Tolerance Values

Ecological

1782

478, 2225, 2445, 6677

6677

1782

3675

478, 3600,

478, 2225, 2445, 6677

478, 1782, 2445, 2975, 6677, 3677

1782

478, 2445, 2975, 6677,

6677

2225, 2975, 4040, 6605,

6492

MA* References**

) ) ) ) 1 ) ) ) 1 ) ) ) J ) )) ) ) ) ) ) ) ) )))

K»

~

f^.t-' /Y-A

Figure 25.47

Figure 25.47 Lateral view of culicine trumpet. Figure 25.48 Dorsal view of culicine pupal terminal abdominal segments. Figure 25.49 Lateral view of culicine trumpet.

1086

Chapter 25 Culicidae

setaS-C

seta 9-C

Figure 25.50

seta 9-Vlll'

tracheation

■jH— Figure 25.53 5-V I 9-VIII

paddle midrib

Figure 25.51

Figure 25.52

>— flared apex

sclerotized ring tracheation

seta 9-VIII

Figure 25.54

Figure 25.55

Figure 25.50 Lateral view of culicine trumpet. Figure 25.51 Dorsal view of Haemagogus sp. paddles and abdominal segments. Figure 25.52 Lateral view of Culex sp. trumpet. Figure 25.53 Dorsal view of segment VIII and paddles of Culex. Figure 25.54 Lateral view of Culiseta sp. trumpet.

Figure 25.56

Figure 25.55 Lateral view of Aedeomyia trumpet with sclerotized ring at base and flared apex (adapted by Lindsay Matter from ptiotographic image by Stephen Doggett - with permission). Figure 25.56 Dorsal view of segment VIII and paddles of Culiseta sp.

Chapter 25 Culicidae

1087

Adults

1.

Proboscis rigid and stout on basal half, apical half tapered toward apex and strongly curved downward (Fig. 25.57). Large species(up to 15 mm long), with broad,

r.

metallic colored scales on head, thorax, abdomen, and legs Proboscis slender, never strongly curved downward on apical half, and of nearly uniform thickness (Fig. 25.58)

2(1').

Toxorhynchites 2

Scutellum evenly rounded on posterior margin (Fig. 25.59); palpi of female nearly as long as proboscis (Fig. 25.60); abdomen bare of scales or only sparsely scaled Anopheles 2'. Scutellum trilobed on posterior margin (Fig. 25.61); palpi less than one-half as long as proboscis (Fig. 25.62); abdomen densely scaled both dorsally and ventrally 3 3(2'). Second marginal cell of wing short, less than half as long as its petiole (Fig. 25.63) .... Uranotaenia 3'. Second marginal cell of wing as long as or longer than its petiole (Fig. 25.64) 4 4(3'). Postnotum with a tuft of setae (Fig. 25.65); wing squama without fringe of hair (Fig. 25.66) Wyeomyia 4'. Postnotum bare (Fig. 25.67); wing squama with fringe of hair (Fig. 25.68) 5 5(4'). Spiracular bristles present(Fig. 25.69) 6 5'. Spiracular bristles absent(Fig. 25.70) 7 6(5). Postspiracular bristles present(Fig. 25.71); tip of abdomen pointed (Fig. 25.72) Psorophora 6'. Postspiracular bristles absent(Fig. 25.73), tip of abdomen blunt(Fig. 25.74) Culiseta 7(5'). Anterior pronotal lobes large, collar-like, almost joining dorsally (Fig. 25.75); abdominal scales bright metallic violet or silver Haemagogus 7'. Anterior pronotal lobes small, widely separated dorsally (Fig. 25.76); abdomen without bright metallic scales 8 8(7'). Postspiracular bristles present(Fig. 25.71) 9 8'. Postspiracular bristles absent(Fig. 25.73) 10 9(8). Wing scales very broad, mixed brown and white (Fig. 25.77); tip of abdomen blunt (Fig. 25.74) MansonialCoquillettidia 9'. Wing scales narrow (rarely moderately broad)(Fig. 25.78); tip of abdomen pointed (Fig. 25.72) Aedes 10(8'). Antenna much longer than proboscis, 1st flagellar segment longer than the next 2 segments combined (Fig. 25.79) Deinocerites 10'. Antenna not longer than proboscis, or only slightly so, 1st flagellar segment about as long as each succeeding segment(Fig. 25.80) 11 11(IO'). Scutum bicolored with narrow longitudinal lines of white scales (Fig. 25.81); penultimate (next to the last) segment of front tarsi very short, only about half as long as wide (Fig. 25.82) Orthopodomyia 11'. Scutum without longitudinal lines of white scales (Fig. 25.83); penultimate segment of front tarsi much longer than wide (Fig. 25.84) 12 12(11'). All adult females with short, thick antennal flagellomeres (Fig. 25.85); large, apical tuft of scales on the mid and hind femur (Fig. 25.86); wings covered with yellow, white and brown scales Aedeomyia 12', Normal length antennal flagellomeres(Fig. 25.80); wing scales narrow and uniformly dark (Fig. 25.78) Culex

1088

Chapter 25 Culicidae

scutellum

proboscis

Figure 25.57 Figure 25.59 palpus

Figure 25.60 proboscis

proboscis

Figure 25.58

proboscis

scutellum

palpus

Figure 25.62

Figure 25.61

second marginal cell

petiole

Figure 25.63

Figure 25.57 Lateral view of head showing

Figure 25.60 Dorsai view of head showing Anopheles

Toxorhynchites sp. proboscis. Figure 25.58 Lateral view of head showing typical cuilcine proboscis. Figure 25.59 Dorsai view of thorax showing Anopheles sp. scutellum (adapted by Zel Stoltzfus from Darsie and Ward 2005).

sp. palpi.

Figure 25.61

Dorsal view of thorax showing typical

culicine scutellum (adapted by Zel Stoltzfus from Darsie and Ward 2005).

Figure 25.62 Dorsal view of head showing typical culicine palpi.

Figure 25.63 Uranotaenia sp. wing.

Chapter 25 Culicidae

1089

second marginal cell

petiole

Figure 25.64

squama

Figure 25.66 abdomen

Figure 25.65

postnotum

abdomen

squama with fringe

Figure 25.67 Figure 25.68 spiracular bristles absent spiracular bristles present

scutum

scutum

Figure 25.69 Figure 25.70

Figure 25.64 Typical culicine wing. Figure 25.65 Dorsal view of Wyeomyia sp. postnotum. Figure 25.66 Wyeomyia sp. squama. Figure 25.67 Dorsai view of typical culicine postnotum.

Figure 25.68 Typical culicine squama. Figure 25.69 Lateral view of cuiicine thorax. Figure 25.70 Lateral view of culicine thorax, spiracular bristles absent.

1090

Chapter 25 Culicidae

spiracular bristles

postspiracular

tip of abdomen

bristles

Figure 25.72

pronotal lobes

Figure 25.71

postspiracular bristles absent

Figure 25.76 Figure 25.73

anterior pronotal lobe

Figure 25.75

tip of abdomen

Figure 25.74

wing scales

Figure 25.77

Figure 25.71 Lateral view of culiclne thorax, showing postspiracular bristles present. Figure 25.72 Dorsal view of pointed culiclne abdomen. Figure 25.73 Lateral view of culiclne thorax. Figure 25.74 Dorsal view of blunt culiclne abdomen.

Figure 25.75 Dorsal view of pronotal lobes of Haemagogus sp. Figure 25.76 Dorsal view of culicine thorax. Figure 25.77 Enlarged view of Mansonia sp. broad wing scale, mixed brown and white.

Chapter 25 Culicidae

1091

proboscis

1st flagellar segment wing scales

Figure 25.78

Figure 25.79

proboscis

• antenna

scutum

scutum

1st ffageliar segment

Figure 25.81

Figure 25.83

Figure 25.80 penultimate segment .

penultimate segment

Figure 25.84

Figure 25.82

Figure 25.78 Enlarged view of cullcine narrow wing scales.

Figure 25.79 Dorsal view of head showing Deinocerites sp. antenna.

Figure 25.80 Dorsal view of culiclne head showing typical mosquito antenna.

Figure 25.81 Dorsal view of Orthopodomyia sp. scutum (adapted by Zel Stoltzfus from Darsie and Ward 2005). Figure 25.82 Orthopodomyia sp. tarsus. Figure 25.83 Dorsal view of culicine scutum (adapted by Zei Stoltzfus from Darsie and Ward 2005). Figure 25.84 Typical culiclne front tarsus.

1092

Chapter 25 Culicidae

tuft of scales

thickened antenna!flageliomeres

Figure 25.86

Figure 25.85

Figure 25.85 Lateral view of female Aedeomyia with short, thickened antennal flageliomeres (adapted by Lindsay Matter from APHC 2016).

Figure 25.86 Lateral view of hind femur of Aedeomyia with large, apical tuft of scales (adapted by Lindsay Matter from APHC 2016).

ADDITIONAL TAXONOMIC REFERENCES

North Carolina: Slaff and Apperson (1989); Harrison et al. (1998). Northwestern United States: Gjullin and Eddy (1972).

General Lane (1953); Carpenter and LaCasse (1955); Breland (1958); Darsie

Oklahoma: Rozeboom (1942). Southeastern United States: King et al.(1960).

and Ward (1981, 1989), Stone (1981); Darsie (1995); Sallum and Forrattini (1996); Teng and Apperson (1996); Reinert

Texas: Randolph and O'Neill (1944); Reeves and Darsie (2003).

et al. (1997); Andreadis and Munstermann (1997); Eldridge et al. (1998); Harbach and Kitching (1998); Moore (1999); Ginnig and Eldridge (1999); Darsie and Ward (2000); Ginnig (2000); Reinert(2000a); Reinert(2000b); Fonseca et al. (2001); Reinert(2001); Darsie et al.(2002); Savage and

Virginia: Gladney and Turner (1969). Washington: Sames and Pehling (2005).

Strickman (2004); Reinert et al(2004); Darsie and Ward (2005); Edman (2005); Wilkerson et al(2015); BurkettCadena and Blosser (2017). http://www2.inbio.ac.cr/papers/ culicidaejarvas/intro.html. http://mosquito-taxonomic-

inventory.info/simpletaxonoray/term/6062. APHC- https://phc.amedd.army.mil/PHC%20Resource%20 Library/TG369_AFRICOMMosquitoKey.pdf(2016)

Regional faunas Alaska: Gjullin etal.(1961). Arizona: McDonald et al.(1973). Arkansas: Carpenter (1941). British Columbia: Curtis (1967).

California: Bohart and Washino (1978); Meyer and Durso (1993); Wekesa etal.(1996) Canada: Wood et al. (1979). Colorado: Harmston and Lawson (1967). Connecticut: Andreadis et al. (2001). Florida: Breeland and Loyless(1982); Ansell,(1994, 1995); Betts (1994); O'Meara et al.(1997); Darsie and Shroyer (2004). Illinois: Ross and Horsfall (1965); Lampman et al.(1997); Jenson et al.(1999). Indiana: Siverly (1972). Iowa: Knight and Wonio (1969). Maryland: Sardelis and Turell (2001). Massachusetts: Dennehy and Livdahl (1999). Michigan: Wilmot et al. (1990). Minnesota: Barr (1958). Montana: Quickenden and Jamison (1979). Nebraska: Janousek and Kramer (1999); Moore (2001). New Jersey: Headlee (1945); Crans and Crans(1998); Peyton et al. (1999). New York: Means(1979, 1987); Oliver et al.(2003); Kokas and Lee (2005).

Utah: Nielsen and Rees (1961).

Wisconsin: Dickinson (1944).

Wyoming: Denke et al.(1996); Nielsen and Blackmore (1996); Moore (2001).

Regional species list Arkansas: Jamieson et al.(1994). Colorado: Harmston (1949). Connecticut: Andreadis(2001); Andreadis et al.(2005). Delaware: Darsie et al (1951). District of Columbia: Good (1945). Florida: O'Meara and Evans(1997); Darsie and Ward (2000). Idaho: Brothers (1971). Kentucky: Quinby et al (1944). Manitoba: Trimble (1972). Maryland: Bickley eta/. (1971).

Michigan: Cassani and Newson (1980). Mississippi: Harden et al (1967); Goddard and Harrison (2005). Nebraska: Moore (2001).

New Jersey: Crans and McCuiston (1999). New Mexico: Sublette and Sublette (1970). New York: Jamnback (1969). North Dakota: Darsie and Anderson (1985). Northeastern United States: Stojanovich (1961). Ohio: Parsons et al.(1972). Oklahoma: Parsons and Howell (1971). Ontario: James et al. (1969). Pennsylvania: Wilson et al.(1946). South Dakota: Gerhardt(1966). Southeastern United States: Stojanovich (1961). Texas: Sublette and Sublette (1970). Utah: Nielsen (1968).

Virginia: Dorer et al (1944). Vermont: Graham et al.(1991).

West Virginia: Amrine and Butler (1978); Joy et al (1994). Wyoming: Nielsen and Blackmore (1996); Moore (2001).

o VO

i

J ) ) )) ) ) > )) ) J ) ) )

Family Genus

Species

back-waters)

gatherers

filterers and

swimmers

(limnetic);

lotic— depositional (pools and

Generally collectors—

Habit

Generally Generally lentic—littoral planktonic—

Habitat

Trophic Relationships

■* Emphasis on trophic relationships

' SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic

(173)

Culicidae

Diptera - Mosquitoes

Order

(number of species in parentheses)

Taxa

E. D. Walker, Fl. D. Newson, R. W. Merritt, J. Davis, E. Fligh, and M. Flutchinson.)

North

Widespread

Distribution

American SE

UM

M

MA*

Ecological

(continued)

914,2147, 2372, 2937, 3179,4599, 5831, 6266, 5402, 6496, 6711, 1054, 1311,6231, 6495, 19,4114, 6230, 1954, 1027, 3363, 3580,4005, 4032, 179, 6619,4498

References**

CULICIDAE

NW

Tolerance Values

Table 25A Summary of ecological and distributional data for Culicidae (Diptera).(For definition of terms see Tables 6A-6C; table prepared by J. Wallace,

J

Family

Anopheles(22)

Aedes(87)

Genus

Continued

Species gatherers and filterers

(divers,

feeding in many habitat

(temporary ponds and pools)

planktonicswimmers

(limnetic);

deposltional

lotic—

Collectorsfilterers

Neustonic/

Lentic— littoral

zones)

Collectors—

Swimmers

Lentic

Habitat

** Emphasis on trophic relationships

North

Widespread

Widespread

Trophic American Relationships Distribution

* SE = Southeast, UM = Upper Midwest, M = Midwest, NW — Northwest, MA : : Mid-Atlantic

Order

parentheses)

(number of species in

Taxa

Table 25A

9.1

Tolerance Values

323, 401, 914, 1053,2147, 2734, 3148, 4833, 4930, 4017, 5831, 6634, 5402, 6711, 120, 5538,6279, 6280, 6278, 6400,6205, 6.0

5430

131, 914, 1051, 1447, 2147, 2665, 3148,4030, 4833, 3222, 5831, 6229, 5402, 6711, 1027, 124, 316, 1579, 2249, 3497, 3577, 4253, 1187,4346, 2707, 5430 8,0

vo Ch

Family

Mansonia (2) {Coquillettidia)f

Haemagogus

Deinocerites(3)

Culiseta (8)

Culex(30)

Continued

equines

Species CoNectors-

filterers

roots of

hydrophytes

plants. piercing respiratory siphon)

Collectors—

gatherers and

Clingers (stems and

Lentic—

t {Coquillettidia is used as the genus of Mansonia in some species level keys)

** Emphasis on trophic relationships

Widespread

Southern Texas

Collectors—

Extreme South

Widespread

Widespread

filterers

vascular

Lentic (tree holes) Planktonic

marine

intertidal

Collectors—

filterers

Planktonic

(crab holes)

Beach zone—

filterers

gatherers and

(ponds and divers ground pools) (feeding at bottom)

Collectors—

Planktonic—

swimmers.

Lentic—

filterers

limnetic

ground pools)

ditches, and

(lakes, ponds,

Planktonic—

swimmers

Lentic—

Habit

limnetic

Habitat

North

Trophic American Relationships Distribution 10.0

8,0

Ecological

(continued)

6711

914, 2147, 3148, 5831, 5402, 6677,

661

3148, 5402

914, 2147,

877, 2147, 2665, 3148, 4820, 4833, 5402, 6711, 2672, 3673

914, 1305, 1306, 1307, 1308, 2147, 2319, 3148, 4833, 5395, 5402, 5831, 6711, 6400, 3578, 6205, 558, 794

References**

CULICIDAE

Tolerance Values

» ; r ) ) ) ) ) ) ) ) ) )) ) ) i ) )

* SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA : : Mid-Atlantic

Order

species in parentheses)

(number of

Taxa

Table 25A

j

o\

o

Family

Lentic

Wyeomyia (3)

Uranotaenia (4)

Collectors—

in living and dead plants)

pitcher plants

plants)

and dead

pools and water in living

and small

** Emphasis on trophic relationships

J

SB

UM

M

NW

Tolerance Values

914, 1832, 2147, 2862, 19, 3148, 5395, 1953, 5402, 6711, 4254, 649

914, 2147, 3148, 5831, 5402, 6711

914, 2147, 3148, 5831, 5402, 6677, 3548, 4253

6711

914, 2147, 3148, 5831, 5402, 6677,

558, 794

2371

3148, 6711,

914, 2147,

Ecological MA* References**

i ) ) ) ) ) ) ) ))) ^ ) ) ) ) ) )

Southeast

East, Collectorsmicrohabitats

mats in

filterers

Planktonic (in

Lentic (bog

West and

intermountain)

Widespread (except for

East, South

Widespread

South Florida

East, South

(ponds and ground pools)

Collectors-

filterers

Planktonic— swimmers

(engulfers)

Predators

filterers

collectors—

(piercers):

Predators

filterers

Collectors—

filterers

Lentic—

Planktonic

North

Trophic American Relationships Distribution

limnetic

phytotelmata)

other

containers,

water

artificial

holes,

Lentic (in tree and rock

Planktonicswimmers

Lentic

Planktonic

Planktonic

Habit

(temporary ponds and pools)

(ditches and canals)

Toxorhynchites

squamipennis

(3)

Psorophora (12)

Aedeomyia

containers)

water

artificial

wooden

Lentic (tree holes and

Habitat

Orthopodomyia

Species

(3)

Continued

* SE = Southeast, UM = Upper Midwest, M : : Midwest, NW = Northwest, MA = Mid-Atlantic

Order

species In parentheses)

(number of

Taxa

Table 25A

t

w

«^-»N .4'%^"^^'^ te^S

V' ■

SIMULIIDAE Peter H. Adler

Ckmson University, Clemson, South Carolina

Douglas C. Currie Department of Natural History, Royal Ontario Museum and Department ofEcolopfy &Evolutionary Biology, University of Toronto, Ontario, Canada

INTRODUCTION

Blaek flies are among the most widespread and ubiquitous macroinvertebrates in flowing waters. They also are among the most abundant; the greatest secondary production for any stream macroinvertebrate has been recorded for black flies below lake

outlets. In the terrestrial stage, the females are well known for their pestiferous habits of swarming about people and animals, often biting to acquire blood for egg maturation. About 256 species of black flies have been recorded from North America north of Mexico,

representing roughly 11% of the approximately 2,300 named species in the world. Eggs of black flies are deposited in or near run ning water. They generally cannot withstand desicca

tion and perish within a few hours of drying. The eggs of many species, however, experience dry streambeds, but presumably are protected in moist sediments of the stream bottom. Eggs of some species hatch in a few days, whereas those of other species undergo an obligatory diapause and do not hatch for 6 months or more. Hatching is triggered by temperature, oxygen tension, and perhaps photoperiod. Escape from the egg is aided by an egg burster on the head of the flrst-instar larva.

Larval black flies are found exclusively in flowing freshwaters, including seepages, large rivers, sulfur springs, hot springs, glacial meltwatcrs, slow-flowing swamps,impoundment outflows, waterfalls,intermit tent streams, and subterranean Hows. Black flies are

often viewed as intolerant of pollution, but a few species do well in polluted flows such as those that are organically fouled. An average ofthree to four species generally inhabit a stream site at any one time despite the presence of many species in the surrounding geographic area; rarely, as many as 10 species can occupy a short section of stream. The areas of North

America with the greatest elevational variation provide the widest diversity of habitats for black flies and are richest in species. Among the most important factors influencing larval development is temperature. Each species has an optimal range of water temperature. Temperatures above 30°C, however, rarely provide a suitable ther mal regime for development. The larvae of some spe cies develop through the winter, even beneath ice and snow. Species that are better adapted to a broad range oftemperatures can complete as many as seven gener ations per year in southern areas of the continent. These multivoltine species belong to the genus Simulium. About 63% of North American species are univoltine, completing a single generation annually. Larvae anchor themselves to the substrate by means of tiny hooks on their posterior proleg, which are enmeshed with a silken pad spun from the silk glands. The larvae distribute themselves evenly, ran domly, or in bands and clumps, depending on age, species, and environmental factors. The majority of larval life is dedicated to feeding, and these aggrega tion patterns often optimize water velocities for filter-feeding. During filter-feeding, larvae open their labral fans to the current and twist their bodies 90-180°.

Larvae have been referred to as ecosystem engineers because they filter fine particulate matter from the water column and subsequently egest larger pellets that can be used as food by the benthic microbial and invertebrate communities. Though conventionally viewed as filter feeders, larval black flies practice a variety of additional feeding modes, including graz ing, predation, deposit feeding, and engulfing of algal filaments, each of which can provide a substantial means of obtaining food, even in species with fully developed labral fans. Ingested particles range from

1097

1098

Chapter 26 Simuliidae

0.09 to 350 litn in diameter and include bacteria, dia toms, leaf fragments, pollen, fecal pellets, protozoa, and minute arthropods, as well as inorganic matter. Larvae can develop to adults on a diet of bacteria, and in some systems, protozoa make up a significant part of the diet. Larvae also can use surface films and dis

solved organic matter that has flocculated. Filtering efficiency is influenced by temperature, current veloc ity, particle size and concentration, labral-fan struc ture,and parasitism. Most species can be reared easily from eggs or larvae to adults if adequate food and aeration are provided. Larvae move to new sites by looping inchworm fashion or by drifting downstream, often on silk strands, especially around dusk and during the night. Drift is caused by foraging predators, ultraviolet radi ation, changes in discharge, and anchor ice lifting larvae from the substrate as water temperature

increases. During floods and other unfavorable con ditions, larvae move into protected areas such as the hyporheic zone. The population dynamics of black flies are affected by abiotic and biotic factors such as tempera ture and the quality and quantity of food, as well as natural enemies and other symbiotic organisms. Larvae are hosts of numerous symbiotes, including

bacteria, fungi, helicosporidia, ichthyosporeans, mermithid nematodes, microsporidia, nematomorphs, protists, stramenopiles, and viruses. Predation by invertebrates and vertebrates, particularly birds and fish, is opportunistic and often intense. Ectoparasitic water mites inhabit the pupal cocoons, where they await emergence ofthe adult flies on which they will feed. Black flies typically have seven larval instars, although within a species, the number can vary from 6 to 11 in relation to environmental factors such as

food availability and parasitism. Within about a week to nearly a year after hatching, depending on water temperature and species, the larva begins its molt to the pupa. During the molting process, the pupa remains hidden in the larval skin and is referred to as

pharate. The pharate pupa retains the larval appear ance,continues to feed,locates a suitable pupation site

in the water, scrapes the pupation area clean, spins the cocoon, and finally sheds the larval cuticle. Male lar vae generally develop faster than female larvae and pupate and emerge a few days before females. The pupal stage usually lasts a few days to a few weeks. Adults typically emerge from the pupal skins in the morning, and are active during the day. Mating takes place shortly after emergence. Four North American species lack males and are obligately parthenogenetic. At least six North American species

couple on the ground and all but one of these are associated with northern environments or high eleva tions. Most sexual species probably include aerial coupling in their mating behavior. Males form loose aggregations over or beside a landmark, such as a waterfall or riparian vegetation, where they intercept females; the coupled pair flies out of the swarm or drops to the ground. The females of North American species can disperse up to 225 km in search of blood, although distances of less than 15 km are probably typical of most species. Females subsequently seek an appropri ate habitat for oviposition, depositing their eggs during flight over water or while landed on a wetted stone or trailing vegetation. Oviposition generally occurs under low illumination, especially toward the end of daylight. The number of eggs matured in one ovarian cycle varies from 20 in northern species to more than 800 in more temperate species, with an average of 150-600 eggs per cycle. Females of some species might undergo up to six cycles of egg matura tion. Adults usually live 10-35 days in nature. Water,sugar, and blood constitute the adult diet. Most adults probably imbibe water and feed on sugar, which provides energy for flight and other activities. Sugar is acquired from plant nectar and the honeydew of aphids and related insects. Only females suck blood, with about 90% of North American species being capable of taking blood. Species incapable of sucking blood live at high elevations and northern latitudes, and are obligately autogenous, developing eggs without benefit of blood. Some species are facul tatively autogenous, maturing at least the first batch of eggs without feeding on blood. Most species are probably anautogenous for at least one ovarian cycle, requiring blood to produce their eggs. Blood is acquired exclusively from birds and mammals. To locate and feed on appropriate hosts, females evaluate a series of habitat features and host attributes

such as size, shape, color, odor (especially carbon dioxide), temperature, and various phagostimulants.

Through their blood feeding, black flies can transmit the agents of various filarial, protozoan, and viral diseases to domestic animals and wildlife. The more common diseases in North America are bovine oncho-

cerciasis in cattle, leucocytozoonosis in birds, and vesicular stomatitis in various domestic mammals. In

North America, black flies are not responsible for the transmission of any causal agents of human diseases. Reviews of the behavior, ecology, natural his tory, sampling, and medical-veterinary importance of black flies are provided by Laird (1981), Kim and Merritt(1988),Crosskey(1990),Adler and McCreadie (2002), Adler et al.(2004), and Adler (2005).

Chapter 26 Simuliidae

1099

EXTERNAL MORPHOLOGY

Pigmentation patterns of the antennae provide useful

Larval black flies are recognized by their external head capsule, typically bearing a pair of labral fans, and an elongate body with a single posterior proleg. The silken cocoon and thoracic gills are distinctive features of the compact pupae. Adult black flies are recognized by their distinctive wing venation,antennal shape, and arched thorax. Morphological details and illustrations ofall life stages of black flies are provided by Adler et al. (2004). The structural uniformity of the family Simuliidae, while peiTnitting ready identification of any black fly as a member of the family, can frustrate efforts to identify species and even genera. Further complicating

taxonomic characters.

the taxonomy and identification of black flies in all life

stages is the existence of cryptic (or sibling) species— structurally similar, if not identical, species with unique ecologies. These cryptic species are often most reliably identified by analysis of the giant polytene chromosomes in the larval silk glands(Adler et al. 2004).

Eggs The eggs are rather oval in outline, especially when viewed dorsally, but in lateral view, they are often somewhat triangular with rounded angles. Their color changes from whitish when freshly laid to brown before the larvae hatch. Egg size among species varies

about fourfold (0.15-0.54 mm). The largest eggs are found in autogenous, northern species that produce relatively few eggs. The egg has an exochorion, which is an outer adhesive, gelatinous layer. When the exo chorion is removed with a clearing agent, the shell or endochorion is revealed. It has little surface sculpture other than minor pitting. A micropyle is located at the blunt end of the egg, allowing the entry of sperm. In

some species, a micropyle is absent, the sperm proba bly entering the egg enzymatically. Larvae

Larvae have a well-sclerotized, external head cap sule and an elongate body (3-15 mm), slightly expanded posteriorly, with one prothoracic proleg and one posterior proleg (Fig. 26.1). The head bears some of the most useful eharacters for identification

(Fig. 26.7). Its posterior margin is defined by a thin rim, the postocciput, which either encloses a pair of cervical sclerites or has a gap that leaves the cervical sclerites free. The pattern of head spots on the frontoclypeal apotome is important for species identi fication. In most black flies, each antenna consists of a proximal, medial, and distal article, the latter

bearing a terminal, cone-shaped sensory structure.

The lateral and ventral areas of the head are com

posed primarily of the postgenae. Each side of the head bears a pair of pigmented eyespots, or oeelli. Ventrally, the head has two valuable taxonomic fea tures, the hypostoma and the postgenal cleft. The hypostoma is an anteriorly toothed plate important for generic identification (Figs. 26.8-26.12). The teeth of the hypostoma consist of a single median tooth bounded on each side by sublateral, lateral, and paralateral teeth. Intermediate teeth, best developed in the prosimuliines, are on either side of one or more of the sublateral teeth. The postgenal cleft is an area of thin, unpigmented cuticle that varies in size from a tiny triangle to a deep, clear area extended anteriorly to the posterior margin of the hypostoma (Fig. 26.6). One of the most conspicuous larval features is a pair of labralfans that filter particulate matter from the current (Fig. 26.1). In mature larvae, these fans eonsist of about 20-80 primary rays. Labral fans are seeondarily absent in Gymnopais and Twinnia. Addi tional mouthparts include mandibles, maxillae, a labrum, and a fused labium and hypopharynx. The brushes on the mandibles remove filtered matter from the labral fans and direct it into the mouth. The man

dibles also help graze food from the substrate, espe cially in fanless larvae. The maxillae, each with a one-segmented maxillary palpus, are of rather uni form structure across the family. Silk is extruded through the common opening ofthe silk glands, which lies between the lower labial and upper hypopharyngeal lobes. The banding patterns of the giant chromo somes in the silk glands are important in taxonomy and identifieation. The labrum appears as an anterior continuation of the frontoclypeal apotome. The body consists of three thoracie segments and nine apparent abdominal segments. Spiracles, though present, are nonfunctional. Pigmentation of the body is highly varied, ranging from white to various shades of green, red, brown, and black. Colors are particu larly apparent in freshly fixed specimens, and the color and distribution of pigment is useful in speeies identi fication. The final instar is recognized by the dark gill histoblast—the future gill of the pupa—on either side of the thorax (Fig. 26.1). The histoblast can be dis sected out and uneurled to provide additional taxo nomic information. The thorax has a single ventral prothoracic proleg of two articles. The distal article, which is sometimes retracted within the basal article in

fixed larvae, bears a pair of lateral sclerites and an apical ring of tiny hooks (Fig. 26.15). The abdomen expands either gradually toward the posterior or rather abruptly at the fifth segment. The larvae of

1100

Chapter 26 Simuliidae

Ectemnia have a nearly prehensile abdomen that expands markedly at segment V (Fig. 26.18). The last abdominal segment of all larvae bears a single poste rior prolog that appears as a continuation of the abdomen and is armed with a ring of minute hooks that anchor the larva to its silk pad. Anterodorsal to the ring of hooks is the anal sclerite (Fig. 26.19). It is usually X-shaped, but in a few taxa it is rectangular, Y-shaped, or absent. In a few taxa, the posteroventral arms ofthe anal sclerite form a continuous ring(Gigantodax) or nearly continuous ring (Parasimulium) around the last segment (Fig. 26.17). Anterior to the anal sclerite are the unpigmented, eversible rectal papillae that function in osmoregulation and are often withdrawn in fixed larvae. They are comprised ofthree simple or three compound lobes with numerous smaller lobules. Paired ventral tubercles arise from the

last segment of many black flies (Fig. 26.19), and a transverse midventral bulge occurs on the final segment of several black flies (e.g., Stegopterna, Fig. 26.16).

Pupae The body ofthe pupa is uniform in shape through out the family and reflects the shape of the adult. The antennal sheaths of female pupae reach or exceed the posterior margin of the head, whereas those of male pupae extend one-half to three-quarters of the distance

to the posterior margin ofthe head.The pupal abdomen has nine visible segments. Anterior to the first abdomi nal segment is a small dorsal postscutellar bridge. Each side of the pupal abdomen has a striate pleural mem brane, which contains large pleurites on the fourth and fifth segments of Prosimulium and Helodon(Fig. 26.20). The paired spiracular gills(respiratory organs)are among the most important taxonomic characters of the pupa(Fig. 26.21). Each gill arises from the antero-

conspicuous features are eight (rarely six) recurved hooks along the posterior margin of each of the third and fourth tergites. The ninth abdominal segment of most species has a pair of terminal spines that can be long and slender (e.g., in most prosimuliines) or short (e.g., in most species of Simulium). The sternites of the third to seventh segments have small recurved hooks. The pleural region of the eighth and ninth segments usually has variously shaped setae, for example anchor- or grapnel-shaped in Metacnephia (Fig. 26.28). Finer setae are scattered over the abdomen. The pupal armature of most species also includes dorsal,transverse rows ofposteriorly directed spine combs on the anterior margins of, at most, the fourth through ninth tergites (Fig. 26.20). All black flies produce a silk cocoon that can be categorized as either shapeless or well-formed. In Parasimulium, the prosimuliines, and some simuliines,the cocoon is a shapeless, sac-like sleeve covering all or part of the pupa (Fig. 26.21). This shapeless cocoon can be as simple as a small, ventral pad of silk that persists after the initial cocoon enclosing the pupa quickly disintegrates {Gymnopais). In three genera {Ectemnia, Metacnephia, and Simulium), the cocoon is well formed with rather rigid walls. Wellformed cocoons are either slipper-shaped if the anterior margin lacks a collar and is flush with the substrate (Fig. 26.24), or boot-shaped if the anterior margin is raised as a collar (Fig. 26.23)(shoe-shaped if the collar is short). Well-formed cocoons can be finely or coarsely woven and sometimes have addi tional features. For example, they can be borne on silk stalks up to 30 mm long {Ectemnia, Fig. 26.22), bear an anterodorsal projection of various lengths (e.g., some Simulium species), or have large open win dows in their sides or loops of silk arising from the anterior margin (Fig. 26.25).

lateral comer ofthe thorax and consists of branches of

varying number, length, and thickness, which are typ ically filamentous but often tubular or club-like. The number of filaments per gill varies from two to more than ICQ. Intraspecific variation in the number of filaments is common, especially in species with many filaments. The surface sculpture of the gill varies from smooth to variously furrowed or tuberculate. The cuticle ofthe head and thorax dorsally can be smooth and shiny or covered with microtubercles ranging from rounded granules to thin spines. The cuticle also can be finely wrinkled or strongly rugose. Sensory hairs, the trichomes, arise from the head and thorax.The thorax typically has 4-7 pairs oftrichomes, butin some species,it has a dense covering oftrichomes. The abdomen has numerous hooks, spines, setae, and combs to secure the pupa in its cocoon. The most

Adults

Adults are small and compact, with short

cigar-shaped antennae, an arched thorax, and broad wings. Most species are blackish but some are reddish brown, gray, or orange. Most males have prominent, reddish compound eyes divided into large upper fac ets and small lower facets. The eyes of females are separated by a distinctfrons and have only small fac ets. Ocelli are absent in adults, but a shiny tubercle, the stemmatic bulla, is located near the posterior mar gin of each compound eye in Parasimulium and some

prosimuliines (Fig. 26.37). The antennae resemble slender, inverted cones or strings of beads. Most spe

cies have nine flagellomeres per antenna, in addition to the scape and pedicel, but a few species have seven

Chapter 26 Simuliidae

or eight flagellomeres. Female mouthparts of about 90% of North American species are designed for blood feeding and have minutely serrated mandibles and toothed maxillary laciniae. The mouthparts of males and of non-bloodsucking females lack serra tions and teeth. The long, slender maxillary palps are composed of five segments. The wings are hyaline or smokey but never patterned, whereas the halteres are white, yellow, or brownish. Wing venation is significant in the identifi cation and classification of black flies (Figs. 26.3026.35). The most conspicuous veins are concentrated along the leading margin of the wing and include the costa(Q,subcosta (Sc), anterior branch of the radius (jR,)> and posterior branch ofthe radius(i.e., the radial sector, Rs). The radial sector can be unbranched or have a long fork (i.e., with widely separated R2 + R^ and + R^ veins); more rarely, the radial sector can have a short, obscure, apical fork. The branched media(M,and Mj), two anterior cubital veins {CuA^ and CUA2), posterior cubitus (CuP), and two anal veins {A^ and A2) are weakly expressed. A false vein (medial-cubital fold) lies between M2 and CuAj. Parasimulium and the prosimuliines have only thin hair-like setae on the costa (Fig. 26.35), whereas most other taxa have short, usually dark spiniform setae among the hair-like setae (Fig. 26.33). The legs of most black flies are uniform in color, whereas those of the more derived Simulium are dis

tinctly patterned or banded. The basitarsus lies distal to the tibia, and on the hind leg it bears an inner apical

1101

flattened flange, the calcipala (Fig. 26.41). Each tar sus consists of four tarsomeres. The upper surface of the first tarsomere of the hind leg in the genus Simulium has a variously incised area, the pedisulcus, which can be long and shallow or short and deep (Fig. 26.42). In some genera of the tribe Simuliini (e.g., Stegopterna and Ectemnid), it is represented by faint wrinkling. The distalmost leg segment consists of a pair of claws. A thumb-like lobe on the female claw (Fig. 26.43)indicates that birds are the principal hosts, whereas a simple claw (Fig. 26.39) or one with a small subbasal tooth indicates that mammals are the

main hosts or that the species does not take blood. The claws of males have large, grooved lobes ofcuticle (grappling hooks) dorsally. The elongate abdomen consists of 11 segments. The terminalia of males and females provide the most definitive means of species identification for adults. The terminalia of the female include the eighth seg ment with its pair of hypogynial valves that form the functional ovipositor, the Y-shaped ninth sternite {genitalfork),the tenth abdominal segment with a pair ofanal lobes(paraprocts),and a pair ofone-segmented cerci(Fig. 26.50). The internal sperm-storage recepta cle, or spermatheca, is sclerotized and typically darkly pigmented (Fig. 26.51). The male terminalia consist of the genitalia, the tenth segment with its small tergite, and the cerci. The genitalia are composed ofthe claspers, or gonopods {gonocoxite + gonostylus), and the aedeagus with its sclerotized plates (e.g., ventral plate) and associated parameres(Fig. 26.47).

KEYS TO THE GENERA OF SIMULIIDAE

(Modified from Adler et al. 2004; figures reprinted from The Black Flies(Simuliidae) ofNorth America, by Peter H. Adler, Douglas C. Currie and D. Monty Wood. Copyright © 2004 by Cornell University. Used by permission of the publisher, Cornell University Press.) Larvae 1. V.

2(1). 2'.

3(1').

Labral fans absent. Head tapered anteriorly (Figs. 26.2 and 26.3) 2 Labral fans present(often closed, but with stalks obvious). Head with sides nearly parallel (Figs. 26.4-26.7) 3 Frontoclypeal apotome with posterolateral head spots(Fig. 26.2). Labrum with prominent bulge on each lateral margin Gymnopals Stone Frontoclypeal apotome without posterolateral head spots(Fig. 26.3). Labrum without bulges on lateral margins Twinnia Stone and Jamnback Anal sclerite with posterior arms nearly or completely encircling base of posterior proleg (Fig. 26.17), or if posterior arms difficult to resolve, then body of larva completely unpigmented. Western mountains

3'.

4(3).

4

Anal sclerite, if present, with posterior arms absent or extended ventrally no farther than about 1/2 distance around base of posterior proleg (Fig. 26.19), or if posterior arms difficult to resolve, then body of larva variously pigmented. Widely distributed 5 Body unpigmented. Head without ocelli or head spots. Pacific Northwest ...Parasimulium Malloch

gill histoblast

antenna

Figure 26.1 iabral fan

prothoracic proleg

posterior proleg

labrum

f

1

' '4/

s ocelli

posterolateral head spots

Figure 26.2

Figure 26.3

Figure 26.4

stalk of Iabral fan

frontoclypeal apotome

4 hypostoma hypostomal groove

postgena /'/f , / '

r

postocciput

cervical sclerite

Figure 26.7

postgenal cleft

Figure 26.5 Figure 26.6

Figure 26.1 Lateral view of larva of Simulium venustum. Figure 26.2 Dorsal view of larval head of Gymnopais holopticus (from Adier ef a/. 2004). Figure 26.3 Dorsal view of larval head of Twinnia tibblesi(from AdIer ef al. 2004). Figure 26.4 Dorsal view of larval head of Prosimulium mixtum (from AdIer etal. 2004). 1102

Figure 26.5 Dorsal (a) and ventral (b) views of larval head of Stegopterna mutata (from AdIer ef al. 2004). Figure 26.6 Dorsal (a) and ventral (b) views of larval head of Metacnephia sommermanae (from Adier ef al. 2004). Figure 26.7 Dorsal view of larval head of Simulium tribulatum (from AdIer ef a/. 2004).

Chapter 26 Simuliidae

1103

lateral tooth

Figure 26.8

Figure 26.9

Figure 26.10

Figure 26.11

median tooth

paralateral

intermediate

teeth

tooth

sublateral teeth

Figure 26.13

Figure 26.12

Figure 26.8 Ventral view of larval hypostoma of Stegopterna mutata. Figure 26.9 Ventral view of larval fiypostoma of TIalocomyia ramifera. Figure 26.10 Ventral view of larval hypostoma of Ectemnia invenusta.

Figure 26.11 Ventral view of larval hypostoma of Cnephia dacotensis. Figure 26.12 Ventral view of larval hypostoma of SimuHum vittatum.

Figure 26.13 Ventral view of anterior margin of larval hypostoma of Prosimulium mixtum (from Adier et al. 2004).

1104

Chapter 26 Simuliidae

4'.

Body pigmented. Head with ocelli and head spots. Southwest Gigantodax Enderlein [1 species: G. adleri Moulton]

5(3').

Antenna with proximal and medial articles transparent, colorless, contrasting with dark brown distal article (Fig. 26.4). Hypostoma with intermediate teeth between primary teeth; paralateral teeth absent(Fig. 26.13)

5'.

Antenna with proximal, medial, or both articles lightly to darkly pigmented (Figs. 26.5 and 26.7), or if colorless, then postgenal cleft extended anteriorly to, or beyond, hypostomal groove(Fig. 26.6). Hypostoma without intermediate teeth; paralateral teeth present(Fig. 26.12) Prothoracic proleg with lateral sclerite a narrow bar parallel to base of hooks, extended at most 1/3 distance to base of apical article (Fig. 26.14)(requires dissection if apical

6(5).

article is withdrawn)

6

7

Helodon Enderlein

6'.

Prothoracic proleg with lateral sclerite extended 1/2 or more distance to base of apical article (Fig. 26.15)(requires dissection if apical article is withdrawn) Prosimulium Roubaud

7(5').

Hypostoma with lateral and sublateral teeth not clustered on prominent,common lobes (Figs. 26.10-26.12)(median and lateral teeth can be extended beyond sublateral teeth, but not on prominent lobes); in doubtful specimens (i.e., subgenus Hellichiella Rivosecchi and Cardinali, in part), heavily sclerotized anterior portion of hypostoma occupying at most 1/6 total length of hypostoma Hypostoma with 1 or more sublateral teeth and 1 or more paralateral teeth clustered on prominent, common lobes, giving hypostoma trilobed appearance (consisting of median tooth and 2 lateral lobes); heavily sclerotized anterior portion of hypostoma occupying 1/3 or more total length of hypostoma (Figs. 26.8 and 26.9)

7'.

8

11

8(7).

Hypostoma with apex of median tooth extended anteriorly to, or beyond, apices of lateral teeth; sublateral teeth variously but distinctly posterior to median and lateral teeth (Fig. 26.12) 5i»ih//h/m Latreille

8'.

Hypostoma with apex of median tooth posterior to apices of lateral teeth, or all teeth uniformly small; sublateral teeth with apices extended to various levels (Figs. 26.10 and 26.11) 9 Postgenal cleft extended anteriorly to, or slightly beyond, hypostomal groove (Fig. 26.6). Antenna with transparent, colorless proximal and medial articles contrasting with dark brown distal article Metacnephia Crosskey Postgenal cleft typically extended anteriorly 1/2 or less distance to hypostomal groove, broadly rounded or pointed anteriorly. Antenna with proximal and medial articles variously

9(8').

9'.

pigmented, but not entirely transparent

10

10(9').

Abdomen without abrupt lateral and ventral expansion at segment V,and without pair of ventral tubercles on segment IX. Hypostoma with anterior margin in form of 3 short, convex lobes(Fig. 26.11) Cnephia Enderlein

10'.

Abdomen with abrupt lateral and ventral expansion at segment V, and with pair of ventral tubercles on segment IX (Fig. 26.18). Hypostoma with anterior margin distinctly concave (Fig. 26.10) Ectemnia Enderlein Abdominal segment IX with pair of prominent ventral tubercles (as in Fig. 26.19) Greniera Doby and David Abdominal segment IX without pair of ventral tubercles, but with at most 1 transverse, midventral bulge (Fig. 26.16) 12 Antenna longer than stalk of labral fan by about 1/2 length of distal article (Fig. 26.5). Hypostoma with outer margin of lateral cluster of teeth typically sloped inwardly (Fig. 26.8) Stegopterna Enderlein

11(7'). 11'.

12(11').

Chapter 26 Simuliidae

1105

lateral sclerlte

Figure 26.15

Figure 26.14

anal sclerlte

Figure 26.16 midventral

Figure 26.17

bulge

sclente

rs

Figure 26.18

ventral

ventra

tubercle

Figure 26.14 Lateral view of larval prothoracic proleg of Helodon onychodactylus (after Peterson 1970). Figure 26.15 Lateral view of larval prothoracic proleg of Prosimulium mixtum (after Peterson 1970). Figure 26.16 Lateral view of larval abdomen of Stegoptema mutata.

Figure 26.19

tuberc e

Figure 26.17 Lateral view of posterior portion of larval abdomen of Gigantodax adieri. Figure 26.18 Lateral view of larval abdomen of Ectemnia invenusta.

Figure 26.19 Lateral view of posterior portion of larval abdomen of Simulium bracteatum.

1106

Chapter 26 Simuliidae

12'.

Antenna equal in length to, or marginally extended beyond, stalk of labral fan. Hypostoma with outer margin of lateral cluster of teeth either approximately straight, or sloped somewhat outwardly (Fig. 26.9) Tlalocomyia Wygodzinsky and Diaz Najera

Pupae

1.

Cocoon rudimentary or shapeless and sac-like (Fig. 26.21)

1'.

Cocoon shaped like slipper, shoe, or boot, with definitely formed, rigid walls (sometimes coarsely woven)(Figs. 26.22-26.25) 10 Gill of 3 filaments arising from elongate base. Pacific Northwest Parasimulium Malloch

2(1). 2'. 3(2'). 3'.

2

Gill of 2 or more filaments arising from short base. Widespread 3 Gill of 2 to 4(rarely 5)filaments. Cocoon a small ventral pad (typically not collected with pupa) Gymnopais Stone Gill of more than 4 filaments. Cocoon covering at least part of pupal dorsum 4

4(3').

Gill of 5 inflated tubes each bearing 0-10 tiny, secondary filaments (Fig. 26.26). Southwest Gigantodax Enderlein [1 species: G. adleri Moulton]

4'.

Gill of6 or more slender filaments or inflated tubes without tiny, secondary filaments. Widespread

5

5(4').

Abdominal segments IV and V each with large pleurite in lateral striate membrane (Fig. 26.20)

5'.

Abdominal segments IV and V without large pleurites, or with at most, minute pleurites in lateral striate membrane

6(5).

7

Abdominal tergites VI-IX each with transverse row of spine combs along anterior margin (Fig. 26.20); tergites III and IV each with 4 pairs of recurved hooks

6'.

6

Prosimulium Roubaud, Helodon Enderlein

Abdominal tergites without spine combs; tergites III and IV each with 3 pairs of recurved hooks

Twinnia Stone and Jamnback

7(5').

Abdominal segments VIII and IX laterally with hook-shaped setae (Fig. 26.27). Gill of 17-50 filaments Cnephia Enderlein

7'.

Abdominal segments VIII and IX laterally either with straight or slightly curved setae, or

8(7'). 8'.

Gill of 10 or 12 filaments arising from base on 2 or 3 slender trunks Stegoptema Enderlein Gill offewer than 10 filaments or of 15-30 filaments; if gill of 12 filaments, then base rather inflated and filaments arising in 4 or 5 main groups 9 Gill of 6-12 moderately inflated, rigid filaments, typically radiated laterally from base Tlalocomyia Wygodzinsky and Diaz Najera Gill of 15-30 slender, delicate filaments, typically projected forward Greniera Doby and David Gill of 8 or 10 stout filaments arising close to base and converging anteriorly toward common point. Cocoon attached to silk stalk (Fig. 26.22) Ectemnia Enderlein

without setae (Fig. 26.29). Gill of 6-30 filaments

9(8'). 9'. 10(1'). 10'.

8

Gill with various numbers of filaments, but if 8 or 10, then filaments not inflated nor

converging anteriorly toward common point. Cocoon not attached to silk stalk (Figs. 26.23-26.25)

II

11(IO'). Pleural region of abdominal segments VIII and IX with numerous anchor- or grapnel-shaped setae (Fig. 26.28). Cocoon typically boot-shaped, loosely or tightly woven, but without definitely formed anterior apertures or loops(Fig. 26.23). Canada, Alaska, and western mountains Metacnephia Crosskey

Chapter 26 Simuliidae

1107

recurved

hooks spine 'comb

pleurltes

Figure 26.20 trichome

Figure 26.21

Figure

r\

ill

nii^' ^ r '/ C^/

/'"Ci -r 1*

Figure 26.23

Figure 26.20 Lateral view of pupa of Prosimulium clandestinum; cocoon removed (from Adier ef al. 2004). Figure 26.21 Lateral view of pupa of Prosimulium ursinum.

Figure 26.22 Lateral view of pupa of Ectemnia invenusta (from AdIer ef al. 2004). Figure 26.23 Lateral view of pupa of Metacnephia villosa (from AdIer ef al. 2004).

,// V ,

Figure 26.24

f-i '/

^ \

''

^ - /

Figure 26.25

hook-shaped seta

Figure 26.26

Figure 26.27 grapnel-shaped

termina

seta

m Figure 26.29 Figure 26.28

Figure 26.24 Lateral view of pupa of Simulium vittatum. Figure 26.25 Lateral view of pupa of Simulium arcticum (from Adier et al. 2004). Figure 26.26 Frontal view of pupal gills of Gigantodax adieri.

Figure 26.27 Lateral view of posterior portion of pupai abdomen of Cnephia dacotensis (from Adier etal. 2004). 1108

Figure 26.28 Lateral view of posterior portion of pupal abdomen of Metacnephia borealis (from Adier et al. 2004). Figure 26.29 Lateral view of posterior portion of pupal abdomen of Stegopterna mutata (from Adier et al. 2004).

Chapter 26 Simuliidae

1 r.

1109

Pleural region of abdominal segments VIII and IX at most with unbranched setae. Cocoon variously shaped (Fig. 26.24), but if boot-shaped, then with definitely formed apertures, loops, or perforations anteriorly (Fig. 26.25)except in 2 southwestern species. Widespread Simulium Latreille

Adults

I.

Wing vein R]joined to costa near middle of wing (Fig. 26.30). False vein (m-cu fold) unforked apically. Pacific Northwest(PARASIMULIINAE) Pamsimulium Malloch

r.

Wing vein Rjjoined to costa beyond middle of wing (Figs. 26.31-26.33). False vein (m-cu fold) forked apically. Widespread(SIMULIINAE)

2

2(1').

Radial sector with fork longer than its stem (Fig. 26.31). Costa with hair-like setae only (Fig. 26.35)(PROSIMULIINI)

3

2'.

Radial sector unforked (Fig. 26.32), or with fork shorter than its stem (Fig. 26.33). Costa with spiniform setae interspersed among hair-like setae (Fig. 26.33)(spiniform setae thin and pale in certain species of Greniera)(SIMULIINI)

6

Antenna with 7 flagellomeres (Figs. 26.36 and 26.37). Eye with shiny, dark, raised tubercle (stemmatic bulla) behind posterior margin (Fig. 26.37). Claws of female toothless

4

Antenna with 8 or more flagellomeres, or if with 7 flagellomeres, then posterior margin of eye without stemmatic bulla. Claws offemale toothless (Fig. 26.38), or each with variously sized basal or subbasal tooth or thumb-like lobe (Figs. 26.43)

5

3(2). 3'.

4(3).

Vestiture of head and body consisting of short, sparse, erect hairs. Clypeus bare except for few erect hairs laterally (Fig. 26.36) Gymnopais Stone

4'.

Vestiture of head and body consisting of long, dense, recumbent hairs. Clypeus covered with hair (Fig. 26.37) Twinnia Stone and Jamnback

5(3').

Male: Ventral plate in lateral and ventral views flattened, with lip absent, short, or slender (Fig. 26.47). Female: Claws each with basal or subbasal tooth or thumb-like lobe (Fig. 26.43) Helodon Enderlein

5'.

Male: Ventral plate in lateral and ventral views not markedly flattened, typically with prominent lip (Fig. 26.46). Female: Claws toothless (Fig. 26.38), or each with minute subbasal tooth

Prosimulium Roubaud

6(2').

Wing vein CuA2 straight. Mountains of Arizona and New Mexico Gigantodax Enderlein [1 species: G. adleri Moulton]

6'.

Wing vein CuA2 sinuous (Figs. 26.32-26.34). Widespread

7(6').

Wing without basal medial cell (Fig. 26.34); radius with or without hair dorsobasally. Tarsomere I of hind leg with pedisulcus deep (Fig. 26.42) Simulium Latreille (in part)

7'.

Wing with basal medial cell, although sometimes small (Figs. 26.32 and 26.33); radius with hair dorsobasally. Tarsomere I of hind leg with pedisulcus absent or represented by shallow wrinkles (Figs. 26.39-26.41)

8(7'). 8'.

9'.

8

Costa with only pale setae, some of which may be short and stiff but neither dark nor fully spiniform Greniera Doby and David (in part) Costa with short, stout, black, spiniform setae interspersed among longer, paler, hair-like setae (the former more prevalent near apex of costa)(Fig. 26.33)

9(8').

7

9

Radial sector bifurcated at apex, with branches separated by membrane(Fig. 26.33). Rocky Mountains westward Tlalocomyia Wygodzinsky and Diaz Najera Radial sector unbranched (Fig. 26.34), or if bifurcated, then branches closely approximated and not separated by membrane. Widespread

10

10(9').

Male

11

10'.

Female

16

1110

Chapter 26 Simuliidae

radial

sector(Rs) |- I

. .

■■

r

X.

^ iV

■ ■,

/

false vein

X,.

Figure 26.31

Figure 26.30

:r.7i,

basal medial cell

'%-f ''"°^VU-.,

Figure 26.32

costa(C)

subcosta (So) radius(R)

Figure 26.33

Figure 26.34 radial sector

Figure 26.35

basal section of radius

Figure 26.30 Figure 26.31 Figure 26.32 Figure 26.33

Wing of Parasimulium stonei male. Wing of Gymnopais holopticus female. Wing of Greniera humeralis female. Wing of TIalocomyia ramifera female.

Figure 26.34 Wing of Simulium venustum female. Figure 26.35 Anterior portion of wing of Prosimulium ursinum female.

Chapter 26 Simuliidae

1111

frons stemmatic antenna

flagellomere

clypeus

i maxillary palp

Figure 26.37

Figure 26.36 tarsomere i

Figure 26.38

Figure 26.39

. basltarsus

claw

calcipaia

Figure 26.41

Figure 26.40 pedisulcus

Figure 26.42

Figure 26.45

thumb-like lobe

Figure 26.43

Figure 26.44

Figure 26.36 Anterior view of head of Gymnopais holopticus female (after Wood 1978). Figure 26.37 Lateral view of head of Twinnia tibblesi

Figure 26.41 Distal portion of hind leg of Simulium anatinum female (from Adier et al. 2004). Figure 26.42 Distal portion of hind leg of Simulium

female.

meridionale female.

Figure 26.38 Distal portion of hind leg of Prosimullum

Figure 26.43 Claw and apex of tarsus of Helodon

ursinum female.

decemarticulatus female.

Figure 26.39 Distal portion of hind leg of Stegopterna

Figure 26.44 Hind tibiai spur of Stegopterna acra female (from Adier et al. 2004). Figure 26.45 Hind tibiai spur of Ectemnia taeniatifrons female (from Adier et al. 2004).

mutata female.

Figure 26.40 Distal portion of hind leg of Metacnephia saskatchewana female.

1112

Chapter 26 Simuliidae

iip of ventral plate

gonocoxite

paramere

gonostylus

Figure 26.46

ventral

plate

Figure 26.47

'/

spinule

Figure 26.48

Figure 26.49

Figure 26.46 Genitalia of Prosimulium unicum male: (a) ventral view;(b) lateral view. Figure 26.47 Genitalia of Helodon alpestris male:(a) ventral view;(b) lateral view.

Figure 26.48 Ventral view of genitalia of Ectemnia taeniatifrons male.

Figure 26.49 Ventral view of genitalia of Metacnephia sommermanae male.

11(10). 1 r.

Chapter 26 Simuliidae

1113

Ri dorsally with hair-like setae and scattered, black, spiniform setae on distal 2/3 or more; spiniform setae near apex more numerous than, and as stout as, those on costa

12

Ri dorsally with hair-like setae only, or if spiniform setae present, these confined to apical 1/2 or less, and not as stout as those on costa

14

12(11). Gonostylus with 2 or 3 apical spinules (Fig. 26.48) Ectemnia Enderlein 12'. Gonostylus with 1 apical spinule (Fig. 26.49) 13 13(12'). Basitarsus of hind leg with calcipala absent or minute and apically pointed (as in Fig. 26.40). Tarsomere I of hind leg without pedisulcus dorsobasally, although minute wrinkles can be present(as in Fig. 26.40). Anepisternal membrane typically with pale hair dorsally (absent in M. saskatchewana) Metacnephia Crosskey 13'. Basitarsus of hind leg with small but distinct, apically rounded calcipala (as in Fig. 26.41). Tarsomere I of hind leg with pedisulcus represented by shallow, wrinkled depression (as in Fig. 26.41). Anepisternal membrane bare Simulium Latreille (in part)[subgenus Hellichiella Rivosecchi and Cardinali] 14(11'). Hind tibial spurs with pale apices and longer than width of tibia at point of attachment (as in Fig. 26.44) Stegopterna Enderlein 14'. Hind tibial spurs uniformly dark, equal in length to, or shorter than, width of tibia at point of attachment(as in Fig. 26.45) 15 15(14'). Alaska, Yukon, and east of Rocky Mountains Cnephia Enderlein 15'.

Mountains from southern British Columbia to

California

Greniem Doby and David (in part)[G. humeralis Currie, Adler, and Wood]

16(10'). 16'. 17(16).

Ri dorsally with hair-like setae and scattered, black, spiniform setae on distal 1/2. Wing hyaline 17 R] dorsally with hair-like setae only, or if spiniform setae present, then wing rather smokey 18 Basitarsus of hind leg with calcipala absent or minute and apically pointed (Fig. 26.40). Tarsomere I of hind leg without pedisulcus dorsobasally (Fig. 26.40). Anepisternal membrane typically with pale hair dorsally (absent in M. saskatchewana) Metacnephia Crosskey 17'. Basitarsus of hind leg with small but distinct, apically rounded calcipala (Fig. 26.41). Tarsomere I of hind leg with pedisulcus represented by shallow, wrinkled depression (Fig. 26.41). Anepisternal membrane bare Simulium Latreille (in part)[subgenus Hellichiella Rivosecchi and Cardinali] 18(16'). Claws toothless (Fig. 26.39). Hind tibial spurs with pale apices, and longer than width of tibia at point of attachment(Fig. 26.44) Stegopterna Enderlein 18'.

Claws each with small subbasal tooth or large, basal, thumb-like lobe (as in Fig. 26.43). Hind tibial spurs uniformly dark, equal in length to, or shorter than, width of tibia at point of attachment (Fig. 26.45)

19

19(18').

Spermatheca elongate, with large unpigmented area at junction with spermathecal duct (Fig. 26.52) Ectemnia Enderlein

19'.

Spermatheca spherical or reniform, with small or no unpigmented area at junction with spermathecal duct(Figs. 26.50 and 26.51)

20

20(19').

Spermatheca large, spherical, wrinkled (Fig. 26.51). Alaska, Yukon, and east of Rocky Mountains Cnephia Enderlein

20'.

Spermatheca small, reniform, smooth (Fig. 26.50). Mountains from southern British Columbia to

California

Greniera Doby and David (in part)[G. humeralis Currie, Adler, and Wood]

1114

Chapter 26 Simuliidae

spermatheca

genital fork

spermatheca!

Figure 26.51

duct

v/iimu

8

hypogynial va ve

Figure 26.52

Figure 26.50

Figure 26.50 Ventral view of genitaiia of Greniera

Figure 26.52 Spermatheca of Ectemnia invenusXa

humeratis female.

female.

Figure 26.51 female.

Spermatheca of Cnephia dacotensis

Chapter 26 Simuliidae

ADDITIONAL TAXONOMIC REFERENCES

Most of the following references are taxonomically outdated but are included for the sake of completeness. Readers should use these references in consultation with Adler et al. (2004) to ensure the use of currently accepted species concepts and nomenclature.

General Coquillett (1898); Malloch (1914); Johannsen (1903, 1934); Dyar and Shannon (1927); Vargas (1945); Nicholson and Mickel (1950); Stone and Jamnback (1955); Wirth and Stone (1956); Carlsson (1962); Davies et al.(1962); Wood et al. 0963); Stone (1964a, 1965); Stone and Snoddy (1969); Crosskey (1973, 1988, 1990); Peterson (1981); Adler and Kim (1986); Currie and Walker (1992); Peterson and Kondratieff (1995); Crosskey and Howard (1997); Adler et al.(2004); Adler and Crosskey (2018).

Regional faunas Alabama: Stone and Snoddy (1969). Alberta: Fredeen (1958); Abdelnur (1968); Currie (1986). Alaska: Stone (1952); Sommerman (1953); Peterson (1970); Hershey et al.(1995a); Currie (1997). British Columbia: Hearle (1932); Currie and Adler (1986). California: Wirth and Stone (1956); Hall(1972,1974); Tietze and Mulla(1989). Canada: Shewed (1958); Peterson (1970); Currie (2014). Colorado: Ward and Kondratieff(1992); Peterson and Kondratieff (1995). Connecticut: Stone (1964a). Delaware: Sutherland and Darsie (1960). Eastern Canada: Twinn (1936). Florida: Pinkovsky and Butler (1978). Idaho: Twinn (1938). Kansas: Snyder and Huggins (1980); Mock and Adler (2002). Maine: Bauer and Granett(1979). Manitoba: Fredeen (1958); Crosskey (1994). Maritime Provinces: Lewis and Bennett(1979).

1115

Michigan: Gill and West (1955); Merritt et al.(1978). Minnesota: Nicholson and Mickel (1950). Missouri: Doisy et al.(1986). Montana: Newell (1970). Nebraska; Pruess and Peterson (1987). Newfoundland: Lewis and Bennett (1973). New Jersey: Crans and McCuiston (1970); Carle (2011). New York: Stone and Jamnback (1955). North America: Adler et al.(2004). Northeastern United States: Cupp and Gordon (1983). Ontario: Davies et al. (1962); Wood et al. (1963). Pennsylvania: Frost(1949); Eckhart and Snetsinger (1969); Adler and Kim (1986). Prince Edward Island: Minhaus et al.(2005). Quebec: Back and Harper (1978); Gadreau and Charpentier (2011). Rhode Island: Dimond and Hart(1953). Saskatchewan: Fredeen (1958,1981, 1985). South Carolina: Arnold (1974); Noblet et al.(1979). Southeastern United States: Snoddy and Noblet(1976). Utah: Twinn (1938); Peterson (1955, 1960). Washington: Bacon and McCauley (1959). Western United States: Stains and Knowlton (1943). Wisconsin: Anderson (1960). Yukon: Currie (1997).

Taxonomic treatments at the generic level (L = larvae; P = pupae; A = adults) All genera (all species): Adler et al.(2004)-L, P, A. GymnopaLs: Wood (1978)-L, P, A. Ectemnia: Moulton and Adler (1997)-L, P, A. Gigantodax-. Moulton (1996)-L, P, A. Parasimulium: Peterson (\9n)-A. Prosimulium: Peterson (1970, 1989)-L, P, A. Simulium: Nicholson and Mickel(1950)-A; Stone and Jamnback (1955)-L,P, A;Peterson (1960,1993)-L, P, A; Davies et al.(1962)-P, A; Wood et al.(1963)-L; Stone(1964a)-L,P, A;Stone and Snoddy(1969)-L, P, A; Currie (1986)-L, P; Peterson and Kondratieff(1995)-L, P, A; Moulton and Adler (1995)-L, P, A; Moulton (1998)-L, P, A.(Also see references under headings General, and Regionalfaunas, above.) Twinnia: Wood (1978)-L, P, A.

Family

Genus

Simuliidae

(256)

collectors—

gatherers

end of abdomen

Clingers

Clingers Clingers

Lotic—erosional

Lotic—erosional Lotic—erosional Lotic—erosional

Ectemnia (4)

Gigantodax

Greniera (7)

Gymnopais(5)

Scrapers

Collectors—^filterers

Collectors—filterers

Collectors—^filterers

Midwest, NW = Northwest, MA = Mid-Atlantic

Clingers

Clingers

threads)

In silk

anchored

and proleg

Collectors—filterers

facultative

terminal

minute

predators (engulfers) and

some scrapers,

collectors—filterers;

Generally obligate

Trophic Relationships

hooks on

Generally clingers (ring of

Habit

Lotic—erosional

Generally lotic— erosional

Habitat

Cnephia (4)

adierl

Species

SE = Southeast, UM = Upper Midwest, M : * Emphasis on trophic relationships

Gnats

Diptera - Black Flies and Buffalo

Order

parentheses)

(number of species in

Taxa

R. W. Merritt, K. W. Cummins, and B. V. Peterson.)

North

6.0

NW

MA*

Ecological

152, 968, 2094, 3332,

969, 1216, 1279, 2937, 2975, 4669, 4675, 5129, 5354, 6266, 5005, 6743, 5133, 6740, 6745, 3139, 21, 26, 27, 1736, 1813, 3704, 4018, 4100, 4136, 6751, 6847, 6848, 4058, 4070, 6371

References**

Alaska

Canada and

Widespread

U.S.

Southwestern

Mountains

east of Rocky

Canada and U.S.

Rocky

1355, 5428, 6677, 6331, 6705, 26

1349, 26

4163, 26

5428, 26

6735, 26

5.0

M

Mountains

UM

3333, 4019, 5129, 5132, 3698, 5882,

4.0

SE

U.S. east of

Alaska, Canada,

Distribution

American

Tolerance Values

Table 26A Summary of ecological and distributional data for Simuliidae (Diptera).(For definition of terms, see Tables 6A-6C; table prepared by P. H. Adier, D. C. Currie,

) ) ) )))) ) ) ) ) ) ) ) ) ) ) ) ) )))) ) ^ )

^

Family

Lotic—erosional Clingers (hyporheic zone)

Lotic—erosional

Parasimulium (5)

Prosimulium (38)

Clingers

Clingers

Lotic—erosional

Metacnephia (7)

Habit

C lingers

Habitat Lotic—erosional

Species

Helodon (16)

Genus

Continued

North

Western U.S.

Collectors— filterers; facultative scrapers and predators

SIMULIIDAE

Widespread

Northwest

(engulfers)

Pacific

filterers; facultative predators

Alaska

and Canada;

Collectors—

Collectors—^filterers

(engulfers)

Western North

America

Collectors—

Distribution

American

filterers; facultative predators

Trophic Relationships

SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ■* Emphasis on trophic relationships

Order

parentheses)

(number of species in

Taxa

Table 26A

2.9

SE

3.0

UM

M

Tolerance values

3.0

6.0

NW

MA*

Ecological

(continued)

152, 905, 968, 1356, 2094, 2152, 2213, 2433, 2569, 3332, 3333, 3745, 4019, 4020, 4667, 5129, 5133, 5354, 6328, 6331,29, 26, 1677

1156, 1162,4668, 5761, 26

905, 5428, 6738, 6739, 26, 3703, 6752

2569, 4667, 26, 89, 87

References**

1

00

)

Family

Ciingers

Ciingers Ciingers

Lotic—erosiona!

Lotic—erosional Lotic—erosional

TIalocomyia (4)

Twinnia (3)

Ciingers

Habit

Stegopterna (8)

Habitat Lotic-eroslonal

Species

Simulium (154)

Genus

Continued

Scrapers

)

)

)

)

I

)

)

)

)

)

)

)

)

Collectors—filterers

scrapers

filterers; facultative

Collectors—

Collectors—^fiiterers

Trophic Relationships

^ SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic ■* Emphasis on trophic relationships

Order

species in parentheses)

(number of

Taxa

Table 26A

North

)

)

and Canada

northeast U.S.

Western and

America

Western North

Widespread

Widespread

Distribution

American

)

SE

)

UM

)

M

Tolerance Values

)

5.0

NW

)

)

MA*

Ecological

)

6705, 26

)

968, 1355, 2213,

26

4019, 4020, 5129, 5132, 5428, 26

3010

)

3933,4030, 4019, 4119,4183,4240, 4318, 4320, 4665, 4669, 4841, 4919, 5129, 5133, 5353, 5411, 5428, 5882, 6210, 6289, 6328, 6331, 6611, 6734, 6739, 6741, 6744, 6747, 6771, 6832, 23, 29, 26, 1037, 1736, 1902, 3138, 6844, 6849, 1960, 6610, 6737,4442, 1677,

152, 261, 390, 651, 653, 800, 905, 968, 979, 1077, 1082, 1083, 1179, 1534, 1701, 1959, 1961, 2020, 2132, 2135, 2136, 2365, 2520, 2568, 3119, 3271, 3332, 3333, 3340, 3343, 3348, 3346, 3680, 3698, 3748,

References**

I

m

t

««< ■*«»

CfflRONOMIDAE Leonard C. Ferrington, Jr. University ofMinnesota, St. Paul

INTRODUCTION

The family Chironomidae (nonbiting midges) is worldwide in distribution and is an ecologically important group of aquatic insects often occurring in high densities and diversity. The relatively short life cycles and the large total biomass of larvae confer ecological energetic significance on this taxon (as both consumers and prey) and the partitioning of ecological resources by a large number of species pre sumably enhances the biotic stability of aquatic ecosystems.

Although slightly more than 1,200 species are known from the Nearctic Region, it is estimated that as many as 2,000 species may occur, including numer ous undescribed species. More than 5,000 species are described worldwide, however some may not be aquatic, so the actual number of aquatic species cannot be accurately assessed (Ferrington 2008) but estimates of actual species range up to 20,000. Chironomidae is among the most specious families of aquatic Diptera. Presumably, the great species diversity in this family is the product of its antiquity, relatively low vagility (instances of geographic isolation appear to be com mon), and evolutionary plasticity. The overall diver sity of the family also is reflected in the rich chironomid faunas of many aquatic ecosystems. The Chironomidae is unequivocally the most widespread of all aquatic insect families. Chironomids occur in most types of aquatic ecosystems, as well as moist soils, tree holes, pitcher plants, and dung. The range of conditions under which chironomids are found is more extensive than that of any other family

of aquatic insects. Almost the complete range of gra dients of temperature, pH, salinity, oxygen concen tration, current velocity, depth, productivity, altitude, latitude, and other parameters have been exploited, at least by some chironomid species. Species occur from Antarctica at 68°S latitude (Belgica antarctica Jacobs) and subantarctic islands (Parochlus steinenii (Gerke)) (Edwards and Usher 1985; Sugg et al. 1983) to Lake

Martin B. Berg Loyola University Chicago, Illinois

Hazen at 81 °N on Ellesmere Island (Oliver and Corbet 1966). They exhibit extreme elevational ranges, occur ring in a glacial-melt stream at 5,600 m in the Himalaya Mountains (Koshima 1984) to more than 60 m depths in Lake Hovsgol (Hayford and Ferrington 2006) and >1,000 m depths in Lake Baikal (Linevich 1971). They are among the most tolerant of aquatic insects to water and air temperatures, with larvae of Paratendipes thermophilus Townes maturing in hot springs at temperatures of 38.8°C (Hayford et al. 1995) and larvae of Diamesa mendotae Muttkowski surviving freezing at -15°C (Bouchard et al. 2006b). Adults of D. mendotae also are able to depress their freezing point after emergence and survive in air tem peratures less than -20°C (Carrillo et al 2004; Bouchard et al 2006a). This species is also excep tional in that it can survive for extended periods below freezing as an adult (Anderson et al 2013; Mazack etal 2014). The wide ecological amplitude displayed by species of Chironomidae is related to the very exten sive array of morphological, physiological, and behavioral adaptations found among the members of the family. Ecologists have used the partitioning of gradients by various midge species to characterize the overall ecological conditions of lentic and lotic sys tems, including estuarine habitats (Kranzfelder and Ferrington 2016, 2018) and rock pools (Egan and Ferrington 2015). The actual number of species present in a system is the result of the complex of physical, chemical, bio logical, and biogeographic conditions. When sampled comprehensively, species richness of Chironomidae is usually among the highest of aquatic insect families detected in most aquatic settings, often approaching 80 or more species and occasionally exceeding 100 species per site. Coffman (1989) summarized 152 species richness estimates as a function of stream order and concluded that average richness increased with increasing stream order up to third order, then

1119

1120

Chapter 27 Chironomidae

leveled off or decreased in higher order rivers. Mean species richness(and range of estimates) varied from; 26 (10-64) for first order streams; 44 (13-144) for second order; 63(25-157)for third order; 51 (25-83) for fourth order; 47(11-86)for fifth order; 47(10-99) for sixth order; 45 (12-148) for seventh order and higher. By contrast, the least diverse faunas are strongly correlated with extreme conditions when richness may be as low as a dozen or fewer species. Chironomid larvae are known to feed on a great variety of organic substrates:(1)coarse detrital parti cles (leaf- and wood-shredders);(2) medium detrital particles deposited in or on sediments (gatherers and scrapers); (3) fine detrital particles in suspension, transport, or deposited (filter-feeders, gatherers, and scrapers);(4) algae; benthic, planktonic, or in trans port (scrapers, gatherers, and to a lesser extent filter-feeders);(5) vascular plants (miners);(6)fungal spores and hyphae (gatherers);(7) animals (as simple predators, often preying on other chironomid larvae, or as parasites on a variety oftaxa, although the latter may most often be commensal relationships) (Berg 1995). Symbiotic bacteria and fungi that occur in the mid- or hindguts of larvae (Slaymaker et al. 1998) likely contribute to the ability of species to process and assimilate nutrients associated with lignins and/ or celluloses in detrital particles. Most aquatic predators feed extensively on chironomids(larvae, pupae, and/or adults)at some point in their life cycles. As some predaceous fish species increase in size, they may rely less on chironomids (French et al. 2014)or exhibit size-selective predation during winter (Anderson et al 2016). As holometabolous insects, chironomids have four distinct life stages: egg, larva, pupa, and adult (imago). Duration of the larval stage, with four instars, may last from less than 2 weeks to several years, depending on species and environmental conditions.

In general, warm water, high-quality food, and small size of species correlate with shorter life cycles. Although most species appear to be univoltine to trivoltine in seasonal environments (Berg and Hellenthal 1992; Tokeshi 1995), life cycle strategies of some species can be extreme, with Apedilum elachistus Townes maturing from egg to adult in less than 7 days in rock pool habitats in the Brazilian Pantanal(Nolte 1996). In contrast, Butler(1982)proposed a 7-year life cycle for two species of Chironomus in northern tun dra ponds of Alaska. Cold-adapted species may have a more labile life cycle, with two, three or four gener

ations occurring over successive years (Ferrington and Masteller 2015) First larval instars may be planktonic, are often difficult to sample, and may have a unique

morphology (Ward and Cummins 1978) not treated in keys that are designed for fourth instar larvae. Later instars are usually benthic. Toward the end of the fourth instar, the larval thoracic region begins to swell with the formation of the pupal integument and adult tissues. Differing degrees of swelling and visibil ity of developing pupal structures have been used to age fourth instar larvae (e.g., Bouchard et al 2006b; Butler 1982; Wiilker and Gotz 1968). Upon completion offeeding activities, late fourth instar larvae attach themselves with silken secretions

to the surrounding substrates and pupation occurs. The pupal stage begins with apolysis, the separation (not shedding)ofthe larval from the underlying pupal integument. After ecdysis of the larval skin (exuviae) and subsequent maturation of the developing adult tissues, the pupa frees itself from the silken chamber and swims to the surface of the water where eclosion

(adult emergence) takes place from within the pupal skin (or exuviae). The exuviae fills with air and by virtue of an outer waxy layer of the cuticle(which has nonwettable properties) it remains floating on the water surface until bacteria begin to decompose the wax layer (see surface-floating pupal exuviae below) (Kavanaugh etal 2014). Technically,the adult stage begins with the pupaladult apolysis (pharate adult), which occurs a short time before eclosion. Chironomid adults usually live a few days, although some species may survive for several weeks at low air temperatures (Ferrington et al 2011). The adult stage performs the functions of reproduction and dispersal. As a rule, chironomid adults do not need to feed, as reflected by the usual condition of reduced mouthparts and atrophied gut. However, many species (perhaps most) will take liq uid and semiliquid carbohydrate sources such as aphid honeydew and flower nectars. The consump tion of these energy rich substances presumably max imizes the potential for the completion of additional ovarian cycles. Mating takes place in aerial swarms, on the water surface in skating swarms (Ferrington and Ssther 1987), or on solid substrates. Females may broadcast the eggs at the water surface or, more frequently, deposit gelatinous egg masses on the open water or on emergent vegetation. Egg or larval development may be arrested under unfavorable environmental condi

tions. Some species are facultatively or obligatorily parthenogenetic,and larvae parasitized by nematodes or nematomorphs may produce intersexual adults. Other ectoparasites, endoparasites, and/or symbionts include water mites(Smith and Oliver 1976)and fungi of the Coelomomycetaceae (Weiser and McCauley 1971) and Trichomycetes (Slaymaker et al 1998;

Chapter 27 Chironomidae

1121

Ferrington et al. 2000). Excellent reviews of chironomid biology may be found in Oliver (1971), Davies (1976), Finder (1986), and Armitage et al.(1995). In most ecological studies, the larva is the life stage most frequently encountered. However, the quantitative collection of early instars is often diffi cult(Chapter 3). Even when larval populations can be "adequately" sampled (Chapter 3), sorting and particularly retrieval of early instars can require prodigious amounts of time (densities of 50,000 lar

have been developed for use in wadeable streams (Bouchard and Ferrington 2011). SFPE collection protocols take advantage of the fact that pupal exuviae are not wettable for a period of time and float on the surface where they can be

vae per m^ are not uncommon). After sorting, iden

face or the water column, the substrates are not

tification of larvae also proves to be difficult because: (1) only a small percentage of larvae of Nearctic species have been described; (2) larvae of several important groups of genera are essentially inseparable;(3) keys usually will not separate early instars; and (4) slide preparation of specimens is usually required, and even then diagnostic features are often difficult to see.

The collection of surface-floating pupal exuviae (SFPE) has only recently become a widely used method for the investigation of chironomids (e.g., Coffman 1973; Wilson and Bright 1973), although the idea is not new (Thienemann 1910; Lenz 1955). Surface-floating pupal exuviae have been used exten sively to monitor surface water and sediment quality in Western Europe and England (McGill et al. 1979; Ruse 1995a, b; Ruse and Wilson 1984; Ruse et al. 2000; Wilson 1977, 1980, 1987, 1989; Wilson and

Bright 1973; Wilson and McGill 1977; Wilson and Wilson 1983), and in Australia for measuring the effects of stream acidification on Chironomidae

(Cranston et al. 1997). In North America the methodology has been successfully used in studies ofphenology(Coffman 1973; Wartinbee and Coffman 1976), diel emergence patterns

(Coffman 1974), ecology and community composition (Blackwood et al 1995; Chou et al 1999; Ferrington 1998, 2000; Ferrington et al 1995; Kavanaugh 1988), microbial decomposition(Kavanaugh 1988), assessment of effects of point sources of enrichment (Coler 1984; Ferrington and Crisp 1989), heavy metal discharges (Hayford and Ferrington 2005), water and sediment quality (Ferrington 1993b), effects of agricultural prac tices (Barton et al 1995) and extreme flood events (Anderson and Ferrington 2013). The advantages of the SFPE technique are numerous and make it well suited for use in biological monitoring and assessment programs. Most generic and species-level taxa are easily separated in the pupal stage, even if a formal generic or species name cannot be assigned. Large numbers of specimens usually can be collected in a short time and standardized criteria

for subsampling and sampling frequency of SFPE

collected at natural or artificial blockades in streams

and along the windward shore in lentic ecosystems. Quantitative collections can be made using enclosures because all known species rise to the surface for eclosion. Because the collections are made from the sur

disturbed and most emerging species, regardless of larval microhabitat, are collected. Once a voucher

slide series for species present in a system has been identified, most specimens subsequently can be recog nized with a dissecting microscope. Because the pupal exuviae have no tissues (except in the thoracic horns), no clearing is required. All the diagnostic features are usually discernible on every specimen because the specimen is depressed by the cover glass into two dimensions. As there is no ambiguity about the age of the pupal exuviae, the size of a species is determined. On occasion, the larval and pupal exuviae as well as the adult of a specimen are collected entangled with each other, providing ideal material for association. Additional types of studies for which the collection of pupal exuviae is well suited include diversity (richness and actual taxonomic composition of faunas), phe nology (diel and annual), biogeography, size distribu tion of species, sex ratios, and production of adults. EXTERNAL MORPHOLOGY Larvae

Mature (fourth instar) chironomid larvae range in size from about 2 to 30 mm. There are three main

body divisions: head, thorax, and abdomen, all of which have structures that are used for generic-level diagnosis. These divisions may appear as pale yellow or white in preserved larvae or may be variously pigmented or patterned. Common pigmentation includes dark yellow, brown, black, or yellow with a black posterior margin in the case of the head capsule and yellow, green, blue, violet, rose, orange, or brown in the case of the thorax and abdomen. In addition, the

presence of hemoglobin in certain larvae produces a strong "blood red" color in living specimens, which often changes to a more diffuse red, red-orange, or brown color when preserved in ethanol. Head: The head is in the form of a completely sclerotized capsule, and is never fully retractile. Plates that have characters of diagnostic importance associ ated with them are the genae or lateral sclerites, the frontoclypeal apotome or dorsal sclerite, the labrum or

1122

Chapter 27 Chironomidae

anterodorsal sclerite (Figs. 27.142 and 27.143), and the mentum or medioventral sclerite (Figs. 27.1-27.4). Sensory structures situated on the dorsal aspect of the head capsule are the eyespots, antennae, and numerous setae, scales, and lamellae. Eyespots (Figs. 27.1-27.4) occur on the dorsal or dorsolateral aspects of the genae. Antennae (Figs. 27.1-27.3) also arise from the genae but occur anterior to the eyespots, except in Tanypodinae larvae, which have antennae that are retractile into the head capsule(Fig. 27.4). Setae originate from all head capsule sclerites, but in dorsal aspect usually only the S setae of the labrum are of diagnostic importance. The dorsal S setae (Figs. 27.1-27.3) are paired medial setae of the labrum and are referred to as SI through SIV setae. The anteriormost setal pair are the SI setae; setae SII through SIV originate in sequence progressing poste riorly. Various scales or lamellae (Figs. 27.246 and 27.248) may surround or originate in close proximity to the bases of the S setae, particularly SI. Structures associated with feeding show extreme variation among the genera of Chironomidae. In gen eral these structures are concentrated on the antero-

ventral aspect ofthe head capsule including the region of the mentum and the ventral surface of the labrum.

Included in this category are the pecten epipharyngis, premandibles, mandibles, maxillae, prementohypopharyngeal complex,and the mentum.(See Figs. 27.127.4 for the ventral perspective and orientation of these structures.) The pecten epipharyngis and premandibles origi nate from the ventral surface of the labrum. The

pecten epipharyngis generally consists of three scales that may be simple or have numerous apical teeth. In some genera the scales may be fused and the pecten epipharyngis thus appears as a single plate or bar. The premandibles are movable ancillary feeding struc tures and may have a blunt apical cusp or, in the case of some predatory genera, may have from one to several mesally projecting teeth (Fig. 27.142). Premandibles are vestigial or lacking in Tanypodinae and Podonominae larvae.

Paired mandibles and maxillae occur ventral to

the premandibles. The mandibles may have strong but blunt apical and lateral teeth(Figs.27.304-27.306) or may have an elongate and sharply pointed apical tooth (Fig. 27.14); the former is typical of nonpredatory species and the latter, of predatory species. The maxilla, which is usually lightly sclerotized and diffi cult to discern, bears a maxillary palp which is gener ally detectable on its anteroventral surface. In Tanypodinae larvae and in some Chironomini species (specifically "Harnischia Complex" genera such as

Robackia and Beckidia) the maxillary palp is more elongate and often multisegmented. The prementohypopharyngeal complex and the mentum occur on the medioventral aspect of the head capsule. In Tanypodinae larvae, the prementohypo pharyngeal complex is armed apically with a welldeveloped and readily seen ligula. In most other chironomid larvae the mentum may obscure the greater part ofthe prementohypopharyngeal complex and, as no well-developed structure homologous to the tanypod ligula is usually present, this structure may be difficult to locate. The current interpretation of the larval mentum holds that this structure is a

double-walled, medioventral plate consisting of a dorsomentum and a ventromentum. In Chironomini, Pseudochironomini, and Tanytarsini the ventromen

tum is greatly expanded laterally into striated ventromental(paralabial)plates which are quite conspicuous. With the exception of the Tanypodinae, the ventro mentum may be only slightly to moderately expanded laterally in the remaining chironomid genera. When only slightly expanded, the ventromental plates may appear to be absent or vestigial. In some genera the plates may be quite obvious and may even have a cardinal beard (i.e., elongate setae arising in definite rows)originating near the dorsal surface. In most chironomid genera, again excluding Tanypodinae, the dorsomental plate is usually well sclerotized and has conspicuous teeth. In these cases the term "mentum" is simply used to refer to the toothed margin of the dorsomental plate. Notable exceptions occur within certain genera such as' Acamptocladius and Protanypus in which the dorso mental plates do not meet on the ventral midline of the head capsule and the ventromental plate covers all (in the case of Acamptocladius)or most(in the case of Protanypus)ofthe dorsomental teeth. A more detailed discussion of the structure of the mentum is provided by Saether (1971a). In Tanypodinae larvae the ventromental plate appears to be membranous or lightly sclerotized and is difficult to see. The dorsomental plate is much reduced in Fentaneurini genera and is also difficult to discern. In the remaining Tanypodinae genera, the lateral aspect of the dorsomental plate is welldeveloped and gives the appearance of paired, heavily sclerotized, plate-like structures armed with teeth along the anterior margin. Medially the dorsomental plate has reduced sclerotization and often is not read ily distinguishable from the ventromental plate. Thorax: The larval thorax, consisting of the first three segments behind the head capsule, is only clearly demarcated from the remainder ofthe body segments

flagellum

lauterborn organ antennal blade

mandible

antenna

SI!seta SI seta

SI seta

■^^premandlble

labral lamella mandible

igecten epipharyngis

pecten epipharyngi

y-p

premandlble

maxillary palp maxilla

maxillary palp

mentum

maxilla

eyespot mentum

ventromental

eyespot

7

striated

plate length

ventromental plate

Figure 27.2

Figure 27.1 lauterborn organ—«

stalk of lauterborn organ—ij

anterior labral region

y

antennal blade

I'll antenna

1^1

antennal blade

retractile antenna maxillary palp mandible

spur or anteromesal process \

maxilla

campanlform sensillum

antennal tubercle

dorsomental teeth

SI seta

V-dorsomental plate

mandible labral lamella

llgula

pecten epipharyngis-

eyespot

premandlble-

^paraglossa

maxillary palp maxilla mentum

eyespot

^

I

striated ventromental plate

^7 Figure 27.3

Figure 27.1 Generalized Chironomini head capsule (ventral view). Figure 27.2 Generalized Orthocladiinae head capsule (ventral view).

Figure 27.4

Figure 27.3 Generalized Tanytarsini head capsule (ventral view).

Figure 27.4 Generalized Tanypodinae head capsule (ventral view).

1123

1124

Chapter 27 Chironomidae

in the late fourth instar. At this time the larva is often

referred to as a prepupa because the thorax is swollen

by developing pupal structures. Often structures of the pupa,such as the respiratory organ or pronounced setae, are visible, thus allowing association of the larva with the pupal stage. Ventrally, on the first tho racic segment, a pair of prolegs typically arise from a common base and bifurcate apically. However, in some genera no bifurcation may occur and a single proleg appears to exist, or the prolegs may be reduced or vestigial. The prolegs are generally armed apically with sclerotized claws, the size and structure of which

may be of specific importance. Numerous setae of species-level importance may arise from the thoracic segments, most notably a distinct ring of strong setae in some case-building Tanytarsini larvae and some Eukiejferiella larvae. Abdomen: The abdomen of the larva consists of

all post-thoracic segments. Structures ofthe abdomen used for generic identification include setation, prolegs, anal tubules, procerci, ventral tubules, and supraanal setae.

In all Tanypodinae larvae except Pentaneurini

genera, the lateral margins of at least the anterior five abdominal segments are equipped with a dense row of setae termed a lateral hair fringe (Fig. 27.5). In

Natarsia sp. the fringe is somewhat less dense but still distinguishable in most preserved specimens. A dis tinct lateral hair fringe also is present in the Orthocladiinae genera Stackelbergina and Xylotopus (Fig. 27.297). Most other abdominal setae patterns appear to have only species-level significance. As is the case with the prolegs of the thorax (anterior prolegs), those of the abdomen (posterior prolegs) show quite a bit of variation in structure. The posterior prolegs usually occur as paired struc tures with apical claws (Figs. 27.5, 27.298, 27.300), but they may occasionally be fused or extremely reduced (Figs. 27.301-27.303). Anal tubules occur on the anal segment in most chironomid genera and may originate above, between,or occasionally along the basal margin of the posterior prolegs. Anal tubules generally occur as two paired structures, but occasionally may consist of one or three pairs, or rarely be absent. On the dorsum of the preanal segment, a pair of

fleshy tubercles, or procerci, may occur (Figs. 27.346 and 27.347). One to several strong setae originate from the apex of the procerci and generally 0-2 weak lateral setae are present. Occasionally the procerci may have basal or lateral regions of sclerotization or pigmentation; basal spurs also may be present. Rarely the procerci are vestigial or lacking (Figs. 27.30027.303).

In some genera of Chironomini the lateral margins of the antepenultimate abdominal segment are produced into fleshy protuberances that are called ventral tubules. When present, these occur as

one or two pairs. In some species of the genus Goeldichironomus, the anteriormost pair bifurcates just beyond the base (Fig. 27.179). Supraanal setae occur on the anal segment, usually just dorsal to the anal tubules (Figs. 27.26, 27.346-27.347). Although these setae are generally only of species-level diagnostic importance, in the Tanypodinae genus Pentaneura and in the Podonominae genera Boreochlus and Paraboreochlus they appear to have generic diagnostic importance when used in concert with other body characters.

Pupae Chironomid pupae range in length from about 1.5 to 20 mm. There are three main body divisions: head, thorax, and abdomen. The morphology of the

pupa is almost completely external, since it is consid ered to be a modified larval integument within which the adult develops (Figs. 27.348-27.350). The struc tures of the pupa are best seen in the cast skins(pupal exuviae), and the general description below and keys to follow are based on exuviae. (For reference to

particular structures see the glossary.) Head: The head region of chironomid pupal exu viae consists primarily of eye, antenna, and mouthpart sheaths. The area of integument covering the vertex of the pharate adult head is the frontal apotome (Fig. 27.348). Thorax: The thoracic region ofchironomid pupal exuviae bears the leg, wing, and halter sheaths (Figs. 27.351-27.353). The thorax also has several groups of setae and the taxonomically very important thoracic horn (Fig. 27.349). Abdomen: The abdomen of chironomid pupal

exuviae consists of eight similar segments and one or more additional segments modified into anal lobes

and genital sheaths (Fig. 27.350). The terga (and sometimes sterna) often bear distinctive groupings of spines, recurved hooks and shagreen (Fig. 27.350). The anal lobes often bear a fringe of setae and/or two or more spine-like to hair-like macrosetae (Fig. 27.350).

The pigmentation of pupal exuviae is not as var ied as that of larvae. However, most species have a characteristic pigmentation even if it is the absence or near absence of pigment. In others, the thorax and abdomen are yellow, golden yellow, yellow brown, brown, dark brown, or gray. Many have such pig ment in a distinct pattern, especially on the abdomen.

Chapter 27 Chironomidae

Together with size and shape, the pigmentation of pupal exuviae may be an important aid in sorting and identification.

Adults

Adult chironomids range in size from about 1.5 to 20 mm. There are three main body divisions: head, thorax, and abdomen.

Head: The most conspicuous features of the head are the eyes and antennae. The antennae of most male chironomids are plumose(Fig. 27.818). The antennae ofthe females are not plumose and usually are shorter than those of the males and with fewer flagellomeres. However, some male antennae are not plumose and have a reduced number of flagellomeres (e.g., Fig. 27.819). Thorax: The adult thorax bears the legs, wings, and halteres. The legs of chironomids are moderately long, especially the first pair, and consist of a femur, tibia, and five tarsomeres. The distal ends ofthe tibiae

often carry one or two spines that are of taxonomic importance(Figs. 27.797-27.799,27.801-27.802). The fourth and fifth tarsomeres are infrequently cordiform (heart-shaped, Fig. 27.800) or trilobed (Fig. 27.820). The wings ofchironomids have a moderately complex venation, but with few crossveins (Figs. 27.772, 27.775-27.784). Nomenclature of wing veins has been disputed and variously interpreted. Consequently, we have chosen to retain the more traditional nomencla

ture of Sffither (1980). The thorax may bear groups of setae and is usually patterned with brown,black, yellow, or green (Figs. 27.770 and 27.773). Abdomen: The abdomen of adult chironomids

consist of eight segments plus several terminal seg ments modified as genitalia. Groups of setae usually are present on terga, sterna, and pleura and the abdo men may be patterned in brown, black, yellow, blue, or green (Figs. 27.770 and 27.773). The genitalia (termed hypopygium in the male) often have a com plex external and internal morphology and in the male have a large gonocoxite and terminal (usually) gonostylus (Figs. 27.771, 27.774, 27.785-27.796).

1125

pupae have been described. Additionally, the key to pupae includes some taxa that cannot, at present, be assigned to any known genus. Such taxa are desig nated as numbered genera in the keys to pupae; how ever,in some cases these numbered genera may simply be the undescribed pupae of named genera known only as adults, or aberrant pupae belonging to estab lished genera. Some ofthe numbered genera ofthe 4th edition of this book have now been assigned names and are so identified in this edition. To avoid confu

sion, those which are still identified only by number have the same number assigned to them as in the 4th edition even though this has produced gaps in the numbered genera sequence. The reader must keep in mind that the number of undiscovered immature

stages is large, based on the number of species known only from the adult stage and the frequency with which we encounter new forms of immatures. The

importance and value in conducting rearing programs for larvae and/or pupae are crucial to maximize asso ciation of life stages for species descriptions. Because many taxa are poorly described or unknown as lar

vae, a rearing program will provide much needed additional descriptions of immature stages. It is to be expected that large collections and/or collections from unusual habitats or geographical regions will contain species that do not fit the generic concepts used in these keys. The keys do not include genera belonging to subfamilies and tribes that have not yet been recorded from North America. 2. Collections from freshwater and brackish

water should be keyed in the main generic keys for larvae and pupae. Marine specimens are treated (usu ally above the level of genus) in separate keys. 3. The Orthocladiinae, which is the most diverse of

the chironomid subfamilies, has not yet been satisfac torily divided into tribes. Tentative names for a few of the orthoclad tribes are indicated by quotation marks. 4. Slides prepared for identifications may be tem porary mounts with water or, preferably, permanent mounts with any suitable medium such as Euparal or Canada balsam.

Larvae NOTES ON PREPARATION OF SPECIMENS AND USE OF KEYS

General

1. When possible, use the fourth larval instar. These may often be recognized by the swollen thorax of the prepupa.

1. The keys below include: freshwater Nearctic lar vae(to subfamily and genus),freshwater Nearctic pupae (to subfamily and genus), freshwater Nearctic adults (to tribe), and marine Nearctic larvae, pupae, and adults(to tribe). The freshwater keys to genus include all known Nearctic genera for which larvae and/or

side facing upward. The head must often be gently (sometimes not so gently) depressed to expose the mouthparts. Depending on the thickness of the abdo men, the head and abdomen should be placed under the same cover glass. This is usually not possible for

2. Sever the head and mount it with the ventral

1126

Chapter 27 Chironomidae

large larvae since the abdomen prevents the depres sion of the head. In such cases, mount the abdomen

under a separate cover glass, but on the same slide.

d. move the abdomen,dorsal side up(usually), to a position just below the thorax. e. place a cover glass over the specimen, depress ing it just enough to cause the exuviae to flat ten hut not distort.

Pupae 1. Pupae are best determined from pupal exuviae. The identification of pupae (not exuviae) is not easy since the structures are obscured by tissues. If possible, remove pharate adults from their pupal exuviae.

2. Before dissecting pupae or pupal exuviae examine the leg sheath arrangement and type of tho racic horn. Preparation of exuviae often disrupts the leg sheath pattern, and the thoracic horn (particularly of the Chironominae) is often extremely difficult to see.

3. Pupal exuviae are best prepared as follows (these steps should be carried out with the specimen in a drop of mounting medium on a microscope slide): a. separate the head and thorax from the abdomen. b. split the thorax along the mid-dorsal suture and open the thorax so that the two edges of the suture are on opposite sides of the specimen. c. turn the thorax so that the outer side of the

integument is facing upwards and arrange on the slide above the abdomen.

Adults

1. The keys below are designed for adult males. Males can be separated from females by their gener ally more slender abdomen, a rather conspicuous set of genital appendages, and,in most species, character istic plumose antennae. 2. Most adults can be identified to subfamily and many to tribe with the use of a dissecting microscope.To be certain, however,it is best to prepare the specimen for examination with a compound microscope. This is sometimes a tedious and involved process, but has been greatly simplified here for the purposes of a key to tribes. a. Remove and mount:

(1)one of each pair of legs (2) wings b. Remove the abdomen and heat it carefully in 10% KOH to remove the soft obscuring tissues (alternatively, the abdomen will clear standing for 12-24 hours at room temperature in 10% KOH); mount the abdomen dorsal side up, being particularly careful to orient the genitalia dorsal side up.

KEY TO TRIBES OR SUBFAMILIES OF NORTH AMERICAN FRESHWATER CHIRONOMIDAE LARVAE

1.

Antennae retractile (Fig. 27.4); prementohypopharyngeal complex with a 4-8 toothed ligula (Figs. 27.4, 27.13, 27.15, 27.16, 27.41-27.53, 27.63, 27.86); mentum almost entirely membranous or with dorsomental teeth arranged in conspicuous plates (Figs. 27.11, 27.12, 27.17-27.20, 27.25, 27.60, 27.62, 27.64-27.66) or longitudinal rows (Figs. 27.10, 27.13) TANYPODINAE(p. 1127)

r.

Antennae nonretractile (Figs. 27.1-27.3); prementohypopharyngeal complex without a toothed ligula. Mentum usually entirely toothed (Figs. 27.98-27.102, 27.128, 27.132-27.138, 27.145, 27.150-27.167, 27.171-27.178, 27.180-27.189, 27.192-27.195, 27.201-27.203, 27.211, 27.224, 27.225, 27.249-27.296, 27.311, 27.313-27.316, 27.319, 27.320, 27.322, 27.323, 27.332, 27.336-27.339, 27.340-27.342), or occasionally with weakly sclerotized or translucent

portions (Figs. 27.251, 27.252, 27.324, 27.328, 27.334), but never membranous

2

2(1').

Procerci long, at least 5 times as long as wide; premandibles absent... PODONOMINAE(p. 1177)

2'.

Procerci variable, rarely more than 4 times longer than wide; premandibles present (Figs. 27.1-27.3, 27.142, 27.329), usually well-developed and conspicuous

3(2').

Ventromental plates well-developed and with conspicuous striations throughout more than one-half their width (Figs. 27.98-27.102,

3

27.132-27.138, 27.145, 27.150-27.167, 27.171-27.178, 27.180-27.189, 27.192-27.195, 27.201-27.203, 27.211, 27.340-27.342) CHIRONOMINAE .... 4

Chapter 27 Chironomidae

3'.

4(3).

4'.

5(4'). 5'.

1127

Ventromental plates vestigial to well-developed (Figs. 27.249-27.284, 27.311, 27.315, 27.316, 27.322-27.324, 27.328, 27.332-27.334, 27.336-27.339), when well-developed never with striations although occasionally with setae underneath (Figs. 27.285-27.296, 27.313, 27.314) 6 Antenna arising from distinct tubercle (Figs. 27.103-27.107, 27.113, 27.114, 27.116-27.118, 27.131); first antennal segment elongate and usually at least slightly curved (Figs. 27.103-27.114); lauterborn organs very large and conspicuous (Figs. 27.103-27.112, 27.123, 27.131) or occurring at the apex of elongated stalks (Figs. 27.113, 27.114, 27.119, 27.120) TANYTARSINI(p. 1139) Antenna not arising from a distinct tubercle; if first antennal segment elongate then not curved as in Figs. 27.103-27.114, and lauterborn organs not large or occurring at the apex of elongated stalks 5 Outermost lateral teeth of mentum rounded and directed laterally; mentum as in Figs. 27.340-27.342 PSEUDOCHIRONOMINl' Outermost lateral teeth usually pointed and directed anteriorly (Figs. 27.132-27.138, 27.145, 27.150-27.167, 27.171-27.178, 27.180-27.189, 27.192-27.195,

6(3'). 6'. 7(6').

7'.

27.201-27.203, 27.211); mentum never as in Figs. 27.340-27.342 CHIRONOMINI(p. 1145) Mentum as in Fig. 27.322, 27.323, or 27.324 PRODIAMESINAE (p. 1180) Mentum not as in Fig. 27.322, 27.323, or 27.324 7 Third antennal segment with areas of reduced sclerotization which give the appearance of annulations (Figs. 27.330 and 27.331), or mentum similar to Fig. 27.328 and anterior labral region with large scales similar to Fig. 27.329 DIAMESINAE (p. 1176) Third antennal segment occasionally very small (Figs. 27.228, 27.235, 27.238-27.240, 27.244) but never with reduced sclerotization causing an annulated appearance; mentum and anterior labral region never as in Fig. 27.328 or 27.329 ORTHOCLADIINAE(p. 1159)

KEY TO THE GENERA OF NORTH AMERICAN FRESHWATER CHIRONOMIDAE LARVAE

Tanypodinae 1. Abdominal segments with a lateral hair fringe (Fig. 27.5); dorsomental teeth present in well defined plates (Figs. 27.11, 27.12), or arranged in longitudinal rows (Figs. 27.10, 27.13); head ratio 1.0 to 1

2

1'.

Abdominal segments lacking well defined lateral hair fringe; dorsomental teeth absent or extremely reduced; head ratio 1.5 or greater PENTANEURINI.... 16

2(1).

Dorsomental teeth arranged in longitudinal rows (Figs. 27.10, 27.13); ligula with 6-7 teeth (Figs. 27.15, 27.16); head capsule with

2'.

pronounced anterior taper (Fig. 27.6) CLINOTANYPODINI.... 3 Dorsomental teeth present in well defined plates (Figs. 27.11, 27.12); ligula with 4-5 teeth (Figs. 27.43, 27.45, 27.53); head capsule more rounded anteriorly (Fig. 27.7)

3(2).

3'.

4

Ligula with 6 teeth (Fig. 27.15); mandible strongly hooked (Fig. 27.14); ratio of head length to antennal length approximately 1.6 Clinotanypus Kieffer Ligula with 7 teeth (Fig. 27.16); mandible not strongly hooked; ratio of head length to antennal length approximately 2.6 Coelotanypus Kieffer

'In North America the tribe Pseudochironomini is represented by the genera,Pseudochironomus and Manoa,The characteristics used in this key to identify specimens to the tribe level thus also serve to identify to the generic level; no additional key to genus is provided for this tribe. Larvae of Manoa have not been reared and are only provisionally associated, Pseudochironomus can be differentiated from the single species of Manoa by referring to Jacobsen and Perry (2002).

1128

Chapter 27 Chironomidae

4(2').

Dorsomental plates each with 13-15 teeth

ANATOPYNIINI^

4'.

Dorsomental plates with 9-7 or fewer teeth

5(4').

Dorsomental plates with 2-3 teeth (Fig. 27.17); antennae about one-third length of head capsule and ratio of length of first antennal segment to length of remaining segments less than 3.5 NATARSIINI—Natarsia Fittkau

5'.

Dorsomental plates with more than 3 conspicuous teeth (Figs. 27.18-27.20, 27.25); antennae shorter than one-third the head capsule length, or, if longer than one-third the length of head capsule, the ratio of the length of the first antennal segment to length of remaining segments always 3.5 or greater

5

6

6(5').

Mandible with bulbous base and very minute lateral teeth (Fig. 27.54); ligula with 5 pale yellow to light brown teeth; teeth of ligula forming a convex, arch or (less commonly)of equal lengths(Fig. 27.44) TANYPODINI—Tanypus Meigen

6'.

Mandible without conspicuous bulbous base; ligula with 4-5 teeth (Figs. 27.41, 27.42, 27.43, 27.45, 27.53); when 5 pale yellow or light brown teeth are present on ligula the teeth form a concave arch (Figs. 27.41, 27.42)

7(6')

Ligula with 5 or (less commonly)4 black teeth (Figs. 27.53, 27.45); paraglossae with 1 main tooth and 1-7 accessory teeth on each side (Figs. 27.30, 27.31)

7

PROCLADIINI ....8

7'.

Ligula with 5 or 4 light yellow to brown teeth (Figs. 27.41-27.43); paraglossae pectinate (Fig. 27.34); unevenly bifid (Fig. 27.33), or with only accessory teeth on outer margin (Fig. 27.32)

8(7).

Antennal blade more than twice as long as the combined lengths of antennal segments 2 through 4(Fig. 27.22); ligula usually with 4 teeth (Fig. 27.45). . Djalmabatista Fittkau

8'.

Antennal blade subequal in length to the combined lengths of antennal segments 2 through 4(Fig. 27.21); ligula normally with 5 teeth (Fig. 27.53) Procladius Skuse

9(7').

Ligula with 4 teeth (Fig. 27.43); paraglossae pectinate (Fig. 27.34); mandible with row of 4 or more inner teeth (Figs. 27.58, 27.59) MACROPELOPIINT (in part)

10

Ligula with 5 teeth (Figs. 27.41, 27.42); paraglossae not pectinate (Figs. 27.32, 27.33); mandible not as in Figs. 27.58, 27.59

11

9'. 10(9).

Lateral dorsomental teeth closely appressed (Fig. 27.25); small claws of posterior prolegs as in Fig. 27.40 or simple

10'.

Lateral dorsomental teeth not closely appressed (Fig. 27.19); small claws of posterior prolegs basally ovoid (Fig. 27.39); or simple

11(9').

9

Dewtanypus Roback Psectrotanypus Kieffer

Inner margin of dorsomental plate produced medially into one (Fig. 27.18) or two points (Fig. 27.60)

30

11'.

Inner margins of dorsomental plate rounded (Fig. 27.20)

12

12(11').

Mandible with 2 or 3 complete or partial rows of inner teeth (Fig. 27.61); dorsomental plates not interrupted in the middle (Fig. 27.62); 3 middle teeth of ligula all about the same length (Fig. 27.63) FITTKAUIMYIINI—Fittkauimyia Karunakaran

12'.

Mandible with 4 or fewer inner teeth, not forming multiple rows; dorsomental plates separated in the middle; 3 middle teeth of ligula differing in length MACROPELOPIINI(in part)

13(12').

13

Pseudoradula widest near base of m-appendage and tapering in width to apex, granules more coarse in proximal half of pseudoradula (Fig. 27.64) Radotanypus Fittkau and Murray

^Not recorded in North America.

Chapter 27 Chironomidae

1129

head capsule

thorax

Figure 27.7

Figure 27.6

Figure 27.8

Figure 27.9

Figure 27.5 abdomen

Rgure 27.11

Figure 27.10

Figure 27.12 rocercus

poaterior proleg

Figure 27.15

t I

* ^

Figure 27.16

Figure 27.14

Figure 27.13

Figure 27.5 Generalized Tanypodinae larva (dorsal view) showing iateral hair fringe. Figure 27.6 Dorsal view of Clinotanypus sp. head capsule.

Figure 27.7 Dorsal view of Procladius sp. head capsule. Figure 27.8 Dorsal view of Nilotanypus sp. head capsule. Figure 27.9 Dorsal view of Conchapetopia sp. head capsule. Figure 27.10 Longitudinal arrangement of dorsomental teeth in Coelotanypus concinnus (Goquillett).

Figure 27.11

Dorsomental teeth of Tanypus sp.

Figure 27.12 Dorsomental teeth, ventromentum and prementohypopharyngeal element (in ventral view) of Procladius sp. (redrawn from Roback [1977]). Figure 27.13 Longitudinal arrangement of dorsomental teeth on mental region of Clinotanypus sp.(ventral view).

Figure 27.14 Mandible of Clinotanypus sp. Figure 27.15 Ligula of Clinotanypus sp. Figure 27.16 Ligula of Coelotanypus sp.

1130

Chapter 27 Chironomidae

13'.

Pseudoradula narrowest near median edges of dorsomental plates and expanded apically (Fig. 27.65) or pseudoradula of uniform width but granules weaker near base (Fig. 27.66)

14(13').

All teeth of ligula directed anteriorly (Fig. 27.42); length of palpus at least 4 times greater than width at midlength and campaniform sensillum occurring in basal one-third of palpus (Fig. 27.29); pseudoradula of uniform width but granules weaker near base (Fig. 27.66) First lateral teeth of ligula outcurved (Fig. 27.41); length of palpus 2.9-4.1 times

14'.

the width at midlength; campaniform sensillum located near middle of palpus (Figs. 27.23, 27.24); pseudoradula narrowest near median edges of dorsomental plates (Fig. 27.65)

15(14').

Dorsomental plate with 7 teeth; palpus about 4.1 times longer than wide at midlength (Fig. 27.24); 2nd antennal segment 4.2-4.7 times longer than maximum width; all ventrolateral setae of mandible simple (Fig. 27.67)

14

45

15

Alotanypm Roback

15'.

Dorsomental plate with 4-5 teeth (Fig. 27.20); palpus 2.9-3.2 times longer than width at midlength (Fig. 27.23); 2nd antennal segment 2.1-2.5 times longer than maximum width; ventrolateral seta 1 of mandible simple, setae 2 and 3 multibranched (Fig. 27.68) Apsectrotanypus Fittkau

16(1').

Maxillary palp with 2 or more basal segments (Figs. 27.27, 27.28)

17

16'.

Maxillary palp with only one basal segment

18

17(16).

Maxillary palp with 2 basal segments and campaniform sensillum very large, about as large as width of palp, and located between first and second segments (Fig. 27.70); pseudoradula broad, widened toward base of m-appendage (Fig. 27.69); dark claws of posterior prolegs, if present, pectinate or bifid (Fig. 27.35) Zavrelimyia (Paramerina)(in part)

17'.

Maxillary palp with more than two basal segments, or when only two basal segments then campaniform sensillum much smaller than width of palp; pseudoradula narrow, widest near middle and with granules in parallel, longitudinal rows(Fig. 25.71); dark claws of posterior prolegs, if present, all simple Ablabesmyia Johannsen

18(16').

Lauterborn organs' at tip of second antennal segment well sclerotized and solidly fused with segment so that second segment appears to envelope or flank most of the third segment(Fig. 27.72); third and fourth segments of antenna about equal in length

18'.

31

Lauterborn organs small or vestigial, less than 1/3 the length of segment 3, or not fused to apex of segment 2; fourth segment of antenna variable in length, but usually substantially shorter than third segment

19

19(18').

Anterior margin of ligula straight (i.e., all teeth ending at more or less the same point)(Figs. 27.50, 27.51)

20

19'.

Anterior margin of ligula with median tooth distinctly longer than or shorter than the 1st lateral and/or 2nd lateral teeth (Figs. 27.46-27.49, 27.52)

22

20(19).

Posterior proleg with 1 claw darkly pigmented; supraanal seta strong, and originating from distinct papillae; anal tubules longer than posterior prolegs (Fig. 27.26) Pentaneura Philippi

20'.

All claws of posterior prolegs unicolorous; supraanal seta weak, or, if conspicuous, not originating from distinct papillae; anal tubules shorter than posterior prolegs

21

^ (Caution-Sometimes it is difficult to see the structure of the lauterborn organs and the lengths of the distal segments of the antennae. If these structures cannot be seen, continue to couplet 19).

Chapter 27 Chironomidae

1131

dorsomental teeth

\

/

Figure 27.18 Figure 27.19

Figure 27.17

n

^

V

Figure 27.23

Figure 27.20 Figure 27.21

Figure 27.25

Figure 27.24 Figure 27.22

Figure 27.26 Figure 27.27

Figure 27.17 Arrangement of dorsomental teeth in Natarsia sp. Figure 27.18 Arrangement of dorsomental teeth in Brundlniella sp. Figure 27.19 Arrangement of dorsomental teeth In Psectrotanypus dyari (Coqulllett). Figure 27.20 Arrangement of dorsomental teeth in Apsectrotanypus sp. Figure 27.21 Apex of antenna of Procladius sp. Figure 27.22 Apex of antenna of Djalmabatista pulcher(Johannsen). Figure 27.23 Basal segment of maxillary palpus of Apsectrotanypus sp.

Figure 27.28

Figure 27.29

Figure 27.24 Basal segment of maxillary palpus of Alotanypus venustus (Coqulllett). Figure 27.25 Arrangement of dorsomental teeth in Derotanypus alaskensis (Malloch). Figure 27.26 Procerci, supraanal setae, anal tubules, and posterior prologs of Pentaneura sp. Figure 27.27 Maxillary palpus of Ablabesmyia (Ablabesmyia) mallochi (Walley). Figure 27.28 Maxillary palpus of Ablabesmyia (Ablabesmyia) parajanta Roback.

Figure 27.29 Basal segment of maxillary palpus of Macropelopla sp.

1132

Chapter 27 Chironomidae

Figure 27.30

Figure 27.31 Figure 27.33

Figure 27.32

Figure 27.34

Figure 27.38 Figure 27.37

Figure 27.36 Figure 27.35

Figure 27.40 Figure 27.39

Figure 27.30 Paraglossa of Procladius sp. Figure 27.31 Paraglossa of Djalmabatista pulcher (Johannsen). Figure 27.32 Paraglossa of Alotanypus venustus (Coquillett)(redrawn from Roback [1978]).

Figure 27.33 Paraglossa of Macropelopia decedens (Walker)(redrawn from Roback [1978]). Figure 27.34 Paraglossa of Psectrotanypus dyari (Coquillett).

Figure 27.35 Pigmented claws of posterior prologs of Zavrelimyia {Paramerina) smithae (Sublette)(redrawn with modification from Roback [1972]).

Figure 27.36 Pectinate claw of posterior prolog of Nilotanypus sp.

Figure 27.41

Figure 27.42

Figure 27.43

Figure 27.37 Bifid claw of posterior prolog of Labrundinia sp.

Figure 27.38 Bifid claw of posterior prolog of Zavrelimyia sp.

Figure 27.39 Basally ovoid claw of posterior prolog of Psectrotanypus dyari (Coquillett).

Figure 27.40 Smallest claw of posterior prolog of Derotanypus alaskensis (Malloch)(redrawn from Roback [1978]).

Figure 27.41 Ligula of Alotanypus venustus (Coquillett). Figure 27.42 Ligula of Macropelopia sp. Figure 27.43 Ligula of Psectrotanypus dyari (Coquillett).

Chapter 27 Chironomidae

21(20').

21'.

22(19'). 22'.

23(22). 23'.

1133

One daw of posterior proleg bifid (Fig. 27.38); head capsule unicolorous; ratio of length of 1st antennal segments to combined lengths of remaining antennal segments about 3.1 Zavrelimyia Fittkau (in part) 43 All claws of posterior prolegs simple; posterior 1/4 of head capsule with conspicuous blackish brown pigmentation; ratio of length of 1 st antennal segment to combined lengths of remaining antennal segments about 2.5 Zavrelimyia(Paramerina)(in part) Median tooth of ligula longer than 1st lateral teeth (Figs. 27.48, 27.49) 23 Median tooth of ligula shorter than (Figs. 27.46, 27.47) or equal in length to 1st lateral teeth; when equal in length to 1st lateral teeth, then all 3 median teeth shorter than 2nd lateral teeth (Fig. 27.52) 24 Posterior prolegs with 1 bifid claw (Fig. 27.37); 2nd antennal segment usually distinctly more darkly pigmented than 1st segment; ligula as in Fig. 27.48 ... Labrundinia Fittkau Posterior prolegs with 1 pectinate claw (Fig. 27.36); all antennal segments unicolorous; ligula as in Fig. 27.49 Nilotanypus Kieffer

24(22').

Second antennal segment with distinct dark brown pigmentation

24'.

All antennal segments unicolorous

25(24').

Head capsule with granular appearance visible at lOOX magnification; body with undulate wrinkles; ratio of length of 1st antennal segment to combined length of remaining segments 6.0-7.5 Guttipelopia Fittkau Head capsule lacking granular appearance; body smooth; ratio of length of 1 st antennal segment to combined lengths of remaining segments variable,

25'.

Monopelopia Fittkau (in part) 25

but rarely greater than 5.5

26(25'). 26'. 27(26).

26

Mandible with at least 1 large conspicuous tooth along inner margin (Figs. 27.56 and 27.57) Inner margin of mandible smooth or with only very small teeth present Mandible with 1 large, pointed tooth and 1-2 smaller teeth present on inner margin (Fig. 27.56)

27'. 28(26').

27 28

Larsia Fittkau

Mandible with 1 large blunt tooth and 1 small tooth present on inner margin (Fig. 27.57) Krenopelopia Fittkau (in part) Head with posterior half pigmented dark brown; mandible with 1 small tooth and well-developed accessory blade (Fig. 27.55); larvae apparently restricted to mats of blue-green algae occurring on steep rock outcrops over which small

28'.

volumes of water trickle Hudsonimyia Roback Head capsule unicolorous; mandible not as in Fig. 27.55; larvae occurring in a wide variety of habitats

29(28').

29'.

29

Mandible with small but distinct teeth along inner edge; ratio of length of 1st antennal segment to combined lengths of remaining segments 3.4-3.8; campani form sensillum in basal 1/3 of maxillary palp; ventrolateral seta 1 of mandible reduced and in small pit, setae 2 and 3 long and simple (Fig. 27.73) Trissopelopia Kieffer Teeth of inner margin of mandible very minute, indistinct, or lacking; ratio of length of 1st antennal segment to combined lengths of remaining segments variable but usually 3.8-5.3; campaniform sensillum of maxillary palp variable, but usually in apical 1/3; at least ventrolateral seta 3 of mandible bifid or

multibranched (Fig. 27.75)

Thienemannimyia group"* . ... 36

■"The Thienemannimyia group consists of the genera Arctopelopia, Conchapelopia, Helopelopia, Meropelopia, Rheopelopia, Telopelopia and Thienemannimyia. Larvae of these genera are very difficult to identify to genus. Characters given in this key are based on fourth instar larvae, and probably will not work for earlier instars. Earlier instars should only be identified to ^'Thienemannimyia group sp.". Identification should be confirmed by rearing larvae to pupal or adult stages.

1134

Chapter 27 Chironomidae

30(11).

Ventrolateral setae 1 and 3 of mandible simple, seta 2 bifid (Fig. 25.74); campaniform sensillum of maxillary palp located in basal 1/3 Macropelopia (Bethbilbeckia)

30'.

Ventrolateral seta 1 simple, setae 2 and 3 multibranched (Fig. 21.15);

31(18). 31'. 32(31).

32'. 33(32'). 33'.

34(31'). 34'.

35(34). 35'.

campaniform sensillum of maxillary palp located in middle 1/3 Brundiniella Roback Anterior margin of ligula straight (i.e., all teeth ending at more or less the same point) or slightly convex 32 Anterior margin of ligula concave 34 Paraglossae with two inner teeth (Fig. 27.76); two smallest claws of posterior prolegs each with one large inner tooth and some spines on outer margin, three slightly larger claws with a single spine-like inner tooth, and nine largest claws with only fine spines on inner margin (Fig. 27.77) Denopelopia Roback and Rutter Paraglossae with one inner tooth (Fig. 27.33); at least smaller claws of posterior prolegs with several large inner teeth (Fig. 27.78 or 27.81) 33 Second antennal segment brown; campaniform sensillum in apical 1/2 of first antennal segment; some claws of posterior prolegs dark Xenopelopia Fittkau Second antennal segment not dark; campaniform sensillum in basal 1/2 of first antennal segment; all claws of posterior prolegs pale Telmatopelopia Fittkau Pecten hypopharyngis with fewer than 13 unequal teeth (Fig. 27.79); teeth of ligula deeply concave; campaniform sensillum in apical 1/2 of first antennal segment 35 Pecten hypopharyngis with 15 teeth of approximately equal lengths (Fig. 27.80); teeth of ligula only moderately concave; campaniform sensillum in basal 1/2 of first antennal segment Pentaneurella Fittkau and Murray Second antennal segment brown; one claw of posterior prolegs dark, dark claw and some smaller pale claws with large inner teeth (Fig. 27.81) .... Monopelopia Fittkau (in part) Second antennal segment pale, same color as basal segment; all claws of posterior prolegs unicolorous and pale 44

36(29').

B-seta of maxillary palp 2-segmented (Fig. 27.82)

36'. 37(36').

B-seta of maxillary palp 3-segmented (Fig. 27.83) 37 Mandible strongly curved in apical 1/2, no teeth or only one very small inner tooth visible on mandible (Fig. 27.84) Rheopelopia Fittkau Mandible uniformly weakly curved, 2 small inner teeth visible on mandible (Fig. 27.85) 38 Ventrolateral seta 3 and sensillum minusculum clearly located in basal 1/3 of mandible (Fig. 27.85); middle tooth of ligula about as long as width at base

37'.

38(37').

(Fig. 27.86) 38'.

39(36). 39'.

39

Helopelopia Roback

Ventrolateral seta 3 and sensillum minusculum occurring more distally along

outer margin of mandible, generally located in middle 1/3 of mandible (Fig. 27.87); middle tooth of ligula clearly longer than width at base (Fig. 27.47) Conchapelopia Fittkau (in part) Pecten hypopharyngis with about 25 teeth (Fig. 27.88); pseudoradula strongly widened in middle (Fig. 27.89) Arctopelopia Fittkau Pecten hypopharyngis with about 20-22 teeth; pseudoradula either strongly widened proximally (Fig. 27.90) or uniform in width (Fig. 27.91)

40

40(39').

Pseudoradula strongly widened proximally (Fig. 27.90)

41

40'.

Pseudoradula uniform in width (Fig. 27.91)

42

41(40). 41'.

Two inner teeth of mandible about same size (Fig. 27.92). ..Thienemannimyia {Hayesomyia)in part Inner basal tooth of mandible larger than distal tooth (Fig. 27.93) Telopelopia Roback

Chapter 27 Chironomidae

Figure 27.45

Figure 27.44

Figure 27.46

Figure 27.49

Figure 27.46

Figure 27.52

Figure 27.50

1135

Figure 27.47

Figure 27.53

Figure 27.55

Figure 27.56

Figure 27.57

Figure 27.58

'^'9"''®

Figure 27.54

Figure 27.44 Three types of liguia of Tanypus species: (A) Tanypus (Apelopia) neopunctipennis Sublette;(B) Tanypus (Tanypus) punctipennis Meigen;(C) Tanypus (Tanypus) possibly concavus Roback.(Tanypus (Tanypus) possibly concavus Roback redrawn from Roback [1977]). Figure 27.45 Liguia of Djalmabatista pulcher (Johannsen). Figure 27.46 Liguia of Larsia sp. Figure 27.47 Liguia of Conchapelopia sp. Figure 27.48 Liguia of Labrundinia sp. Figure 27.49 Liguia of Nitotanypus sp.

Figure 27.50 Liguia of Pentaneura sp. Figure 27.51 Liguia of Zavretimyia sp. Figure 27.52 Liguia of Ablabesmyia annulata (Say). Figure 27.53 Liguia of Procladius sp. Figure 27.54 Mandible of Tanypus sp. Figure 27.55 Apex of mandible of Hudsonimyia karelena Roback (redrawn with modification from Roback [1979]). Figure 27.56 Mandible of Larsia sp. Figure 27.57 Mandible of probable Krenopelopia sp. Figure 27.58 Mandible of Derotanypus. Figure 27.59 Mandible of Psectrotanypus sp.

Figure 27.62

Figure 27.60

Figure 27.61 Figure 27.63

Figure 27.64 Figure 27.65

Figure 27.67

Figure 27.66

Figure 27.60 Dorsomental plates and m-appendage of Macropelopia (Bethbilbeckia) sp. Figure 27.61 Mandible of Fittkauimyia sp. Figure 27.62 Dorsomental plates and m-appendage of Fittkauimyia sp. Figure 27.63 LIgula of Fittkauimyia sp. Figure 27.64 Dorsomental plates and m-appendage of Radotanypus sp.

1136

Figure 27.68

Figure 27.65 Dorsomental plates and m-appendage of Apsectrotanypus sp.

Figure 27.66 Dorsomental plates and m-appendage of Macropelopia sp.

Figure 27.67 Mandible of Aiotanypus sp. Figure 27.68 Mandible of Apsectrotanypus sp.

T, Figure 27.71

Figure 27.69

Figure 27.70

0 Figure 27.72

Figure 27.73

Figure 27.75 Figure 27.74

Figure 27.76

Figure 27.69 Dorsomentum, m-appendage and pseudoradula of Zavrelimyia (Paramerina) sp. Figure 27.70 Maxillary palp of Zavrelimyia (Paramerina) sp. Figure 27.71 Dorsomentum, m-appendage and pseudoradula of Ablabesmyia sp. Figure 27.72 Lateral and ventral views of apex of second antennal segment of Monopelopia sp.

Figure 27.73 Mandible of Trissopeiopia sp. Figure 27.74 Mandible of Macropelopia (Bethbilbeckia) sp. Figure 27.75 Mandible of Brundiniella sp. Figure 27.76 Paraglossa of Denopelopia sp.

1137

n

Figure 27.77 Figure 27.78

Figure 27.80 Figure 27.79

Figure 27.81

I 61 Figure 27.83

u Figure 27.82

Figure 27.84

Figure 27.85 ^

>< #0.;

ss.;-^*,V • '•

Figure 27.86

Figure 27.77 Two smallest claws, one larger claw and one largest claw of posterior prologs of Denopelopia sp. Figure 27.78 Claws of posterior prologs of Telmatopelopia sp. Figure 27.79 Pecten hypopharyngis of Krenopelopia sp. Figure 27.80 Pecten hypopfiaryngis of Pentaneurella sp. Figure 27.81 Claws of posterior prologs of Monopelopia sp.

1138

Figure 27.82 Maxillary palp and B-seta of Arctopelopla sp. Figure 27.83 Maxillary palp and B-seta of Conchapelopia sp. Figure 27.84 Mandible of Rheopelopia sp. Figure 27.85 Mandible of Helopelopia sp. Figure 27.86 LIgula of Helopelopia sp.

Chapter 27 Chironomidae

42(40').

1139

Distance between origin of ventrolateral setae 2 and 3 of mandible about 1.5 times the distance between ventrolateral setae 1 and 2

(Fig. 27.94) 42'.

Thienemannimyia Fittkau (in part)

Distance between origin of ventrolateral setae 2 and 3 of mandible about 3.0 times the distance between ventrolateral setae 1 and 2(Fig. 27.95)

Meropelopia Roback

43(21).

Length of maxillary palpus 5-6 times greater than width; campaniform sensillum located in basal 0.4 of length of maxillary palpus

Zavrelimyia (Reomyia)

43'.

Length of maxillary palpus approximately 4 times greater than width; campaniform sensillum located 0.5-0.7 from base of maxillary palpus

Zavrelimyia Fittkau

44(35').

All claws of posterior prolegs without large inner teeth

44'.

Three smaller claws of posterior prolegs with large inner teeth

45(14).

Pseudoradula of uniform width but granules weaker toward base (Fig. 27.66); labral sclerite weak or absent

45'.

Krenopelopia Fittkau (in part) Monopelopia (Cantopelopid) Macropelopia Thienemann

Pseudoradula absent basally; labral sclerite present and well-developed (see Niitsuma and Watson 2009) Bilyjomyia Niitsuma and Watson

Tanytarsini 1. Ventromental plates well separated, pointed at anteromedial edge (Fig. 27.99); larvae construct portable cases made of sand grains and/or detritus (Figs. 27.96, 27.97)

1'.

2(1).

2'.

2

Ventromental plates almost meeting at ventral midline of head capsule, anteromedial edge generally rounded or truncate (Figs. 27.98, 27.100-27.102); larvae may construct tubes or filtering structures but never portable cases Lauterborn organs originating alternately at different heights on 2nd antennal segment (Figs. 27.103, 27.104); antennal segment 2 subequal to or longer than the combined lengths of segments 3-5; larval case always straight (Fig. 27.96) Lauterborn organs originating on opposite sides at apex of 2nd antennal segment (Figs. 27.105-27.107); antennal segment 2 distinctly shorter than the combined lengths of segments 3-5; larval case straight or curved (Figs. 27.96, 27.97)

6

3

4

3(2).

Distal lauterborn organ arising preapically on 2nd antennal segment(Fig. 27.104); antennal segment 2 subequal in length to the combined lengths of segments

3'.

Distal lauterborn organ arising from apex of 2nd antennal segment(Fig. 27.103); antennal segment 2 distinctly longer than the combined lengths of segments 3-5(Fig. 27.103) Stempellinella Brundin Antennal tubercle with a conspicuous anteromesally projecting palmate process (Figs. 27.105, 27.131); larval case curved (Fig. 27.97) 21 Antennal tubercle lacking palmate process but with a single strong anteromesally directed spine (Figs. 27.106, 27.107); larval case curved or straight 5 Lauterborn organs on distinct stalks, stalks subequal to or greater in length than antennal segment 3(Fig. 27.106); larval case curved (Fig. 27.97) Constempellina Brundin Lauterborn organs not conspicuously stalked (Fig. 28.107), or if stalks are apparent then much shorter in length than antennal segment 3; larval case straight

3-5(Fig. 27.104)

4(2'). 4'. 5(4'). 5'.

(Fig. 27.96)

Zavrelia Kieffer

Thienemanniola Kieffer

6(1').

Mentum with only 3 distinct teeth (Fig. 27.101); mandible blunt distally, lacking apical and lateral dents (teeth)(Fig. 27.115) Corynocera Zetterstedt

6'.

Mentum with more than 3 distinct teeth; mandible with at least 3 teeth

7

1140

Chapter 27 Chironomidae

Figure 27.88

Figure 27.87

Figure 27.89

Figure 27.91

Figure 27.90

Figure 27.87 Mandible of Conchapelopia sp. Figure 27.88 Pecten hypopharyngis of Arctopelopia sp.

Figure 27.89 Dorsomentum, m-appendage and pseudoradula of Arctopelopia sp.

Figure 27.90 Dorsomentum, m-appendage and pseudoradula of Thienemannimyia (Hayesomyia) sp. Figure 27.91 Dorsomentum, m-appendage and pseudoradula of Thienemannimyia sp.

Chapter 27 Chironomidae

Figure 27.92 Figure 27.93

Figure 27.94

Figure 27.95

Figure 27.92 Mandible of Thienemannimyia (Hayesomyia)senata. Figure 27.93 Mandible of Telopelopia okoboji.

Figure 27.94 Mandible of Thienemannimyia sp. Figure 27.95 Mandible of Meropeiopia sp.

1141

1142

Chapter 27 Chironomidae

Figure 27.98

Figure 27.96

Figure 27.97

Figure 27.100

Figure 27.103

Figure 27.99

Figure 27.104

Figure 27.101

Figure 27.105

Figure 27.96 Portable sand case of Stempellinella sp. Figure 27.97 Portable sand case of Constempellina sp. Figure 27.98 Mentum of Rheotanytarsus sp. Figure 27.99 Mentum of Constempellina sp. Figure 27.100 Mentum of Cladotanytarsus (Lenzlella) sp. Figure 27.101 Mentum of Corynocera sp.(redrawn from Hirvenoja [1961]).

Figure 27.102

Figure 27.106

Figure 27.107

Figure 27.102 Mentum of Sublettea sp. Figure 27.103 Antenna of Stempellinella sp. Figure 27.104 Antenna of Zavrelia sp.(redrawn from modification from Bause [1913]). Figure 27.105 Antenna of Stempellina sp. Figure 27.106 Antenna of Constempellina sp. Figure 27.107 Antenna of Thienemanniola sp. (redrawn wltti modification from Lefimann [1973]).

Chapter 27 Chironomidae

1143

7(6').

Stalks of lauterborn organs appearing annulated along three-fourths of their length (Fig. 27.119) Tanytarsus van der Wulp (in part)

T. 8(7').

Stalks of lauterborn organs never appearing annulated Second antennal segment annulated (Fig. 27.120)

8'.

Second antennal segment not annulated

9(8').

Stalks of lauterborn organs less than 1.2 times combined lengths of antennal segments 3-5 (Figs. 27.108-27.112)

11

9'.

Stalks of lauterborn organs greater than 1.25 times (i.e., distinctly longer than) the combined lengths of antennal segments 3-5 (Figs. 27.113, 27.114)

10

Antennal tubercle with a straight or anteromedially curved spur originating from medial edge (Figs. 27.114, 27.116-27.118)

17

Antennal tubercle without a straight or anteromedially curved spur (Fig. 27.113)

19

11(9).

Lauterborn organs less than 0.40 times the length of their stalks (Fig. 27.112)

14

11'.

Lauterborn organs about as long as or longer than the length of their stalk (Figs. 27.108-27.Ill)

12

10(9'). 10'.

12(11). 12'.

13(12).

13'.

14(11).

14'. 15(14).

8 Tanytarsus van der Wulp (in part) 9

Lauterborn organs about as long as their stalk; lauterborn organs broad and usually with visible longitudinal striations (Figs. 27.108, 27.109) 13 Lauterborn organs at least one-third longer than their stalk (Fig. 27.110); or if subequal in length then organs oval and lacking longitudinal striations (Fig. 27.111); stalks sometimes vestigial or absent(Fig. 27.123) 16 Mentum with 2nd lateral tooth smaller than both 1st and 3rd lateral teeth (Fig. 27.100); 2nd antennal segment membranous in apical half and subequal in length to 3rd antennal segment(Fig. 27.108) ... Cladotanytarsus (Lenziella) Mentum with all lateral teeth decreasing in size in an orderly manner, or if 2nd lateral teeth are reduced then 3rd antennal segment distinctly longer than 2nd segment(Fig. 27.109), and 2nd segment not membranous throughout the apical one half Cladotanytarsus Kieffer Mentum with 5 pairs of lateral teeth 15 Mentum with 4 pairs of lateral teeth; combined length of antennal segments 2-5 subequal in length to segment 1 in 4th instar specimens Neozavrelia Goetghebuer Distal portion of 2nd antennal segment greatly expanded (Fig. 27.122); mentum strongly arched (Fig. 27.102) Sublettea Roback

15'.

Distal portion of 2nd antennal segment only moderately expanded (Fig. 27.121); mentum not strongly arched (Fig. 27.98) Rheotanytarsus Thienemann and Bause

16(12').

Larvae marine; first antennal segment short, about as long as the combined lengths of the remaining segments(Fig. 27.123); pecten epipharyngis consisting of 3 lobes with apical dissections (Fig. 27.124) Pontomyia Edwards Larvae occurring in a wide variety of aquatic habitats; first antennal segment much longer than the combined lengths of the remaining segments (Figs. 27.110, 27.111); pecten epipharyngis consisting of 3-5 scale-like lobes without apical dissections(Fig. 27.125) Paratanytarsus Thienemann and Bause Premandible with 3-4 slender, pointed teeth (Fig. 27.127) Tanytarsus van der Wulp (in part) Premandible with only 2 broader teeth (Fig. 27.126) 18

16'.

17(10). 17'. 18(17'). 18'.

Antennal blade short, subequal in length to antennal segment 3; mentum strongly arched, median 3 teeth set off from remaining 4 lateral teeth (Fig. 27.128). Micropsectra (in part) Antennal blade longer than antennal segment 3, often subequal in length to segment 2; mentum not strongly arched, median tooth often appearing tripartite but well separated from remaining 5 lateral teeth Micropsectra Kieffer (in part)

1144

Chapter 27 Chironomidae

LJ Figure 27.108

Figure 27.110

Figure 27.109

u Figure 27.111 Figure 27.112

Figure 27.121 Figure 27.115

Figure 27.119

Figure 27.116

Figure 27.122

Figure 27.113

Figure 27.114 Figure 27.117

Figure 27.108 Antenna of Cladotanytarsus (Lenziella) sp. Figure 27.109 Antenna of Cladotanytarsus sp. Figure 27.110 Antenna of Paratanytarsus sp. Figure 27.111 Antenna of Paratanytarsus sp. Figure 27.112 Antenna of Rheotanytarsus sp. Figure 27.113 Antenna of Tanytarsus sp. Figure 27.114 Antenna of Micropsectra sp. Figure 27.115 Mandible of Corynocera sp.(redrawn with modification from Hirvenoja [1961]). Figure 27.116 Spur on apex of antenna! tubercle of Micropsectra sp. or Tanytarsus sp.

Figure 27.118

Figure 27.120

Figure 27.117 Spur on apex of antennal tubercle of Micropsectra sp. Figure 27.118 Spur on apex of antennal tubercle of Micropsectra sp. or Tanytarsus sp. Figure 27.119 Apex of antenna of Tanytarsus sp.

Figure 27.120 Apex of antenna of Tanytarsus (redrawn from Roback [1966]). Figure 27.121 Second antennal segment of Rheotanytarsus sp. Figure 27.122 Second antennal segment of Subiettea sp.

Chapter 27 Chironomidae

1145

19(10').

Premandible with 3^ slender, pointed teeth (Fig. 27.127)

22

19.

Premandible with only 2 broader teeth (Fig. 27.126)

20

20(19').

Eighth abdominal segment with well-developed dorsal projection (Fig. 27.129)

Micropsectm (in part)

20'.

Eighth abdominal segment lacking dorsal projection or with only small round bulge [only one species, M. roseiventris (Kieffer), has this bulge but it is not known from North America] Micropsectm Kieffer (in part)

21(4).

Antennal tubercle with a conspicuous dorsally projecting flap that covers base of the first antennal segment and large anteromesally projecting palmate process that nearly reaches midline of head capsule (Fig. 27.131) Neostempellina Reiss

21'.

Antennal tubercle without a conspicuous dorsally projecting flap that covers base of the first antennal segment; anteromesally projecting palmate process less well produced and clearly ending well before midline of head capsule (Fig. 27.105) Thienemann and Bause

22(19).

All claws of posterior prolegs curved and tapering to a single point distally and inner margin of curved area smooth Tanytarsus van der Wulp (in part)

22'.

One or more smaller claws of posterior prolegs not curved to a tapered point distally, and margin with numerous small teeth that usually form two or more partially overlapping rows(Fig. 27.130)

Virgatanytarsus Finder

Chironomini

1. 1'.

Seven anterior abdominal segments subdivided, giving the appearance of a 20-segmented body; larva oligochaete-like

Chernovskiia Saether

Seven anterior abdominal segments not subdivided, not giving the appearance of a 20-segmented body

2(1').

2'.

3(2).

Antenna with 6 segments, lauterborn organs large and alternate at apices of segments 2 and 3 (Figs. 27.140, 27.141)

Mentum with paired median teeth

4(3').

Median teeth of mentum distinctly lighter in color than outer lateral teeth (Figs. 27.132,27.134, 27.187) Median teeth of mentum not less darkly pigmented than outer lateral teeth (Figs. 27.133,27.135, 27.137,27.138)

5(4).

^

6(4').

8 46 4 5

6

Two median teeth of mentum distinctly lighter in color than outer lateral

teeth(Figs. 27.134, 27.187) 5'.

3

Antenna usually with 5, 7, or 8 segments,(Figs. 27.147-27.149) or if 6 antennal segments are present then lauterborn organs not alternate at apices of 2nd and 3rd segments Mentum with a single broad (Fig. 27.136) or narrow (Fig. 27.193) light colored tooth and 5 or 6 pairs of more darkly pigmented lateral teeth

3'.

4'.

2

Four median teeth of mentum distinctly lighter in color than outer lateral teeth (Fig. 27.132) or light median teeth tiered

Antenna elongate and arising from distinct tubercle (Fig. 27.141); anterior edge of ventromental plates straight; mentum as in Fig. 27.137 or Fig. 27.188

47 Paratendipes Kieffer 49

6'.

Not with the above combination of characters

7(6').

Median teeth of mentum deeply separated and strongly recessed within first lateral teeth; ventromental plates produced laterally, more than 4 times wider than long (Fig. 27.211) Fissimentum Cranston and Nolte

7

4^

i

Figure 27.124

Figure 27.125

Figure 27.127

Figure 27.126

Figure 27.123

Figure 27.128

Figure 27.130

Figure 27.131

Figure 27.129

Figure 27.123 Antenna of Pontomyia sp. Figure 27.124 Pecten epipharyngis of Pontomyia sp. Figure 27.125 Pecten epipfiaryngis of Paratanytarsus sp. Figure 27.126 Premandible of Micropsectra sp. Figure 27.127 Premandible of Tanytarsus sp. Figure 27.128 Mentum and ventromental plates of Micropsectra sp.

1146

Figure 27.129 Lateral view of posterior abdominal region of Micropsectra sp. Figure 27.130 Ciaws of posterior proleg of Virgatanytarsus sp. Figure 27.131 Antennal tubercle and palmate process of Neostempellina sp.

Chapter 27 Chironomidae

T. 8(2').

Median teeth of mentum not deeply separated (Figs. 27.133, 27.135); ventromental plates less than 4 times wider than long First antennal segment curved and lateral labral sclerite with anterior margin appearing serrated (Fig. 27.139), mentum as in Fig. 27.138

57

Pagastiella Brundin

8'.

Not with the above combination of characters

9(8').

Mentum with a dome-shaped median tooth, median tooth pale yellow in middle, lateral edges and remaining lateral teeth darkly pigmented; lateral teeth longer than the median tooth, giving the mentum an overall concave appearance (Fig. 27.145) Mentum without a dome-shaped median tooth (Figs. 27.152, 27.171, 27.186), or, if median tooth is dome shaped, then lateral teeth small and not giving the impression of an overall concave mentum (Figs. 27.150, 27.154,

9'.

9

10

27.159,27.160,27.167)

10(9).

Mentum with 5 lateral teeth (Fig. 27.145)

1147

13

Cryptochironomus Kieffer (in part)

10'.

Mentum with 7 lateral teeth

11(IO').

Antenna with 7 segments; blade originating at apex of 3rd segment (Fig. 27.148) Demicryptochironomus henz Antenna with 5 segments; blade originating near apex of 2nd segment(Fig. 27.147) 12 Mentum with 7 sharp and free lateral teeth; S II blade-like,

11'. 12(11').

11

all other S setae reduced

12'.

13(9').

13'.

Gillotia Kieffer

Mentum with 7 lateral teeth, however, 1st lateral teeth incompletely separated from the median tooth (Fig. 27.146), and the outermost 2 lateral teeth fused throughout most of their base; median tooth often with 2 small spines in the center (Fig. 27.146) Cryptochironomus Kieffer (in part) Two outermost lateral teeth of mentum distinctly enlarged as in Figs. 27.151-27.153, and labral sensilla 2-segmented (Fig. 27.143); mentum similar to Figs. 27.151-27.153 14 Two outermost lateral teeth of mentum usually not enlarged, if somewhat enlarged then labral sensilla 3-segmented (Fig. 27.142); mentum not as in Figs. 27.151-27.153

14(13). 14'.

Median tooth of mentum trifid (Fig. 27.153) and antennal blade longer than the combined lengths of segments 2 through 5 Median tooth of mentum broadly rounded, medially notched and often giving the appearance of paired teeth (Fig. 27.152), or with lateral notches (Fig. 27.151); antennal blade shorter than flagellum

15(14').

Median tooth of mentum medially notched (Fig. 27.152)

15'.

Median tooth of mentum broadly rounded or with lateral notches (Fig. 27.151)

16(13').

16

Microchironomus Kieffer

15 Cladopelma Kieffer

Cryptotendipes Beck and Beck Median portion of mentum lacking distinct teeth (Figs. 27.157 and 27.159), with a large dome-shaped tooth (Figs. 27.154 and 27.160) or with 1 to several notches so that the median portion forms a broad convex structure (Figs. 27.155, 27.156, 27.158, 27.161) 17

16'.

Median portion of mentum with distinct teeth (Figs. 27.162, 27.165, 27.173, 27.182)

17(16).

Antenna more than one-third as long as head capsule; mentum and

17'.

ventromental plates as in Figs. 27.156-27.160 Paracladopelma Harnish (in part) Antenna not more than one-third as long as head capsule; mentum and ventromental plates variable

23

18

1148

Chapter 27 Chironomidae

Figure 27.132

Figure 27.133

Figure 27.

#

Figure Figure 27.137

Figure 27.138

Figure 27.139

Figure 27.132 Figure 27.133 Figure 27.134 Figure 27.135

Mentum of Paratendipes sp. Mentum of Stictochironomus sp. Mentum of Microtendipes sp. Mentum of Omisus sp.(redrawn with

modification from Beck and Beck [1970]). Figure 27.136 Mentum of Paralauterborniella sp.

Figure 27.137 Mentum of Lauterborniella sp.

Figure 27.140

Figure 27.141

Figure 27.138 Mentum of Pagastiella sp. Figure 27.139 Antenna and lateral iabral sclerite of Pagastiella sp.

Figure 27.140 Antenna of Microtendipes sp. Figure 27.141 Antenna of Lauterborniella sp.

Chapter 27 Chironomidae

18(17'). 18'.

Antenna! blade longer than flagellum; median tooth of mentum pointed or subtriangular (Fig. 27.161); anal tubules vestigial

Acalcarella Shilova

Antenna! blade shorter than flagellum; median tooth of mentum not pointed or subtriangular

19(18').

1149

19

Mandible with an elongate apical tooth and 4 smaller inner teeth that originate from a common base (Fig. 27.144); mentum as in Fig. 27.150 Nilothauma Kieffer

19'.

Mandible not as in Fig. 27.144 and mentum not as Fig. 27.150

20(19').

Second antenna! segment about as long as 3rd segment and antenna 5-segmented; basal segment of maxillary palp about 4 times as long as wide Harnischia Kieffer

20'.

Second antenna! segment distinctly longer than 3rd segment or antenna 6-segmented and 2nd segment much shorter than 3rd segment; basal segment of maxillary palp at most 3 times as long as broad

21

Antenna 6-segmented; mentum as in Figs. 27.154, 27.155, or 27.203

58

21(20').

20

21'.

Antenna 5-segmented; mentum net as in Figs. 27.154, 27.155, or 27.203

22(21').

Second antenna! segment unsclerotized in basal two-thirds(Fig. 27.149)

22'.

Second antenna! segment fully sclerotized; mentum as in Figs. 27.156-27.160

22 Cyphomella Saether

Paracladopelma Harnish (in part)

23(16').

Mentum with an even number of teeth

24

23'.

Mentum with an odd number of teeth

33

24(23).

Mentum concave and consisting of 8 or 10 darkly pigmented teeth (Figs. 27.165, 27.189); mandible short and robust(Fig. 27.170)

48

24'.

Mentum and mandibles not as in Figs. 27.165, 27.170, 27.189

25

25(24').

Pecten epipharyngis composed of 3 blunt teeth; mentum as in Fig. 27.173; larvae occurring in stems and petioles of aquatic vascular plants

25'.

Hyporhygma Reiss

Pecten epipharyngis either a single individual plate or, more commonly, consisting of 1-3 plates, each with 2 or more teeth; mentum not as in Fig. 27.173

26

26(25').

Apical segment of maxillary palp elongate and well sclerotized (Fig. 27.168); striations of ventromental plates very coarse; mentum as in Fig. 27.162 or 27.163

26'.

Apical segment of maxillary palp not elongate, rarely well sclerotized; ventromental

plates with finer striations and mentum not as in Fig. 27.162 or 27.163

Robackia Saether

27

27(26').

Median pair of mental teeth partially fused and distinctly wider than each of the remaining lateral teeth (Fig. 27.166); anterior margin of ventromental plates coarsely scalloped (Fig. 27.166) Pamchironomus Lenz(in part)

IT.

Median pair of mental teeth not partially fused, or if partially fused then not distinctly wider than each of the remaining lateral teeth; anterior margin of ventromental plates not coarsely scalloped

28(27').

First lateral teeth of mentum much shorter than median and 2nd lateral teeth (Fig. 27.171)

28'.

First lateral teeth of mentum subequal to or longer than the median and/or

Polypedilum Kieffer (in part)

2nd lateral teeth

29(28'). 29'.

Lateral teeth of mentum gradually decreasing in size and length so that anterior margin of mentum appears to be broadly convex (Figs. 27.172, 27.174)

29

32

Second lateral teeth recessed and smaller in size than 1st lateral and 3rd lateral

teeth (Fig. 27.176), or median teeth partially fused (Fig. 27.164) 30(29').

28

Median teeth partially fused (Fig. 27.164); ventromental plates 3-A times wider than their length

30 53

1150

Chapter 27 Chironomidae

3-segmented labral sensilla

2-segmented labral sensilla

M Figure 27.143

Figure 27.144

Figure Figure 27.142

Figure 27.146

Figure 27.150

Figure 27.149 Figure 27.148 Figure 27.147 Figure 27.151

Figure 27.152

Figure 27.142 Anterior view of labral region of Paracladopelma sp. showing position of three-segmented labral sensillae in relation to other labral armature.

Figure 27.143 Anterior view of labral region of Cladopelma sp. showing two-segmented labral sensillae.

Figure 27.144 Mandible of Nilothauma sp. Figure 27.145 Mentum of Cryptochironomus sp. Figure 27.146 Detail of median portion of mentum of Cryptochironomus blarina Townes.

Figure 27.153

Figure 27.147 Antenna of Cryptochironomus sp. Figure 27.148 Antenna of Demicryptochironomus sp. Figure 27.149 Antenna of Cyphomeiia sp.(redrawn with modification from Saether [1977]). Figure 27.150 Mentum of Niiothauma sp. Figure 27.151 Mentum of Cryptotendipes sp. Figure 27.152 Mentum of Cladopelma sp. Figure 27.153 Mentum of Microchironomus sp. (redrawn with modification from Kugler [1971]).

Chapter 27 Chironomidae

Figure 27.154

Figure 27.155

Figure 27.156

Figure 27.157

Figure 27.158

Figure 27.159

Figure

1151

Figure 27.161

Figure 27.162

\m Figure 27.164

Figure 27.154 Mentum of Saetheria tylus (Townes). Figure 27.155 Mentum of Saetheria sp.(redrawn from Jackson [1977]). Figure 27.156 Mentum of Paracladopelma galaptera (Townes)(redrawn from Jackson [1977]). Figure 27.157 Mentum of Paracladopelma rolll (Kirp.) (redrawn with modification from Saether [1977]). Figure 27.158 Mentum of Paracladopelma undine (Townes)(redrawn from Jackson [1977]). Figure 27.159 Mentum of Paracladopelma dorls (Townes)(redrawn with modification from Saether [1977]).

Figure 27.165

Figure 27.160 Mentum of Paracladopelma longanae Beck and Beck (redrawn from Jackson [1977]). Figure 27.161 Mentum of Acalcarella sp. Figure 27.162 Mentum of Robackia clavlger (Townes). Figure 27.163 Mentum of Robackia demeljerel (Kruseman). Figure 27.164 Mentum of Endochlronomus sp. Figure 27.165 Mentum of Stenochlronomus sp.

1152

30'. 31(30').

Chapter 27 Chironomidae

Median teeth not partially fused (Fig. 27.176); ventromental plates not more than 3 times as wide as their maximum length

31

Mola of mandible without serrations, or with only a simple serration at base of seta subdentalis (Fig. 27.190); distance from basal notch of innermost mandibular tooth to insertion of seta subdentalis usually at least 3/4 the distance from basal notch to apical notch of the apical inner tooth

31'.

54

Mola of mandible with 1-3 serrations distinctly separated from seta subdentalis (Fig. 27.191); distance from basal notch of innermost mandibular tooth to insertion of seta subdentalis less than 3/4 the distance from basal notch to

apical notch of apical inner tooth

32(29). 32'.

Tribelos Townes

Ventromental plates more than 3 times wider than their maximum length and with lateral corners rounded (Fig. 27.174)

Polypedilum (Asheum)

Ventromental plates usually less than 3 times wider than their maximum

length, lateral corners never rounded (Fig. 27.172)

Polypedilum Kieffer (in part)

33(23').

Anterior margin of ventromental plates coarsely scalloped; median tooth of mentum about 2 times as wide as 1 st lateral teeth; all lateral teeth pointed and gradually diminishing in size laterally (Fig. 27.167) Parachironomus Lenz(in part)

33'.

Anterior margin of ventromental plates smooth (Figs. 27.178, 27.185) or only finely crenulate (Figs. 27.182, 27.184); median tooth and lateral teeth of mentum variable, but not as in Fig. 27.167

34

34(33').

Median apices of ventromental plates touching or separated from one another by a distance less than the width of the median tooth (Figs. 27.177, 27.178, 27.180,

34'.

Median apices of ventromental plates separated by a distance equal to or greater than the width of the median tooth (Figs. 27.182, 27.186)

38

35(34).

Ventromental plates touching on midline; mentum as in Fig. 27.181 or 27.192

50

35'.

Ventromental plates not touching on midline; mentum not as in Fig. 27.181 or 27.192

36

36(35').

Mentum strongly arched, and with alternating large and small teeth (Fig. 27.180); larvae found in freshwater sponges

27.181,27.192)

35

Xenochironomus Kieffer

36'.

Mentum not strongly arched, not as in Fig. 27.180; larvae found in various habitats, but apparently more common in subtropical regions of the United States

37(36').

Mentum with distinctly overlapping lateral teeth (Fig. 27.177); abdominal segment 8 with a bifurcate anterior ventral tubule (Fig. 27.179) Goeldichironomus Fittkau (in part)

37'.

Lateral teeth of mentum not overlapping (Fig. 27.178) abdominal segment 8 with a nonbifurcate anterior ventral tubule Goeldichironomus Fittkau (in part)

38(34').

Some lateral teeth of mentum projecting at least as far anterior as median tooth, giving the mentum an overall flat (Fig. 27.175) or concave appearance (Fig. 27.194), or median tooth deeply recessed (Fig. 27.193)

51

38'.

Outermost teeth of mentum decreasing in size or length giving the mentum an overall convex appearance (Figs. 27.183, 27.185, 27.186)

39

39(38').

Median tooth of mentum broadly rounded (Figs. 27.182, 27.185) or pointed (Figs. 27.183, 27.184), but lacking lateral notches

40

39'.

Median tooth with lateral notches that give it a trifid appearance (Fig. 27.186)

42

40(39).

Ventromental plate distinctly less than 2 times as wide as long and usually with small crenulations along anterior margin; median tooth and 1 st lateral teeth enlarged and somewhat pointed; 1 st laterals with (Fig. 27.183) or without lateral notches (Figs. 27.184, 27.195)

40'.

37

55

Ventromental plates at least twice as wide as long and anterior margin with or without crenulations; median tooth rounded; 1st lateral teeth variable but

never as in Figs. 27.183, 27.184 or 27.195

41

Chapter 27 Chironomidae

Figure 27.166

1153

Figure 27.167

Figure 27.171

Figure 27.169

Figure 27.170 Figure 27.172 Figure 27.168

Figure

Figure

Figure 27.175

Figure 27.176

Figure 27.166 Mentum of Parachironomus of. frequens (Johannsen) Figure 27.167 Mentum of Parachironomus of. abortivus (Malloch). Figure 27.168 Maxillary palpus of Robackia demeijerei(Kruseman). Figure 27.169 Maxillary palpus of Beckidia tethys (Townes)(redrawn witfi modification from Saether[1977]).

Figure 27.170 Figure 27.171 Figure 27.172 Figure 27.173 Figure 27.174

Mandibie of Stenochironomus sp. Mentum of Polypedilum sp. Mentum of Polypedilum sp. Mentum of Hyporhygma sp. Mentum of Polypedilum (Asheum)

beckae Sublette.

Figure 27.175 Mentum of Beckidia tethys (Townes) (redrawn with modification from Saether [1977]). Figure 27.176 Mentum of Phaenopsectra sp.

1154

41(40').

Chapter 27 Chironomidae

41'.

Median tooth extending anterior to I st lateral teeth; 4th lateral teeth reduced (Fig. 27.185) Einfeldia Kieffer (in part) Median tooth recessed within 1st lateral teeth (Fig. 27.182), or very broad and extending about as far anterior as 1 st lateral teeth, or if projecting farther anterior than 1 st lateral teeth then 4th lateral teeth not conspicuously reduced Glyptotendipes Kieffer

42(39').

Premandible with 3 or more teeth

42'.

Premandible at most bifid

43(42').

Frontal apotome with an oval depression just posterior to antennal bases and 8th abdominal segment with 1 pair of ventral tubules Einfeldia Kieffer (in part)

Kiefferulm (Kiefferulus) Goetghebuer (in part) 43

43'.

Not with the above combination of characters

44(43').

Eighth abdominal segment without ventral tubules and pecten epipharyngis with 3-7 teeth Einfeldia Kieffer (in part)

44

44'.

Not with the above combination of characters

45

45(44').

Eighth abdominal segment lacking ventral tubules, with 1 pair of ventral tubules, or with 2 pairs of ventral tubules; pecten epipharyngis with 10 or more teeth

56

45'.

Eighth abdominal segment with ventral tubules and pecten epipharyngis with 9 teeth; known only from California Kiefferulus(Wirthiella) Sublette (in part)

46(3).

Median tooth of mentum broad and with 6 pairs of more darkly pigmented lateral teeth (Fig. 27.136) Paralauterborniella Lenz

46'.

Median tooth of mentum narrow, recessed and with 5 pairs of more darkly pigmented lateral teeth (Fig. 27.193) Beardius Reiss and Sublette (in part)

47(5).

SI setae well separated, sockets not fused (Fig. 27.209)

47'.

SI setae originating very close to each other, sockets fused medially (Fig. 27.210)

Microtendipes Kieffer Apedilum

48(24).

Mentum with 10 darkly pigmented teeth (Fig. 27.165)

48'.

Mentum with 8 darkly pigmented teeth (Fig. 27.189)

49(6).

Seta posterior to ventromental plates plumose (Fig. 27.137); abdominal segment eight with posteriorly projecting dorsal hump (Fig. 27.196) Lauterborniella Thienemann and Bause

49'.

Seta posterior to ventromental plates simple (Fig. 27.188); abdominal segment eight with anteriorly projecting dorsal hump (Fig. 27.197)

Zavreliella Kieffer

Inner teeth of mandible sharply pointed (Fig. 27.198); head capsule with well developed tubercles lateral to bases of antennae (Fig. 27.199)

Lipiniella Shilova

50(35).

Stenochironomus Xestochironomus Sublette and Wirth

50'.

Inner teeth of mandible low, blunt(Fig. 27.200); head capsule without tubercles

51(38).

Maxillary palp more than 4 times longer than wide, one apical segment elongate (Fig. 27.169); mentum as in Fig. 27.175

Beckidia Saether

Maxillary palp not more than 3 times longer than wide, apical segment not as in Fig. 27.169; mentum as in Fig. 27.193 or Fig. 27.194

52

lateral to bases of antennae

51'. 52(51'). 52'.

Mentum as in Fig. 27.194 Mentum as in Fig. 27.193

53(30).

Oral margin of cardo tuberculate (Fig. 27.201)

53'. 54(31).

Oral margin of cardo not tuberculate (Fig. 27.202) Mandible with 4 inner teeth

54'.

Mandible with 3 inner teeth

Axarus Roback

Kribiodorum Kieffer Beardius Reiss and Sublette (in part) Endochironomus Kieffer Synendotendipes Grodhaus Kieffer Phaenopsectra Kieffer

Chapter 27 Chironomidae

1155

Figure 27.178

Figure 27.177

P\a/ v>/^/Y\

Figure 27.181 Figure 27.180 bifurcate anterior ventrai tubuie

Figure 27.179 Figure 27.182

Figure 27.183

Figure 27.184

Figure 27.185

Figure 27.186

Figure 27.177 Mentum of Goeldichironomus sp. Figure 27.178 Mentum of Goeldichironomus devineyae (Beck)(redrawn with modification from Beck and Beck [1970]). Figure 27.179 Posterior abdominal segments and ventral tubules of Goeldichironomus sp,(redrawn with modification from Fittkau [1965]). Figure 27.180 Mentum of Xenochironomus xenoiabis (Kieffer).

Figure 27.181 Mentum of Axarus festivus (Say). Figure 27.182 Mentum of Giyptotendipes sp. Figure 27.183 Mentum of Dicrotendipes sp. Figure 27.184 Mentum of Dicrotendipes sp. Figure 27.185 Mentum of Einfeidia natchitocheae (Sublette)(redrawn with modification from Subiette [1964]). Figure 27.186 Mentum of Chironomus sp.

Figure 27.187

Figure 27.188

Figure 27.189

Figure 27.191

Figure 27.190

Figure 27.192

Figure 27.193

Figure 27.195

Figure 27.194 of

Apedilum sp.

Figure 27.188 Mentum and ventromental plates

of

Zavreliella sp.

Figure 27.189 Mentum and ventromental plates Xestochironomus sp.

Figure 27.190 Mandible of Phaenopsectra sp. Figure 27.191 Mandible of Tribelos sp.

1156

of

Figure 27.192 Mentum and ventromental plates of Upiniella sp. Figure 27.193 Mentum and ventromental plates of Beardius sp. Figure 27.194 Mentum and ventromental plates of Kribiodorum sp. Figure 27.195 Mentum and ventromental plates of Endotribelos sp.

-

Chapter 27 Chironomidae

Figure 27.196

1157

Figure 27.197

Figure 27.198

Figure 27.199 Figure 27.200

MlTvijfWM

Figure 27.201

Figure 27.202

Figure 27.196 Lateral view of apical segments of abdomen in Lauterborniella sp. Figure 27.197 Lateral view of apical segments of abdomen in Zavreliella sp. Figure 27.198 Mandible of Lipiniella sp. Figure 27.199 Antenna and lateral tubercle of Lipiniella sp.

rs

Figure 27.200 Mandible of Axarus sp. Figure 27.201 Mentum, ventromental plates, maxilla, maxillary palp and cardo of Endochironomus sp. Figure 27.202 Mentum, ventromental plates, maxilla, maxillary palp and cardo of Synendotendipes sp.

1158

Chapter 27 Chironomidae

Figure 27.203

Figure 27.206

Figure 27.205

Figure 27.204

Figure 27.210

Figure 27.209

Figure 27.207

Figure 27.208

Figure 27.211

Figure 27.203 Mentum and ventromental plates of Kloosia sp.

Figure 27.204 Mandible of Endotribelos sp. Figure 27.205 Pecten epipharyngis of Endotribelos sp.

Figure 27.206 Pecten epipharyngis of Dicrotendipes sp. Figure 27.207 Pecten epipharyngis of Einfeldia sp.

Figure 27.208 Figure 27.209 Figure 27.210 Figure 27.211

Pecten epipharyngis of Chironomus sp. SI setae of Microtendipes sp. SI seta of Apedilum sp. Mentum and ventromental plates of

Fissimentum sp.

Chapter 27 Chironomidae

55(40). 55'. 56(45).

1159

Pecten epipharyngis consisting of 3 plates each with 4-5 apical teeth (Fig. 27.205) and mandible and mentum as in Figs. 27.204, 27.195 Endotribelos Grodhaus Pecten epipharyngis consisting of one plate with 3-6 strong or blunt teeth (Fig. 27.206) Dicrotendipes^ Kieffer Pecten epipharyngis with numerous fine teeth in 2-3 partial or complete rows (Fig. 27.207) Einfeldia Kieffer (in part)

56'.

Pecten epipharyngis with stronger teeth in only one row

57(7').

(Fig. 27.208) Chironomus Meigen (in part) First lateral teeth closely appressed to 2nd lateral teeth (Fig. 27.135); antenna distinctly longer than mandible

57'.

Omisus Townes

First lateral teeth not closely appressed to 2nd lateral teeth, much longer than median or 2nd lateral teeth (Fig. 27.133); antenna subequal in length to mandible

Stictochiwnomus Kieffer

58(21).

Mentum as in Fig. 27.154 or 27.155

Saetheria Jackson

58'.

Mentum as in Fig. 27.203

Kloosia Kruseman

Orthodadiinae

1. r. 2(1).

Antenna elongate, more than two-thirds the length of the head capsule Antenna not elongate, equal to or less than two-thirds the length of the head capsule Second antennal segment with extensive dark brown or golden brown pigmentation (Figs. 27.226, 27.227)

2'.

Second antennal segment concolorous with remaining segments, lacking

3(2).

distinct dark brown or golden brown pigmentation Antenna longer than head capsule, 4-segmented, and usually with some

darker pigmentation occurring on the 3rd segment(Fig. 27.226) 3'.

Antenna shorter than head capsule, 5-segmented, and never with dark pigmentation occurring on 3rd segment(Fig. 27.227)

4(2').

Median portion of mentum concave (Fig. 27.255); antenna with large

2 6 3

4

Corynoneum Winnertz Thienemanniella Kieffer

lauterborn organs originating at different levels on 2nd antennal

4'. 5(4').

5'.

segment(Fig. 27.233) Heterotanytarsus Sparck Median portion of mentum not concave; antenna not as in Fig. 27.233 5 Procercus well-developed; 1 procercal seta at lest twice as long as the length of the posterior prolegs; antenna with an elongate blade and whip-like apical segment(Fig. 27.228); body white when preserved Lopesdadius Oliveira Procercus less well-developed; all procercal setae less than twice as long as posterior prolegs; apical segment of antenna not whip-like(Fig. 27.229); body often with distinctive purple or brown pigmented areas when preserved

Rheosmittia Brundin

6(1').

Antenna arising from tuburcle that has a large anteromedially directed spur (Fig.27.230); thoracic segments with conspicuous setae, many of which are apically dissected; larvae that build portable sand cases Abiskomyia Edwards

6'.

Not with the above combination of characters

7(6').

One procercal setae elongate, at least one-quarter the length of the larval body (Fig. 27.298); remaining procercal setae vestigial or absent All procercal setae less than one-fifth the length of the larval body

7'.

7

8 10

' Some species of Dicrotendipes have a mandible similar to Fig. 27,204, but the mentum of these species is similar to Figs, 27,183 or 27,184,

1160

8(7).

Chapter 27 Chironomidae

Mentum consisting of a single broad, dome shaped median tooth with a central cusp and 6 pairs of narrow pointed lateral teeth (Fig. 27.249); apical tooth of mandible longer than the combined widths of the

8'.

lateral teeth (Fig. 27.310) Krenosmittia Thienemann and Kruger Mentum consisting of paired median teeth and 5 paired lateral teeth, the outermost 2 teeth fused throughout most of their length (Fig. 27.250); apical tooth of mandible shorter than combined widths of the lateral teeth (Figs. 27.305, 27.306)

9(8').

9

Mandible with 3 inner teeth (Fig. 27.306)

Pseudorthocladius Goetghebuer

9'.

Mandible with 2 inner teeth (Fig. 27.305)

Pamchaetocladius Wiilker

10(7').

Ventromental plates well-developed and covering all lateral teeth of mentum; 3 small median teeth visible in median concavity of mentum (Fig. 27.251) or median area of mentum lacking teeth (Figs. 27.224 or 27.252); larvae either

15'.

occurring only within colonies of blue-green algae, ectoparasitic on immature mayflies, or occurring within unionid molluscs 70 Mentum not as in Figs. 27.224, 27.251, or 27.252; larvae living in a variety of habitats 11 Cardinal beard present beneath the ventromental plates (Figs. 27.285-27.296, 27.313, 27.314) ....12 Cardinal beard not present beneath the ventromental plates (Figs. 27.252-27.284) 21 SI seta simple (Fig. 27.213) bifid (Fig. 27.214) or bifid with secondary feathering(Fig. 27.215)....13 SI seta palmate (Fig. 27.216), plumose (Fig. 27.217), or with multiple apical dissections (Figs. 27.218, 27.222, 27.223) 18 SI setae simple (Fig. 27.213) 14 SI setae bifid or bifid with secondary feathering (Figs. 27.214, 27.215) 15 Cardinal setae very weak and mostly covered by lateral edges of ventromental plates (Fig. 27.313) or occurring in a single row (Fig. 27.314) 62 Cardinal setae elongate, bristle-like (Fig. 27.295) branched or stellate (Fig. 27.292), and extending well beyond lateral edges of ventromental plates 63 SI setae bifid with secondary feathering; mentum as in Fig. 27.287 Acricotopus Kieffer SI setae bifid; mentum not as in Fig. 27.287 16

16(15').

Mentum with paired median teeth (Fig. 27.291)

16'.

Mentum with an unpaired median tooth (Figs. 27.285, 27.289)

17(16').

Median tooth of mentum broad, dome-shaped, and distinctly lighter in color than remaining lateral teeth (Fig. 27.289)

10'. 11(IO'). 11'. 12(11). 12'.

13(12). 13'.

14(13). 14'.

15(13').

17'.

18(12').

18'.

19(18').

Rheocricotopus Brundin 17

Pamcladius Hirvenoja

Median tooth of mentum narrower, not dome-shaped, and usually

concolorous with 1st lateral teeth (Fig. 27.285) Halocladius(Halocladius) Hirvenoja SI seta palmate (Fig. 27.216); procerci with at least 1 basal spur; apical tooth of mandible longer than the combined widths of the lateral teeth (Fig. 27.307); mentum usually with a broad unpaired but peaked median tooth (Fig. 27.290) or with broad paired median teeth (Fig. 27.288) Psectrocladius Kieffer SI seta plumose (Fig. 27.217) or with multiple apical dissections (Fig. 27.218), but not with the remaining combination of characters 19 Mentum with a broad, truncated median tooth (Fig. 27.286); larvae occurring in marine littoral zones, not presently recorded from North America Halocladius(Psammociadius) Hirvenoja

19'.

Mentum with rounded or pointed median teeth (Figs. 27.293, 27.294, 27.296);

larvae occurring in a variety of freshwater habitats

20(19').

Mandible with 4 inner teeth; mentum as in Fig. 27.293

20

Diplocladius Kieffer

Chapter 27 Chironomidae

20'. 21(11').

Mandible with 3 inner teeth; mentum either with 2-4 pale median teeth (Fig. 27.296) or similar to Fig. 27.294 Procerci present

1161

Zalutschia Lipina 27

21'.

Procerci absent

22

22(21').

Preanal segment strongly bent ventrally so that the anal segment and posterior prolegs are orientated at a right angle to the long axis of the body (Figs. 27.301, 27.302); anal segment and posterior prolegs retractile into preanal segment

23

22'.

Preanal segment at most declivent; anal segment and posterior prolegs not oriented at right angle to main axis of body, and not retractile into preanal segment 24 Posterior prolegs each subdivided, the anterior portion bearing a semicircle of claws (Fig. 27.301); no anal tubules present; antenna as in Fig. 27.236 Gymnometriocnemus Edwards

23(22).

23'.

Posterior prolegs not subdivided (Fig. 27.302); 4 short anal tubules present; antenna similar to Fig. 27.237 Bryophaenocladius Thienemann

24(22').

Larva ectoparasitic on mayflies; mouthparts reduced, mentum as in Fig. 27.252; body robust, especially thoracic segments, giving an overall swollen appearance (Fig. 27.303) Symbiocladius Kieffer (in part)

24'.

Larva not ectoparasitic; mouthparts, mentum, and body not as in Fig. 27.252 or 27.303

25

25(24').

SI seta and SII seta bifid and both well-developed (Fig. 27.221)

74

25'.

SII seta never bifid

26

26(25').

SI seta strong and simple (Fig. 27.219) or with fine apical and lateral serrations (Fig. 27.220); antenna as in Fig. 27.232 or Fig. 27.317; anal tubules elongate and with numerous constrictions throughout their length (Figs. 27.300, 27.321); mentum as in Figs. 27.253, 27.316 or 27.320

65

26'.

27(21).

SI seta not strong and simple or with fine apical and lateral serrations, but usually with multiple apical dissections (Fig. 27.222) or nearly palmate; antenna similar to Fig. 27.238, 27.239 or 27.318; anal tubules not elongate, lacking constrictions; mentum not as in Fig. 27.253 67 Antenna 7-segmented, 3rd segment shorter than 4th segment and 7th segment hair-like or vestigial (Fig. 27.234); mentum with 1 (maeaeri group. Fig. 27.261) or 2 median teeth and 5 pairs of lateral teeth (Fig. 27.263); labral lamella present and divided at least anteriorly (Fig. 27.248); SI seta plumose Hetewtrissocladius Sparck

IT.

Not with the above combination of characters

28

28(27').

Larva with a dense lateral fringe of setae on at least abdominal segments 1 through 5 (Fig. 27.297); mentum as in Fig. 27.259 or 27.315

64

28'.

Larva lacking a dense fringe of setae on abdominal segments; mentum

29(28').

Mentum strongly arched and appearing truncated at apex, only 6 anterior teeth readily discernible when head capsule not strongly depressed (Figs. 27.256, 27.257); larvae phoretic on Ephemeroptera nymphs or living within the gills of bivalve mollusks

30

Mentum, if strongly arched, not as in Figs. 27.256 and 27.257; larvae living in a variety of habitats

31

not as in Fig. 27.259 or 27.315

29'.

30(29). 30'.

29

Body with conspicuous brown setae; mentum as in Fig. 27.257; antenna as in Fig. 27.240; premandible with 2 apical teeth; larvae phoretic on Ephemeroptera ..Epoicocladius Sulc & Zavfel Body lacking conspicuous brown setae; mentum as in Fig. 27.256; antenna as in Fig. 27.231; premandible with 6 apical teeth; larvae living within gills of bivalve mollusks

Baeoctenus Ssther

1162

Chapter 27 Chironomidae

Figure 27.212

Figure 27.216

Figure 27.217

Figure 27.218

Figure 27.213 Figure 27.214 Figure 27.215

Figure 27.221

Figure 27.222

Figure 27.219

Figure 27.223

Figure 27.220

Figure 27.224 Figure 27.225

Figure 27.212 Figure 27.213 Figure 27.214 Figure 27.215

Head capsule of Tempisquitoneura sp. Simple SI seta of Eukiefferiella sp. Bifid SI seta of Cricotopus sp. Bifid Si seta with secondary feathering

in Acricotopus sp.

Figure 27.216 Palmate Si seta of Psectrocladius sp. Figure 27.217 Plumose Si seta of Parametriocnemus sp. Figure 27.218 Si seta of Hydrobaenus sp. with strong apical dissections. Figure 27.219 Strong and simple Si seta of Georthocladius sp.

Figure 27.220 Si seta with fine apicai dissections. Figure 27.221 Bifid Si and Sii setai arrangement in Pseudosmittia sp. Figure 27.222 Si seta with strong apicai dissections in Smittia sp. Figure 27.223 Si seta with weak apicai dissections in Paracricotopus sp.

Figure 27.224 Mentum of Trichochilus sp. Figure 27.225 Mentum of Platysmittia sp.

Chapter 27 Chironotnidae

1163

I

Figure 27.230

u Figure 27.229

Figure 27.227

Figure 27.228

Figure 27.226

LJ Figure 27.231

Figure 27.232

Figure 27.233

Figure 27.226 Antenna of Corynoneura sp. Figure 27.227 Antenna of Thienemanniella sp. Figure 27.228 Antenna of Lopescladius sp. Figure 27.229 Antenna of Rheosmittia sp. Figure 27.230 Antenna of Abiskomyia sp.(redrawn witfi modification from Pankratova [1970]). Figure 27.231 Antenna of Baeoctenus bicotor Saether (redrawn with modification from Saether [1977]).

Figure 27.234

Figure 27.235

Figure 27.232 Antenna of Georthocladius sp. Figure 27.233 Antenna of Heterotanytarsus perennis Saether (redrawn with modification from Saether [1975]). Figure 27.234 Antenna of Heterotrissocladius sp. Figure 27.235 Antenna of Heleniella thienemanni Gowin.

1164

Chapter 27 Chironomidae

n

jV/

0

Figure 27.238

—^

Figure 27.237

^

^

Figure 27.239 Figure 27.240

Figure 27.236

Figure 27.243

o

Figure 27.244 Figure 27.242 Figure 27.247

Figure 27.245

Figure 27.241 Figure 27.248 Figure 27.246

Figure 27.236 Antenna of Gymnometriocnemus sp. Figure 27.227 Antenna of Bryophaenocladius sp. Figure 27.238 Antenna of Smittia sp. Figure 27.239 Antenna of Smittia aquatilis Goetghebuer (redrawn with modification from Thienemann and Strenzke [1941]). Figure 27.240 Antenna of Epoicocladius sp. Figure 27.241 Antenna of Brillia sp. Figure 27.242 Antenna of Parametriocnemus sp. Figure 27.243 Pecten epipharyngis and associated labral setae and other armature of Cricotopus (Cricotopus) sp.

Figure 27.244 Antenna of Paraphaenocladius sp. Figure 27.245 Antenna of Psiiometriocnemus sp. Figure 27.246 SI and labral lamellae of Briliia sp. Figure 27.247 Pecten epipharyngis of Cricotopus (Isocladius) sp. Figure 27.248 81 and labral lamellae of Heterotrissocladius sp.

Chapter 27 Chironomidae

31(29').

31'. 32(31').

Second antennal segment elongate, more than two-thirds the length of the 1st segment; antennal segments 3-5 very reduced, their combined lengths equal to about one-fifth the length of segment 2; antennal blade curved and extending well beyond 5th antennal segment(Fig. 27.235); mentum with 2 broad median teeth and 5 smaller lateral teeth; outermost lateral tooth more elongate than preceding lateral tooth (Fig. 27.254) Heleniella Gowin Antenna and mentum not as in Figs. 27.235 and 27.254 32 Antenna 6-segmented; mentum broadly convex, similar to Fig. 27.264, 27.265, or 27.266; SI plumose or with multiple strong or weak apical dissections 33

32'.

If mentum similar to Fig. 27.264, 27.265, or 27.266, then antenna 4- or 5-segmented

33(32).

Median tooth with distinct median cusp (Fig. 27.265)

33'.

Median tooth without medium cusp

34(33').

Median tooth truncated (Fig. 27.266); SI plumose

34'.

Mentum with paired median teeth (Fig. 27.264); SI with multiple apical dissections SI plumose, labral lamella well-developed (Fig. 27.246); 1st antennal segment slightly or strongly bent(Fig. 27.241, 27.317); mentum similar to Fig. 27.258 or 27.260

35(32').

1165

35

Pamkiefferiella Thienemann 34 Oliveridia Saether

Hydrobaenus Fries

61

35'.

Not with the above combination of characters

36(35').

Mentum with strongly arcuate median tooth and 2 pairs of small lateral teeth (Fig. 27.262); larvae living in submerged decomposing wood Orthocladius(Symposiocladius)

36'.

Mentum not as in Fig. 27.262; larvae occurring in a variety of habitats

37

37(36').

Apical tooth of mandible clearly longer than the combined widths of the lateral teeth (Fig. 27.308) Apical tooth of mandible not longer than the combined widths of the lateral teeth

38 39

37'. 38(37).

36

39(37').

Mentum consisting of a broad median tooth with a single median cusp, and 10 pairs of sharp pointed, distinct, lateral teeth (Fig. 27.278); larvae apparently restricted to high alpine or arctic regions Lapposmittia Thienemann Mentum consisting of a broad median tooth with 2 median cusps, and 6 or fewer pairs of lateral teeth, some of which are often small or indistinct; ventromental plates well-developed, usually extending beyond posterolateral corner of mentum (Fig. 27.277) Nanocladius Kieffer SI seta simple 40

39'.

SI seta bifid, plumose, palmate, or with weak (Fig. 27.223) or strong apical

38.

(Fig. 27.222) dissections

42

40(39).

SI seta thick and heavily sclerotized; median tooth of mentum broad, heavily sclerotized (Fig. 27.267); procercus reduced; 2 procercal setae distinctly larger in diameter and longer in length than the remaining setae Cardiodadiiis Kieffer

40'.

If SI seta thick and heavily sclerotized then median tooth of mentum not broad and heavily sclerotized; procerus not reduced; all procercal setae similar in diameter and length

41(40').

41

Antenna reduced, less than one-half the length of the mandible, or antenna about the length of the mandible and larvae occurring in the leaves of pitcher plants Antenna not reduced, subequal to or longer than the length of the mandible; larvae occurring in a wide variety of habitats, never restricted to the leaves of pitcher plants

69

42(39').

SI seta bifid

43

42'.

SI seta plumose, palmate, or with weak or strong apical dissections

52

41'.

51

1166

Chapter 27 Chironomidae

Figure 27.249

Figure 27.250

Figure 27.252

Figure 27.253

Figure 27.255

Figure

Figure 27.251

Figure 27.254

Figure 27.257

Figure 27.258

Figure 27.249 Figure 27.250 Figure 27.251 Figure 27.252 Figure 27.253 Figure 27.254

Figure 27.259

Mentum of Krenosmittia sp. Mentum of Parachaetocladius sp. Mentum of Acamptocladius sp. Mentum of Symbiocladius sp. Mentum of Georthocladius sp. Mentum of Heleniella thienemanni

Gowin.

Figure 27.255 Mentum of Heterotanytarsus perennis Saether(redrawn with modification from Ssether [1975]).

Figure 27.260

Figure 27.256 Mentum of Baeoctenus bicolor Saether (redrawn with modification from Saether [1977]). Figure 27.257 Mentum of Epoicocladius sp. Figure 27.258 Mentum of Brillia sp. Figure 27.259 Mentum of Xylotopus sp. Figure 27.260 Mentum of Brillia sp.

Chapter 27 Chironomidae

Figure 27.261

Figure 27.262

Figure 27.264

Figure 27.265

Figure 27.266

Figure 27.267

Figure 27.268

Figure 27.269

Figure 27.271

Figure 27.272

1167

Figure 27.443

Figure 27.442 Figure 27.446

frontal warts -

M / tt'r .

Ut

fft'ctU u iili—'IiJi

-X

Figure 27.445 J

^

Figure 27.444

Figure 27.450

V .# \

m Figure 27.448 Figure 27.447

H Figure 27.452

Figure 27.449

Figure 27.451

Figure 27.437 Thoracic horn of Zaiutschia sp. Figure 27.438 Tergum 5 of Paracricotopus sp. Figure 27.439 Anterior thorax and thoracic horn of Nanocladius sp. 1. Figure 27.440 Thoracic horn of Nanocladius sp. 2. Figure 27.441 Thoracic horn of Nanocladius sp. 3. Figure 27.442 Thoracic horn of Nanocladius sp. 4. Figure 27.443 Segment 2 of Nanocladius sp. 1. Figure 27.444 Segment 5 of Nanocladius sp. 1. Figure 27.445 Anterior thorax and thoracic horn of Rheocrlcotopus sp. Figure 27.446 Segment 2 of Rheocrlcotopus sp.

Figure 27.447 Segment 5 of Rheocrlcotopus sp. Figure 27.448 Segment 5 of Psectrocladius (Monopsectrocladius) sp. Figure 27.449 Segment 5 of Psectrocladius (Psectrocladius) sp. 1. Figure 27.450 Frontal apotome with frontal setae and frontal warts of Psectrocladius (Psectrocladius) sp. 1. Figure 27.451 Anal lobes and segment 8 of Psectrocladius (Psectrocladius) sp. 1. Figure 27.452 Segment 8 (ventral view) of male Brillia sp. 1.

1196

36'.

Chapter 27 Chironomidae

Posterior margins of terga 3-6(7) with rows of spines or swollen areas with spines (Fig. 27.435, 27.444, 27.447); frontal apotome with or without warts; embedded spines sometimes present on terga (Fig. 27.436)

37

37(36').

Posterior margins of terga 3-6 with swollen areas that have groups of triangular spines (Fig. 27.435); thoracic horn weakly serrate (Fig. 27.437); terga often with embedded spines (Fig. 27.436); frontal apotome with or without warts Zalutschia (in part)

37'.

Posterior margins of terga 3-6(7) with distinct rows of spines (Figs. 27.444, 27.447); thoracic horn with or without serrations; frontal lobe with or without warts; no

embedded spines on caudolateral angles of terga 6-8 38(37').

38

Anal lobe fringe with a few, very broad setae (Fig. 27.462); wing sheaths with pearl rows

Platysmittia

38'.

Anal lobe fringe setae more numerous or not so broad; wing sheaths without pearl rows

39

39(38').

Frontal setae on prefrons, similar to Fig. 27.609

Rheocricotopus(Psilocricotopus) 40

39'.

Frontal setae on apotome, similar to Fig. 27.610

40(39').

Posterior margins of tergites 3-8 with transverse rows of spines, these are relatively short on tergites 3-5 but much longer on tergites 6-8 (Fig. 27.744) Psectrocladius (Mesopsectrocladius)

40'.

Posterior margins of tergites 3-7 with small spines, 8 without

Nanocladius (Plecopteracoluthus)(in part)

41(7').

Anal lobes with no true macrosetae (Figs. 27.457, 27.467)

42

41'.

Anal lobes with at least one macroseta (Figs. 27.453, 27.465, 27.466, 27.469, 27.474)

44

42(41). 42'.

Thoracic horn short and conical (Fig. 27.745) Nanocladius (Plecopteracoluthus)(in part) Thoracic horn long and/or slender (Fig. 27.456), or broader (Fig. 27.455), often with a distal notch

43(42'). 43'. 44(41'). 44'.

43

45(44').

Abdominal segments with a lateral fringe of setae (Fig. 27.468) Xylotopus Abdominal segments without a fringe Brillia (in part) Anal lobes with one macroseta and 2 insertion scars(Fig. 27.453) Brillia (in part) Anal lobes usually with 3 or more macrosetae (Figs. 27.465, 27.466, 27.473-27.475); when there is only one it is on the surface of the lobe (Fig. 27.469) 45 Two marginal anal macrosetae located distally of the 3rd (Fig. 27.466).. ..Mesocricotopus (in part)

45'.

Anal lobe macrosetae not as above

46(45').

Anal lobe with 3 macrosetae located on the surface (Fig. 27.469); long blunt spines on the posterior margins of most terga (Fig. 27.469) Psectrocladius (Allopsectrocladius)

46'.

47(46'). 47'. 48(47').

48'. 49(48'). 49'. 50(49'). 50'.

Not as above

46

47

Anal lobe with 1-3 macrosetae on the surface, when 3, 2 are in the distal half similar to Fig. 27.469; PSB 11 well-developed and with spinules (Fig. 27.470) Monodiamesa Anal lobe with more than 3 macrosetae (Figs. 27.473-27.475); PSB II, when present, without spinules 48 Terga 4 and 5, at least, with central patches of spines (Fig. 27.449) Psectrocladius (Psectrocladius)(in part) Only shagreen present on terga 49 Segment 8 extended as lobes parallel to the anal lobes(Fig. 27.474) Odontomesa Segment 8 without caudolateral lobes 50 Anal lobes with 4 macrosetae, all of which terminal (Fig. 27.475) Prodiamesa Anal lobes with 3 terminal and 1 more-basal macroseta (Fig. 27.465), or anal lobe with more than 4 macrosetae

125

Chapter 27 Chironomidae

1197

Figure 27.454 Figure 27.456 Figure 27.455

0

: v|

Figure 27.457

\ lifi Figure 27.453 Figure 27.458

Ul Figure 27.460 Figure 27.461

Figure 27.459

Figure 27.463

ht//!/ Figure 27.465 Figure 27.462

Figure 27.464 Figure 27.466

Figure 27.453 Anal lobes and segment 8 of Brillia sp. 1. Figure 27.454 Segment 8 (ventral view) of female Brillia sp. 1. Figure 27.455 Thoracic horn of Brillia sp. 1. Figure 27.456 Thoracic horn of Briilia sp. 1. Figure 27.457 Anal lobes and segment 8 of Brillia sp. 2. Figure 27.458 Shagreen (partially Indicated) on segment 6 of Briiiia sp. 2. Figure 27.459 Anal lobes and segment 8 of Heterotrissociadius sp.

Figure 27.460 Posterior margin of sternum 8 of male (upper) and female (lower) of Heterotrissociadius sp.

Figure 27.461 Segment 2 of Heterotrissociadius sp. Figure 27.462 Anal lobes and segment 8 of Platysmittia sp. Figure 27.463 Segments 2 and 3 of Mesocricotopus sp. Figure 27.464 Segment 2 of Platysmittia. Figure 27.465 Anal lobes and segment 8 of Genus 4. Figure 27.466 Anal lobes and segment 8 of Mesocricotopus sp.

1198

51(1').

Chapter 27 Chironomidae

Anal lobes without macrosetae (Figs. 27.478, 27.481, 27.483, 27.484, 27.489, 27.490), although short spines other than reduced macrosetae may be present (Figs. 27.478, 27.483, 27.489, 27.495); Note: some taxa with very small "macrosetae," which may be overlooked, may be keyed either way from this couplet—in doubtful cases specimens should be run in both directions from this couplet

52

Anal lobes with macrosetae that may be hair-like (Figs. 27.488, 27.492, 27.503), spine-like (Figs. 27.523, 27.526, 27.531), or long and slender (Figs. 27.508, 27.516, 27.555)

66

Distinct, nearly circular groups of spines on the central surface of at least some terga (Figs. 27.479, 27.481,27.482)

53

52'.

Terga without such central groups of spines

54

53(52).

Spine groups on terga 4-6(Fig. 27.479); thoracic horn weakly serrate (Fig. 27.477); some thoracic setae branched (Fig. 27.477)

53'.

Spine groups on terga as in Figs. 27.481, 27.482; thoracic horn long and slender without serrations (Fig. 27.480); all thoracic setae simple Orthocladius (Euorthocladius)(in part)

54(52'). 54'.

Anal lobes elongate, narrow and tapering to a point(Fig. 27.484) Anal lobes, if pointed, not shaped as above (Figs. 27.483, 27.489, 27.495)

55(54).

Posterior margins of terga 2-4 with a pair of broad scale-like setae (Figs. 27.485, 27.746);

55'. 56(54').

Posterior margins of terga with simple setae; thoracic horn a short, stout sac(Fig. 27.486)....Genus 5 Posterior margins of terga 4-6, at least with a single row of distinct spines that may

51'. 52(51).

thoracic horn absent

be blunt or sharp (Figs. 27.487-27.489)

56'.

57(56).

57'. 58(57').

Posterior margins of terga 4-6 without a posterior row of spines or with more than 1 row (Figs. 27.490, 27.492, 27.493, 27.495, 27.496, 27.498)

59(56').

55 56 Rheosmittia

57

59

Spines on posterior margins of terga forming a continuous row (Fig. 27.487); the anterior third of most terga with fine needle-like spinules (Fig. 27.487); anal lobes with or without macrosetae (Fig. 27.488) Metriocnemus(in part) Spines on posterior margins in a less continuous row (Fig. 27.489) 58 Anal lobes short and somewhat triangular, often with a few short spines at the apex (Fig. 27.489)

58'.

Abiskomyia

Anal lobes of male somewhat crescent shaped, lobes with a few terminal spines or more numerous spines which extend anteriorly along the dorsal ridge of each lobe (Figs. 27.747, 27.748)

Nearly transparent exuviae; terga with fine shagreen (Fig. 27.490) or shagreen coarse and sometimes increasing in size posteriorly; when shagreen fine, then conjunctiva 2/3-5/6,(or 7/8) usually with groups of weak spinules in middle third (Fig. 27.490)

Georthocladim

Doithrix

126

59'.

Not with above combination of characters

60(59').

Anterior abdominal segments wide, diminishing to a narrow 8th (Fig. 27.491);

60'. 61(60).

Reduction in width of posterior segments much less 62 Anal lobes terminating in spined processes (Fig. 27.493); terga covered with fine dense shagreen; a weak anal macroseta sometimes present (Fig. 27.493).... Cricotopus (Nostococladius) Anal lobes without terminal processes (Fig. 27.492); terga with posterior bands of shagreen; tiny "macrosetae" present(Fig. 27.492) Symbiocladius

especially in female

61'. 62(60').

62'.

60

61

PSB well-developed on segments 2 and 3(Fig. 27.494); anal lobes with many terminal spines (Fig. 27.495); thoracic horn as in Fig. 27.480 Orthocladius {Euorthocladius) in part PSB, at most, developed on segment 2; anal lobes without spines; thoracic horn, when present, not as above 63

Chapter 27 Chironomidae

1199

Figure 27.470

Figure 27.468 Figure 27.469

Figure 27.467

Figure 27.471

Figure 27.472

Figure 27.475

Figure 27.473 Figure 27.476

Figure 27.474

Figure 27.480

Figure 27.477 Figure 27.479

Figure 27.478

Figure 27.467 Anal lobes and segment 8 of Xylotopus sp. Figure 27.468 Lateral margin of segment 5 of Xylotopus sp. Figure 27.469 Anal lobes and posterior margin of segment 8 of Psectrocladius {Allopsectrocladius) sp. Figure 27.470 Right half of segment 2 of Monodiamesa sp. 1.

Figure 27.471 Thoracic horn of Monodiamesa sp. 1. Figure 27.472 Thoracic horn of Monodiamesa sp. 2. Figure 27.473 Anal lobes and segment 8 of Psectrocladius (Psectrocladius) sp. 2. Figure 27.474 Anal lobes and segment 8 of Odontomesa sp.

Figure 27.475 Anal lobes and segment 8 of Prodiamesa sp.

Figure 27.476 Segment 6 of Abiskomyia sp. Figure 7.1 All Anterior thorax and thoracic horn of Abiskomyia sp. Figure 27.478 Anal lobes and segment 8 of Abiskomyia sp. Figure 27.479 Segment 5 of Orthociadius (Euorthociadius) sp. 1. Figure 27.480 Thoracic horn of Orthociadius (Euorthociadius) sp. 1.

1200

63(62').

Chapter 27 Chironomidae

Posterior margins(or conjunctiva) to terga 3-5 with a row of large recurved hooks (Fig. 27.496) Eukiefferiella or Tokunagaia (in part)

63'.

No such large recurved hooks on conjunctiva

64(63').

Posterior margins of, at least, some terga with groups of several rows of weak to strong spines (Figs. 27.498, 27.501, 27.503); thoracic horn a small yellow (Fig. 27.499) or brown sac(Fig. 27.502) Orthocladius (Euorthocladius)(in part)

64

64'.

Not with above combination of characters

65(64').

Anal lobes as in Fig. 27.504; terga with fine dense shagreen that is slightly larger in posterior bands Bryophaenocladius (in part)

65'.

Anal lobes not shaped as above; terga with coarse shagreen;(Note: difficult to separate semiaquatic genera) Bryophaenocladius (in part), Gymnometriocnemus(Gymnometriocnemus), Smittia

66(51').

Terga 4 and 5, at least, with central groups or rows of stronger spines or spinules (coarse shagreen) in addition to smaller shagreen that may be present (Figs. 27.505 and 27.506); thoracic horn usually present (e.g., Figs. 27.507, 27.509, 27.510) but not as in Fig. 27.519, when absent, spines on tergum 4 as in Fig. 27.505; no booklets on conjunctiva as in Figs. 27.539, 27.568, 27.570)

67

66'.

Terga 4 and 5 usually without central groups or rows of spines or coarse spinules, but anterior and posterior bands or rows of strong spines may be present; if such spines are present, then the thoracic horn is either absent or present; when thoracic horn is present it is not as in Figs. 27.507, 27.509, 27.510; if thoracic horn is absent, the tergal armature is not as in Fig. 27.505; booklets may be present on some conjunctiva (Figs. 27.539, 27.568, 27.570)

71

65

67(66).

Tergites 3-6 with a median transverse band of strong spines (e.g.. Fig. 27.438) .... Paracricotopus(in part)

67'.

Tergites not armed as above

68(67).

Median spine fields of terga 2-7 often in 4 somewhat vaguely to sharply defined groups Fig. 27.505); posterior margins of terga with very long spines(Fig. 27.505); thoracic

68

horn absent or present, when present it is a very small nearly spherical, smooth, colorless sac Cardiocladius (in part)

68'.

Median spine fields on terga 2-7 in 2 groups (Fig. 27.506); posterior margins of terga without long spines; thoracic horn not as above (Figs. 27.507, 27.509, 27.510)

69(68').

Thoracic horn an elongated, stalked sac without spinules(Fig. 27.507) (Eudactylocladius)

69'.

Thoracic horn as in Fig. 27.509 or Fig. 27.510

70(69'). 70'.

Thoracic horn slender and tapering (Fig. 27.509) Thoracic horn broad (Fig. 27.510)

71(66').

Anal lobes with 8-12 spine-like macrosetae (often lost)(Fig. 27.511); thoracic horn large (Fig. 27.512)

IV.

Not with above characters

72

72(71').

Anal lobes extremely long, narrow and pointed with 2-3 small macrosetae inserted subterminally, near the midpoint of the lateral margin (Figs. 27.513, 27.514)

73

Anal lobes not as above, when pointed, not as long and tapering (Figs. 27.517, 27.532); anal macrosetae usually larger and inserted closer to the apex (Figs. 27.517, 27.524, 27.535, 27.553, 27.555)

74

72'.

69 Orthocladius 70

Acricotopus Orthocladius {Pogonocladius) Protanypus

73(72).

Thoracic horn a more or less spherical sac similar to Fig. 27.519

73'.

Thoracic horn long and covered with scale-like ridges(Fig. 27.515)

Genus 5

74(72').

Anal lobes terminating in sharp points (Figs. 27.516, 27.517, 27.520, 27.521); thoracic horn usually a more or less spherical sac (Fig. 27.519) but may be more elongate (Fig. 27.522)

Krenosmittia

75

mm-

Figure 27.482

Figure 27.481 Figure 27.483

Figure 27.485

Figure 27.487

Figure 27.486 Figure 27.484

V- .

\j

'V Figure 27.488

Figure 27.491

Figure 27.489 Figure 27.490

Figure 27.493

Figure 27.481 Anal lobes and segment 8 of Orthocladius {Euorthocladius) sp. 1. Figure 27.482 Segment 5 of Orthocladius (Euorthocladius) sp. 2. Figure 27.483 Anal lobes and segment 8 of Orthocladius (Euorthocladius) sp. 2. Figure 27.484 Anal lobes and segment 8 of Rheosmlttia sp. 1. Figure 27.485 Posterior margin of segment 3 of Rheosmlttia sp. 1. Figure 27.486 Anterior thorax and thoracic horn of Genus 5.

Figure 27.487 Segment 4 of Metriocnemus sp.

Figure 27.492

Figure 27.494

Figure 27.488 Anal lobes and segment 8 of Metriocnemus sp.

Figure 27.489 Anal lobes and segment 8 of Georthociadius sp. Figure 27.490 Anal lobes and segments 5-8 of Pseudosmittia sp. Figure 27.491 Abdomen of Symbiociadius sp. Figure 27.492 Anal lobes and segment 8 of Symbiociadius sp. Figure 27.493 Anal lobes and segment 8 of Cricotopus (Nostocociadius) sp. Figure 27.494 Segments 2 and 3 of Orthocladius (Euorthocladius) sp. 3.

1201

1202

Chapter 27 Chironomidae

74'.

Anal lobes not as above; thoracic horn very rarely as above, frequently absent

78

75(74).

Tergites and sternites with anterior transverse dark lines (apophyses)(Fig. 27.749)

76

75'.

Tergites and sternites without such dark lines

76(75).

Tergite 2 with a very small but strongly elevated lobe-like pad of hooks on the posterior margin (Fig. 27.750); anal lobe macrosetae about as long as the anal lobes Acamptocladius

76'.

Tergite 2 without an elevated pad of hooks although a broad less elevated group of hooks may be present; anal lobe macrosetae as long as above, or shorter

77(76').

Pamkiefferiella

77

Abdominal segments sometimes with more than 4 lateral setae (e.g., Fig. 27.516); when 4 or fewer, the macrosetae of the anal lobes are not situated on tubercles

and are usually much shorter than the anal lobes

Epoicocladius

17.

Abdominal segments never with more than 4 lateral setae; anal lobe macrosetae situated on large tubercles (Fig. 27.520) and nearly as long as the anal lobes. Note: not yet known from the Nearctic Lapposmittia

78(74').

One-3 short, spine-like anal macrosetae, none hair-like (Figs. 27.523, 27.524, 27.526-27.528, 27.531-27.534); Note: variation between species of a genus and among specimens of a species, as well as, problems in the interpretation of spine-like and hair-like, make it necessary to key in both directions from this couplet in doubtful cases

78'.

79

One or more long (Figs. 27.543, 27.555, 27.567, 27.575, 27.589) or short (Figs. 27.535, 27.537, 27.541) and hair-like anal macrosetae

86

79(78).

Thoracic horn present

79'.

Thoracic horn absent; Note: the thoracic horn-like structure on the thorax of

80(79).

Boreoheptagyia is not considered to be a thoracic horn in this key Posterior margin of tergum 8 with a row(s) of spines (Figs. 27.524, 27.526-27.528)

80'. 81(80).

Posterior margin of tergum 8 without a row of spines (Fig. 27.523) Thoracic horn as in Fig. 27.525

81'. 82(79'). 82'. 83(82).

Thoracic horn as in Figs. 27.529 or 27.530 Chaetocladius (in part) Anal lobes with 2-3 approximately equal macrosetae (e.g., Figs. 27.532, 27.534) 83 Anal lobes with 2-3 unequal macrosetae (Fig. 27.531) Synorthocladius Anal lobe macrosetae insert on a narrow, pointed, terminal process(Fig. 27.532); posterior margin of tergum 8 with large spines (Fig. 27.532) Parachaetocladius Anal lobe macrosetae not so inserted on a process (Figs. 27.533, 27.534) 84 Anal lobe macrosetae insert terminally (Fig. 27.533) Halocladius Anal lobe macrosetae insert laterally (Fig. 27.534) 85 A conspicuous, dark, triangular thoracic horn-like structure which is covered by small "hairs" is located on the thorax close to the position where the thoracic horn would be found; anal lobe macrosetae more than half the lobe length and somewhat sinuate (Fig. 27.534) Boreoheptagyia No such structure on the thorax; macrosetae of anal lobes very short and thorn-like, much less than half the length of the lobes (Fig. 27.751) Antillocladius Anal lobe macrosetae all short and hair-like (Figs. 27.535, 27.537, 27.541, but not

83'. 84(83'). 84'. 85(84').

85'. 86(78').

80

Baeoctenus Eukiefferiella (in part)

Fig. 27.538), sometimes inconspicuous, and sometimes with more than or less than 3(Fig. 27.503)

86'. 87(86).

82 81

Anal lobe macrosetae longer and usually strong (Figs. 27.543, 27.555, 27.567, 27.575, 27.589), but not spine-like; almost always with 3 macrosetae

87

95

Anal lobes with 4 weak macrosetae (Fig. 27.752); tergites 2-9 with transverse anterior and posterior bands of strong spines (Fig. 27.752) Camptocladius

Figure 27.496

Figure 27.497

Figure 27.495

Figure 27.499 Figure 27.501

Figure 27.500 Figure 27.498 mmm

Figure 27.502 Figure 27.503 Figure 27.504

Figure 27.508

lAWj'ki'i'i&it'.

Figure 27.505

Figure 27.510 Figure 27.509

Figure 27.507 Figure 27.506

Figure 27.495 Anal lobes and segment 8 of Orthocladius (Euorthodadius) sp. 3. Figure 27.496 Segment 4 of Tokunagaia sp. 1. Figure 27.497 Anal lobes and segment 8 of Tokunagaia sp. 1. Figure 27.498 Anal lobes and segment 8 of Orthocladius (Euorthodadius) sp. 4. Figure 27.499 Anterior thorax and thoracic horn of Orthodadius (Euorthodadius) sp. 4. Figure 27.500 Anal lobes and segment 8 of Orthodadius (Euorthodadius) sp. 5. Figure 27.501 Segments 4 and 5 of Orthodadius (Euorthodadius) sp. 5. Figure 27.502 Anterior thorax and thoracic horn of Orthodadius (Euorthodadius) sp. 5.

Figure 27.503 Anai lobes and segment 8 of Orthodadius (Euorthodadius) sp.6 Figure 27.504 Anal lobes and segment 8 of Bryophaenodadius sp. Figure 27.505 Segment of Cardiodadius sp. Figure 27.506 Segment 4 of Orthodadius (Eudactyiodadius) sp. Figure 27.507 Anterior thorax and thoracic horn of Orthodadius (Eudactyiodadius) sp. Figure 27.508 Anal lobes and segment 8 of Orthodadius (Eudactyiodadius) sp. Figure 27.509 Anterior thorax and thoracic horn of Acricotopus sp. Figure 27.510 Anterior thorax and thoracic horn of Orthocladius (Pogonodadius)sp.

1203

Figure 27.512 Figure 27.513 Figure 27.514 Figure 27.511

Figure 27.515

Figure 27.517 Figure 27.516 Figure 27.518

Figure 27.519 Figure 27.522

Figure 27.521

Figure 27.524

Figure 27.520

Figure 27.523

Figure 27.511 Anal lobes and segment 8 of Protanypus sp. Figure 27.512 Anterior thorax and thoracic horn of Protanypus sp. Figure 27.513 Anal lobes and segment 8 of Genus 5 (see Fig. 27.486). Figure 27.514 Anal lobes and segment 8 of Krenosmittia sp. Figure 27.515 Anterior thorax and thoracic horn of Krenosmittia sp. Figure 27.516 Anal lobes and segments 7 and 8 of Epoicocladius sp. 1. Figure 27.517 Anal lobes and segment 8 of Parakiefferieiia sp. 1.

1204

Figure 27.518 Anal lobes and segment 8 of Parakiefferieiia sp. 2. Figure 27.519 Anterior thorax and thoracic horn of Parakiefferieiia sp. 1. Figure 27.520 Anal lobes and segment 8 of Acamptocladius sp. (after Saether 1971). Figure 27.521 Anal lobes and segment 8 of Lapposmittia sp. (after Thienemann 1944). Figure 27.522 Anal lobes and segment 8 of Lapposmittia sp. (after Thienemann 1944). Figure 27.523 Anal lobes and segment 8 of Baeoctenus sp. (after Ssether 1977). Figure 27.524 Anal lobes and segment 8 of Eukiefferiella sp. 1.

Figure 27.526 Figure 27.527

Figure 27.528

Figure 27.525

Figure 27.529

Figure 27.530 Figure 27.531

Figure 27.532

Figure 27.534 Figure 27.533

Figure 27.536

Figure 27.535 ir'f'r'i'l

Figure 27.538

Figure 27.539

Figure 27.540

Figure 27.537

Figure 27.525 Anterior thorax and thoracic horn of Eukiefferiella sp. 1. Figure 27.526 Anal lobes and segment 8 of Chaetocladius sp. 1.

Figure 27.527 Anal lobes and segment 8 of Chaetocladius sp. 2. Figure 27.528 Anal lobes and segment 8 of Chaetocladius sp. 3. Figure 27.529 Anterior thorax and thoracic horn of Chaetocladius sp. 2.

Figure 27.530 Anterior thorax and thoracic horn Chaetocladius sp. 1. Figure 27.531 Anal lobes and segment 8 of Synorthociadius sp. Figure 27.532 Anal lobes and segment 8 of Parachaetociadius sp.

of

Figure 27.533 Anal lobes and segment 8 of Halocladlus sp. Figure 27.534 Anal lobes and segment 8 of Boreoheptagyla sp. Figure 27.535 Anal lobes and segment 8 of Paraphaenociadius sp. Figure 27.536 Anterior thorax and thoracic horn of Paraphaenociadius sp. Figure 27.537 Anal lobes and segment 8 of Pseudorthocladius sp. Figure 27.538 Anal lobes and segment 8 of Eukiefferiella sp. 2. Figure 27.539 Segment 5 of Eukiefferiella sp. 2. Figure 27.540 Anterior thorax and thoracic horn of Eukiefferiella sp. 2.

1205

1206

Chapter 27 Chironomidae

87'.

Anal lobes with no more than 3 macrosetae

88(87').

89'.

Anal lobes with 2 widely spaced and hair-like macrosetae (Fig. 27.753); anal lobes distally truncate (Fig. 27.753) Gymnometriocnemus(Raphidodadius) Macrosetae and anal lobe shape not as above 89 Posterior margins of terga 3(4)-8 with distinct groups of spinules (Figs. 27.501, 27.503); thoracic horn as in Fig. 27.502 Orthodadius (Euorthodadius)(in part) Posterior margins of terga 3(4)-8 without such spines; thoracic horn, when present,

90(89').

Thoracic horn present

91

90'.

Thoracic horn absent

92

91(90).

Wing sheaths with pearl rows similar to Fig. 27.429; PSB developed, at most, on segment 2; anal lobe with 2 weak macrosetae (Fig. 27.535); thoracic horn with a few spinules (Fig. 27.536)

88'. 89(88').

88

not as above

90

Paraphaenodadius

91'.

Wing sheaths without pearl rows; PSB developed on segments 2 and 3(Fig. 27.494); anal lobe with 2-3 short macrosetae (Fig. 27.495); thoracic horn without spinules (Fig. 27.480) Orthodadius {Euorthodadius)(in part)

92(90').

Anal lobes without spines (Figs. 27.488, 27.497, 27.538) Anal lobes with numerous spines(Fig. 27.537)

92'. 93(92).

93 Pseudorthodadius

Some dorsal abdominal conjunctiva usually with a few large booklets (Fig. 27.496)

Eukiefferiella (in part)

93'.

Dorsal conjunctiva without booklets

94

94(93'). 94'.

Terga with posterior rows of spines (Fig. 27.488) Terga without posterior rows of spines (Fig. 27.541)

95(86').

Posterior margins of most terga and usually sterna with heavy spines that are mostly in a single row (Figs. 27.543, 27.545-27.547, 27.549); thoracic horn a thin filament (Fig. 27.542) or a short spur (Figs. 27.544, 27.548) or absent

96

95'.

Not with above combination of characters

98

96(95).

Posterior margins of terga and sterna (3)4-7(8) with heavy spines (Figs. 27.543, 27.545); thoracic horn usually a filament(Fig. 27.542) but may be spur-like (Fig. 27.544)

97

Metriocnemus (in part) Cricotopus {Cricotopus)(in part)

96'.

Posterior margins of only the terga with heavy spines (Figs. 27.546, 27.547, 27.549); thoracic horn spur-like (Fig. 27.548) or absent Pseudokiefferiella

97(96).

A strong tubercle on the thorax between the thoracic horn and the anterior central thoracic seta (Fig. 27.754); many thoracic and abdominal setae long, dark and thick; thoracic horn a long filament Syndiamesa

97'.

No such tubercle on the thorax (Figs. 27.542, 27.544); thoracic and abdominal setae usually shorter and paler (Figs. 27.542-27.544); thoracic horn a thin filament(Fig. 27.542) or spur-like (Fig. 27.544) Diamesa

98(95').

Anal lobes broad, usually with short, terminal (subterminal) pointed projections (Figs. 27.55027.554); no thoracic horn; lateral abdominal setae often branched (Figs. 27.550-27.553) 99

98'.

Not with above combination of characters

103

99(98).

Some lateral abdominal setae branched (Figs. 27.550-27.553)

100

99'.

No branched abdominal setae (Fig. 27.554)

101

100(99). 100'. 101(99').

Anal lobe with a 4th seta along inner margin (Figs. 27.551, 27.552) Anal lobes without a 4th seta (Figs. 27.550, 27.553) Anal lobes distally rounded or truncate

101'.

Anal lobe with inset, distally pointed processes (Figs. 27.550, 27.553, 27.554)

Pagastia Potthastia (in part) Lappodiamesa 102

Chapter 27 Chironomidae

1207

Figure 27.542 Figure 27.544

Figure 27.541

Figure 27.543

Figure 27.545

r Figure 27.547 Figure 27.548

Figure 27.549

Figure 27.551

Figure 27.546

Figure 27.550

Figure 27.552

Figure 27.553

Figure 27.554 Figure 27.555

Figure 27.541 Anal lobes and segment 8 of Cricotopus sp. 1. Figure 27.542 Anterior thorax and thoracic horn of Diamesa sp. 1.

Figure 27.543 Anal lobes and segment 8 of Diamesa sp. 1. Figure 27.544 Anterior thorax and thoracic horn of Diamesa sp. 2.

Figure 27.545 Segment 5 of Diamesa sp. 2. Figure 27.546 Anal lobes and segment 8 of Pseudokieffehelia sp. 1.

Figure 27.549 Segment 5 of Pseudokiefferieiia sp. 2. Figure 27.550 Anal lobes and segments 7 and 8 of Potthastia sp. 1. Figure 27.551 Anal lobes and segment 8 of Pagastia sp. 1.

Figure 27.552 Anal lobes and segment 8 of Pagastia sp. 2.

Figure 27.553 Anal lobes and segment 8 of Potthastia sp. 2. Figure 27.554 Anal lobes and segment 8 of Pseudodiamesa sp.

Figure 27.547 Segment 5 of Pseudokiefferieiia sp. 1.

Figure 27.555 Anal lobes and segment 8 of

Figure 27.548 Anterior thorax and thoracic horn of Pseudokiefferieiia sp. 1.

Limnophyes sp.

1208

Chapter 27 Chironomidae

"'•"HiDiSlfti"*'"'""

Figure 27.557

/ 4

Figure 27.559

Figure 27.558 Figure 27.556

Figure 27.560

r

1

Figure 27.561 Figure 27.562

Figure 27.563

Figure 27.565

> w- ' 66tH 6 «;«»»« ••««*»

Figure 27.566 Figure 27.564 Figure 27.567

V

:

: V :'

Figure 27.568

Figure 27.569

Figure 27.570

Figure 27.556 Anal lobes and segment 8 of Diplocladius sp. Figure 27.557 Anterior thorax and thoracic horn of Diplocladius sp. Figure 27.558 Anal lobes and segment 8 of

Figure 27.563 Anal lobes and segment 8 of Eukiefferieiia sp. 4.

Figure 27.564 Thoracic horn and precorneal setae of Eukiefferieiia sp. 4.

Figure 27.565 Anal lobes and segment 8 of Tokunagaia sp. 2.

Helenieiia sp. Figure 27.559 Segment 2(ventral view) of Heienieiia sp.

Figure 27.566 Segment 5 of Tokunagaia sp. 2. Figure 27.567 Anal lobes and segment 8 of

Figure 27.560 Anterior thorax and thoracic horn of

Eukiefferieiia sp. 5.

Heienieiia sp. Figure 27.561

Figure 27.568 Segment 5 of Eukiefferieiia sp. 6. Figure 27.569 Anterior thorax and thoracic horn of

Segment 5 of Eukiefferieiia sp. 3.

Figure 27.562 Anterior thorax and thoracic horn of

Eukiefferieiia sp. 6.

Eukiefferieiia sp. 3.

Figure 27.570 Segment 5 of Eukiefferieiia sp. 7.

Chapter 27 Chironomidae

1209

102(101'). Terminal anal lobe projections with scale-like spines (Fig. 27.553) Potthastia (in part) 102'. Terminal projections smooth, without scale-like spines (Fig. 27.554) Pseudodiamesa 103(98'). Posterior margins of terga 2-8 with rows of very long, needle-like spines (Fig. 27.555); thoracic horn absent Limnophyes 103'. Posterior margins of terga with or without long spines, when present then thoracic horn is present (Fig. 27.562) 104 104(103'). Anal lobes subcylindrical (e.g.. Fig. 27.556); posterior margins of terga without a row(s) of larger spines

104'.

105

If anal lobes are subcylindrical then posterior margins of most terga with a row(s) of strong spines

107

105(104). Tergites and sternites demarcated by dark transverse lines (apophyses); anal lobes and thoracic horn as in Figs. 27.556 and 27.557; frontal setae present Diplocladius 105'. 106(105'). 106'. 107(104').

Tergites and sternites without dark lines; frontal setae present or absent

106 Frontal setae present; thoracic horn a weak, unarmed, elongated sac Parorthocladius (in part) Frontal setae absent; thoracic horn elongate with small spines Plhudsonia Sterna 2 and 3 with anterior groups of needle-like spines (Fig. 27.559); thoracic horn present (Fig. 27.560)

107'.

Sterna 2 and 3 without needle-like spines, or, if present, thoracic horn absent

Heleniella

108

108(107'). Terga 3-4, 3-5, or 4-5 with rows of large recurved booklets on conjunctiva (Figs. 27.561,

108'.

27.566, 27.568, 27.570, 27.572); thoracic horn often "onion" shaped (Figs. 27.562, 27.569, 27.571, 27.573, 27.574), sometimes absent; wing sheaths without pearl rows Terga 3-5 without such booklets, or, when present thoracic horn as in Figs. 27.580 and 27.582 and wing sheaths with pearl rows similar to Fig. 27.429

109 112

109(108). Terga 3-5 with recurved booklets on conjunctiva (Figs. 27.561, 27.566, 27.568, 27.570, 27.572); thoracic horn usually present (Figs. 27.562, 27.569, 27.571, 27.573, 27.574) 110 109'. Terga 3-4 or 4-5 with recurved booklets on conjunctiva; thoracic horn "onion" shaped (Fig. 27.577) or absent

Ill

110(109). Thoracic horn absent and tergites and dorsal pleurites nearly covered with fields of shagreen Tokunagaia (in part) 110'.

Thoracic horn usually present (e.g.. Figs., 27.564, 27.569, 27.571, 27.573); tergites and dorsal pleurites less extensively and less uniformly covered with shagreen fields Eukiefferiella (in part)

111(109'). Terga 3-4 with recurved booklets on conjunctiva; thoracic horn present(Fig. 27.577); anterior areas of thorax with dark tubercles (Fig. 27.577) Cardiocladius (in part) 11 r. Terga 4—5 with recurved booklets on conjunctiva; thoracic horn absent; anterior areas of thorax without tubercles (also see Fig. 27.578) Tokunagaia (in part) 112(108). Thoracic horn "onion" shaped (Figs. 27.580, 27.582); a few recurved booklets usually present on conjunctiva 3/4-5/6 (Figs. 27.581, 27.583); wing sheaths with pearl rows (similar to Fig. 27.429) Tvetenia 112'.

Not with above combination of characters

113

113(112'). Anal lobes cylindrical with 3 strong macrosetae inserted at tips (Figs. 27.584, 27.586, 27.589); thoracic horn present or absent

114

113'.

117

Anal lobes broader with macrosetae inserted terminally or somewhat subterminally

114(113). Thoracic horn present (Figs. 27.585, 27.587, 27.588)

115

114'.

116

Thoracic horn absent

115(114). Thoracic horn a pale, spineless sac (Fig. 27.585); abdominal segments without long setae (Fig. 27.584) Parorthocladius (in part)

1210

Chapter 27 Chironomidae

■*

Figure 27.571 /

«»!»##•

Figure 27.572

Figure 27.574

Figure 27.573 y

Figure 27.577

Figure 27.575

Figure 27.580

Figure 27.579 Figure 27.576 Figure 27.578

Figure 27.582 :

Figure 27.583

Figure 27.584

Figure 27.581

Eukiefferiella sp. 8. Figure 27.572 Segment 5 of Eukiefferiella sp. 8.

Figure 27.578 Anal lobes and segment 8 of Tokunagala sp. 4. Figure 27.579 Anal lobes and segment 8 of Tvetenia

Figure 27.573

sp. 1.

Figure 27.571

Anterior thorax and thoracic horn of

Anterior thorax and thoracic horn of

Eukiefferiella sp. 9.

Figure 27.574

Anterior thorax and thoracic horn of

Eukiefferiella sp. 10.

Figure 27.575

Anal lobes and segment 8 of

Figure 27.580 Anterior thorax and thoracic horn of Tvetenia sp. 1. Figure 27.581 Segment 5 of Tvetenia sp. 1. Figure 27.582 Anterior thorax and thoracic horn of

Tokunagala sp. 3.

Tvetenia sp. 2.

Figure 27.576

Figure 27.583 Posterior margin of segment 5 of Tvetenia sp. 2. Figure 27.584 Anal lobes and segment 8 of Parorthocladlus sp.

Anal lobes and segment 8 of

Cardlocladlus sp.

Figure 27.577

Anterior thorax and thoracic horn of

Cardlocladlus sp.

Chapter 27 Chironomidae

115'.

Thoracic horn yellowish, long and slender with some spines (Fig. 27.587); abdomen with long setae (Fig. 27.586)

1211

Genus 8

116(114'). Posterior margins of terga 2-8 with single rows of spines which are large on 2-5 and small on 6-8(Fig. 27.590); sterna 5-7 with posterior rows of sharp spines Genus 9 116'. Posterior margins of terga 2-8 with rows of spines which are approximately the same size on all terga (Fig. 27.589); sterna 2-8 with posterior spines; all leg sheaths straight (Fig. 27.353) Lopescladius 117(113'). Thoracic horn brown, contrasting strongly with the thorax (Figs. 27.592, 27.593); exuviae small(3 mm or less)

Stilocladius

117'. Thoracic horn present or absent, when present and brown, exuviae larger than 3 mm 118 118(117'). PSB II elongate and pointed (Fig. 27.430) Parametriocnemus (in part) 118'. PSB II, when present, not as above (Fig. 27.599) 119 119(118'). Most terga with a posterior row of blunt spines similar to Fig. 27.487, thoracic horn similar to Fig. 27.480; tergites 6-8 with median fields of shagreen Thienemannia 119'. Terga without posterior rows of blunt spines, but sharp spines may be present; when present, thoracic horn not as above

120

120(119'). Frontal setae and frontal warts present(Fig. 27.755), and, abdominal tergites 2-8 with transverse posterior rows of sharp teeth

120'.

Psilometriocnemus

Frontal setae and frontal warts present or absent, but, when present tergites 2-8 without posterior rows of sharp teeth

121

121(120'). Posterior margins of tergum 8 with rows of sharp spines (Fig. 27.528); thoracic horn as in Figures 27.529 and 27.530 Chaetocladius (in part) 121'.

Only shagreen on posterior margins of tergum 8

122

122(121'). Thoracic horn short and broad (Fig.27.594); frontal setae on large tubercles Paracladius 122'. Thoracic horn not as above (e.g.. Figs. 27.595, 27.600, 27.601, 27.605-27.607); frontal setae, when present, not on tubercles although warts may be present(Fig. 27.602).... 123 123(122'). Abdominal segments 2-6 with 2-3 multibranched lateral setae Stackelbergina 123'.

Lateral setae of abdominal segments not branched

124

124(123'). Frontal setae usually large (Fig. 27.601); thoracic horn usually large and well pigmented (Figs. 27.595, 27.600, 27.601), never ovoid as in Fig. 27.606; recurved booklets on tergum 2 almost always in more than 2 rows (Fig. 27.599); anal lobes often with terminal spines (Figs. 27.597, 27.598); exuviae frequently yellow-golden-brown; middle abdominal segments sometimes with chitinous rings (Fig. 27.596); conjunctiva sometimes with a reticulate pigmentation similar to Fig. 27.583; frontal setae, when present, always on frontal apotome (Fig. 27.602); frontal warts sometimes present (Fig. 27.602) Orthocladius {Orthocladius and Symposiocladius)(in part) 124'. Frontal setae usually small(Figs. 27.609, 27.610); thoracic horn usually small and weakly pigmented (Figs. 27.605-27.607), sometimes absent; recurved booklets on tergum 2 almost always in 2 rows(Fig. 27.603); anal lobes, at most, with tiny terminal spines(Fig. 27.608); exuviae often with little pigment, but may be yellow or brown; terga never with chitinous rings; conjunctiva rarely with reticulate pigmentation; frontal setae on frontal apotome (Fig. 27.610) or prefrons (Fig. 27.609) or absent; frontal warts absent; anal lobe macrosetae sometimes unequal

(Fig. 27.604)

Cricotopus {Cricotopus)(in part) and Cricotopus(Paratrichocladius)

125(50').

Anal lobes with 3 terminal and 1 more-basal macrosetae (Fig. 27.465)

Genus 4

125'.

Anal lobes with more than 4 macrosetae(some with 7 or more macrosetae)

Pwpsilocerus

126(59).

Tergites II-VII with similar-sized fine spinules covering most of tergites, frontal setae often on prefrons or absent

Hydrosmittia

Figure 27.585

Figure 27.588

Figure 27.587

Figure 27.586

Figure 27.592 Figure 27.589

Figure 27.590 Figure 27.591 Figure 27.593

Figure 27.594

Figure 27.595 Figure 27.596

Figure 27.597

Figure 27.600

Figure 27.599

Figure 27.598

Figure 27.585 Anterior thorax and thoracic horn of

Figure 27.594 Anterior thorax and thoracic horn of

Parorthocladius sp.

Paracladius sp.

Figure 27.586 Anal lobes and segment 8 of Genus 8. Figure 27.587 Anterior thorax and thoracic horn of

Figure 27.595 Anterior thorax and thoracic horn of Orthocladius (Orthociadius) sp. 3. Figure 27.596 Lateral margin of segment 2 of Orthociadius (Orthocladius) sp. 3. Figure 27.597 Anal lobes and posterior margin of segment 8 of Orthociadius (Orthocladius) sp. 3. Figure 27.598 Anal lobes and segment 8 of Orthocladius (Orthocladius) sp. 4. Figure 27.599 Segments 2 and 3 of Orthocladius (Orthocladius) sp. 4. Figure 27.600 Anterior thorax and thoracic horn of Orthocladius (Orthocladius) sp. 5.

Genus 8.

Figure 27.588 Anterior thorax and thoracic horn of Psilometriocnemus sp.

Figure 27.589 Anai lobes and segment 8 of Lopescladius.

Figure 27.590 Posterior margins of segments 5 and 7 of Genus 9.

Figure 27.591

Anal lobes and segment 8 of

Stilocladius sp.1.

Figure 27.592 Anterior thorax and thoracic horn of Stilocladius sp. 1.

Figure 27.593 Anterior thorax and thoracic horn of Stilocladius sp. 2.

1212

Chapter 27 Chironomidae

1213

126'. Shagreen on tergites II-VII coarse and clearly larger at least posteriorly 127 127(126'). Shagreen on tergites II-VII progressively more coarse posteriorly; antepronotal lobes enlarged; frontal setae absent; small lobe-shaped respiratory organ present Chasmatonotus 127'.

Shagreen on tergites II-VIl with anterior and posterior spinules clearly larger than median spinules, resulting in a transversely striped appearance; frontal setae usually present on frontal apotome; respiratory organ absent

128

128(127'). With 2-4 reduced, hair-like anal macrosetae; median hair-like seta often present on anal lobe; male genital sac often with apical papilla

128'.

No reduced anal macrosetae, or when these occasionally present, no median seta and male genital sac without papilla

Allocladius Pseudosmittia

Chironominae

1.

r.

2(1). 2'. 3(2). 3'.

4(3). 4'.

5(4').

5'.

Thoracic horn always unbranched (e.g., Figs. 27.613, 27.618, 27.625, 27.628, 27.631); wing sheaths almost always with a subterminal tubercle ("Nase")(Fig. 27.663); if subterminal tubercle of wing sheath is absent, then at least some terga with conspicuous groups of spines (e.g.. Figs. 27.611, 27.622, 27.652) Thoracic horn almost always with 2 or more branches (Figs. 27.703, 27.706, 27.720); when unbranched, most abdominal terga have large circular areas of fine spinules (Fig. 27.665) and anal lobes without fringe (Fig. 27.666); wing sheaths almost never with a "Nase" Tergum 4(and some others) with one or more distinct groups of short and/or long spines (e.g.. Figs. 27.611,27.614, 27.620,27.622, 27.629, 27.652) Tergum 4(and others) with a more or less uniform field of shagreen, although this may be loosely divided into 2-4 subfields (Figs. 27.658, 27.661) Tergum 4, and sometimes 3 and 5 with conspicuous groups of long needle-like spines (e.g., Figs. 27.611, 27.612, 27.614, 27.616, 27.619) Terga 4 and 5 without groups of needle-like spines (e.g.. Figs. 27.626, 27.629, 27.636, 27.640, 27.652) although some long spines may be present on tergum 3 and tergum 4 and 5 may have groups of short spines

2

19 3

18 4

8

Tergum 4 with 2 longitudinal rows of needle-like spines that are angled medially at their anterior ends and usually meet(Figs. 27.611, 27.612) Micropsectm (in part) Tergum 4 not as above i.e., the 2 spine rows are not joined anteriorly (e.g.. Figs. 27.617, 27.619, 27.621), sometimes additional spine groups are present (e.g., Figs. 27.614, 27.616) or the rows are transverse (Fig. 27.622) 5

Tergum 4 with 2 longitudinal rows of spines(although sometimes very weakly developed) and 1-2 anterior median groups of spines (Figs. 27.614, 27.616); wing sheaths with or without a pearl row similar to Fig. 27.429 Pamtanytarsus (in part) Tergum 4 with 2 longitudinal rows of spines but without median groups (Figs. 27.617, 27.619-27.621) or, tergum 4 with spine rows transverse (Fig. 27.622); wing sheath without pearl rows

6

6(5').

Tergum 4 with transverse rows of spines(Fig. 27.622)

6'.

Tergum 4 with 2 longitudinal rows of spines (Figs. 27.617, 27.619-27.621)

7(6').

Longitudinal spine rows of tergite 4 sinuate, with anterior spines much shorter than posterior spines and directed postero-medially (Fig. 27.756); anal comb very broad (Fig. 27.757) Tanytarsm (in part)

7'.

If longitudinal spine rows of tergite 4 are sinuate, the anterior spines are about the same length as the posterior spines(Fig. 27.621); anal comb much less broad

(e.g.. Fig. 27.654)

Sublettea

7

Tanytarsus (in part)

1214

Chapter 27 Chironomidae

n/igutm.

Figure 27.601

Figure 27.602 Figure 27.603

Figure 27.604

Figure 27.605

Figure 27.607

Figure 27.606

Figure 27.608 Figure 27.609 Figure 27.610

Figure 27.611 Figure 27.612

Figure 27.613

Figure 27.614

Figure 27.616

Figure 27.615

Figure 27.601 Anterior thorax and thoracic horn of Orthocladius (Orthocladius) sp. 6. Figure 27.602 Frontal apotome of Orthocladius {Orthocladius) sp. 7. Figure 27.603 Segments 2 and 3 of Cricotopus sp. 2. Figure 27.604 Anal lobes and posterior margin of segment 8 of Cricotopus sp. 3. Figure 27.605 Anterior thorax and thoracic horn of Cricotopus sp. 4.

Figure 27.606 Anterior thorax and thoracic horn of Cricotopus sp. 5.

Figure 27.607 Anterior thorax and thoracic horn of Cricotopus sp. 6.

Figure 27.608 Anal lobes of Cricotopus sp. 7.

Figure 27.609 Frontal apotome of some Cricotopus spp.

Figure 27.610 Frontal apotome of some Cricotopus spp.

Figure 27.611 Segments 3-5 of Micropsectra sp. 1. Figure 27.612 Groups of spines of segment 4 of Micropsectra sp. 1. Figure 27.613 Thoracic horn of Micropsectra sp. 1. Figure 27.614 Segments 3-5 of Paratanytarsus sp. 1. Figure 27.615 Anal lobes and posterior margin of segment 8 of Paratanytarsus sp. 1. Figure 27.616 Segment 4 of Paratanytarsus sp. 2.

Chapter 27 Chironomidae

1215

J

1

Figure 27.618

Figure 27.619

Sf Figure 27.621

Figure 27.620

Figure 27.617 Figure 27.625 Figure 27.623

U-

n i

1

Figure 27.628

rr

Figure 27.627

Figure 27.624

/ Figure 27.622

Figure 27.631

r~ Figure 27.626 \

^ >■

Figure 27.630

4 -!

Figure 27.629

Figure 27.632 Figure 27.633 Figure 27.634

Figure 27.617 {Tanytarsus) sp. Figure 27.618 (Tanytarsus) sp. Figure 27.619

Segments 3-6 of Tanytarsus 1. Thoracic horn of Tanytarsus 1. Segments 3 of Tanytarsus (Tanytarsus)

sp. 1.

Figure 27.620 (Tanytarsus) sp. Figure 27.621 (Tanytarsus) sp. Figure 27.622 Figure 27.623

Segments 3-6 of Tanytarsus 2. Segments 3 and 4 of Tanytarsus 3. Segments 2-5 of Sublettea sp. Anal lobes and segment 8 of Sublettea

sp.

Figure 27.624 Posterior margin of segment 8 (ventral view) of Sublettea sp. Figure 27.625 Thoracic horn of Sublettea sp.

Figure 27.626

Segments 4 and 5 of Paratanytarsus

sp. 3.

Figure 27.627 Anal lobes and segment 8 of Paratanytarsus sp. 3. Figure 27.628 Thoracic horn of Paratanytarsus sp. 3. Figure 27.629 Segments 4 and 5 of Stempelllna sp. Figure 27.630 Anal lobes and segment 8 of Stempelllna sp. Figure 27.631 Frontal apotome and anterior thorax with thoracic horn of Stempelllna sp. Figure 27.632 Frontal apotome and anterior thorax with thoracic horn of Constempelllna sp. 1. Figure 27.633 Segment 5 of Constempelllna sp. 1. Figure 27.634 Anal lobes and segment 8 of Constempelllna sp. 1.

1216

8(3'). 8'. 9(8'). 9'.

10(9). 10'.

11(9'). 11'.

12(11').

Chapter 27 Chironomidae

Tergum 4 with a single median group of short spines (e.g., Fig 27.626).... Paratanytarsus (in part) Tergum 4 with paired groups of short spines (e.g., Figs. 27.629, 27.638, 27.640 27.645, 27.655)... 9 Strong lateral spines on segment 8(Figs. 27.630 and 27.634); shagreen and spine groups on terga as in Figs. 27.629, 27.633, and 27.636 10 Spines on segment 8 usually in the form of a caudolateral comb (e.g., Figs. 27.637, 27.646, 27.648, 27.650); however, when single (e.g., Fig. 27.641), tergal spine groups never as above 11 Frontal setae not spine-like (Fig. 27.631); spine and shagreen groups on terga 4 and 5 as in Fig. 27.629 Stempellina Frontal setae spine-like (Figs. 27.632, 27.635) and the spine and shagreen groups on tergum 5 as in Fig. 27.633 or 27.636 Constempellina

Terga 5 and 6 with large oval groups of small spines (Fig. 27.638); anal lobes with fringe setae limited to distal ends (Fig. 27.637); generally lotic; 2-3 mm in length Terga 5 and 6 without such large areas of spinules but smaller groups may be present (Figs. 27.640, 27.645, 27.652, 27.655); anal lobes usually with more complete fringe (Figs. 27.641, 27.646, 27.648, 27.650, 27.654) Caudolateral spine on segment 8 usually a simple spur (Fig. 27.641)less often with a small accessory spine

12'.

Neozavt-elia

12

Rheotanytarsus

Caudolateral spines on segment 8 in the form of a comb (Figs. 27.646, 27.648, 21.650,21.653,11.eSA)

13

Each anal lobe with 1 or 2 dorsal seta (e.g.. Fig. 27.646), if 2, then paired patches of short spines are present on terga 3-6 and most tergites and pleurites are strongly shagreened

14

13'.

Each anal lobe with 2 dorsal setae (Figs. 27.648, 27.650, 27.654); when paired patches of short spines are present on terga 3-6 the tergites and pleurites are not strongly shagreened

15

14(13). 14'.

Paired groups of short spines on terga (2)3-6 (Fig. 27.644) Paired groups of short spines on terga 4-5 or 4-6(Fig. 27.645)

15(13').

Combs on caudolateral corners of segment 8 wide (Fig. 27.648); precorneal setae lamellar and inserting on a mound (Figs. 27.647, 27.649)

Cladotanytarsus

15'.

Combs on 8 usually less wide (Figs. 27.650, 27.653); precorneal setae not as above

16

16(15'). 16'. 17(16).

Fringe of anal lobe limited to no more than the distal half of the margin (Fig. 27.650) 17 Fringe of anal lobes usually extending along entire margin (e.g.. Fig. 27.654). . . Tanytarsus (in part) Thoracic horn with long hairs (Fig. 27.758) Micmpsectra (in part) Thoracic horn without setae (Fig. 27.651) Corynocera Terga 2-6 with shagreen occupying about one-third to one-half of the surface; fields of pleura with, at most, weak shagreen (Fig. 27.658) Stempellinella Terga 2-6 with larger fields of shagreen, those on tergite 2 as large as those on tergite 3 (Fig. 27.759);[Note; a few unplaced, undescribed or poorly described species of Tanytarsini (e.g.. Figs. 27.661, 27.662) may key here, including possibly some specimens of the species Neostempellina reissi Caldwell] Zavrelia Thoracic horn unbranched (Fig. 27.664); terga with large circular areas of fine spinules (Fig. 27.669); and lobes without fringe (Fig. 27.666) Pseudochironomus (in part) Thoracic horn with 2 or more branches (e.g., Figs. 27.674, 27.703, 27.706, 27.720); anal lobes with, at least, a partial fringe (e.g.. Figs. 27.676, 27.693, 27.711, 27.716) 20 Row of booklets on posterior margin of segment 2 distinctly interrupted (Figs. 27.669, 27.671, 27.679) or, rarely, absent 21 Row of booklets on segment 2, at most, very narrowly interrupted (Figs. 27.691, 27.709, 27.710, 27.717, 27.728), always present 31

13(12').

17'. 18(2').

18'.

19(1'). 19'.

20(19'). 20'.

Micmpsectra (in part) Micmpsectra (in part)

Chapter 27 Chironomidae

Figure 27.635

1217

Figure 27.636 Figure 27.638 Figure 27.637

Figure 27.639

/ Figure 27.640 Figure 27.641

Figure 27.642

Figure 27.643

Figure 27.Mi

Figure 27.645

Figure 27.644

Figure 27.646

Figure 27.651

Figure 27.648

Figure 27.649

Figure 27.650 Figure 27.652

Figure 27.635 Frontal apotome and anterior thorax with thoracic horn of Constempellina sp. 2. Figure 27.636 Segment 5 of Constempellina sp. 2. Figure 27.637 Anai lobes and segment 8 of Neozavrelia sp. Figure 27.638 Segments 5 and 6 of Neozavrelia sp. Figure 27.639 Thoracic horn of Neozavrelia sp. Figure 27.640 Segments 5 and 6 of Rheotanytarsus sp. 1.

Figure 27.646 Anai iobes and segment 8 of Micropsectra sp,. 2. Figure 27.647 Anterior thorax and thoracic horn of Cladotanytarsus sp. 1. Figure 27.648 Anal lobes and segment 8 of Cladotanytarsus sp. 1. Figure 27.649 Anterior thorax and thoracic horn of Cladotanytarsus sp. 2. Figure 27.650 Anal lobe and segment 8 (left side) of

Figure 27.641 Anal lobes and segment 8 of Rheotanytarsus sp. 1. Figure 27.642 Anterior thorax and thoracic horn of Rheotanytarsus sp. 1. Figure 27.643 Thoracic horn of Rheotanytarsus sp. 2.

Corynocera sp. Figure 27.651 Thoracic horn and precorneal setae of Corynocera sp.

Figure 27.644 Segment 4 of Micropsectra sp. Figure 27.645 Segment 5 of Micropsectra sp. 2.

Figure 27.652 Segment 2-5 of Tanytarsus (Tanytarsus) sp. 4.

1218

Chapter 27 Chironomidae

Figure 27.655 Figure 27.653

Figure 27.657 Figure 27.654 Figure 27.656

Figure 27.663 Figure 27.658

Figure 27.659

Figure 27.662

Figure 27.660 Figure 27.665 '.rl

J Figure 27.661

Figure 27.669

Figure 27.664 Figure 27.667

Figure 27.666

Figure 27.668

Figure 27.662 Anal lobes and segment 8 of

Figure 27.653 Spines on caudolaterai margin of segment 8 of Tanytarsus (Tanytarsus) sp. 4. Figure 27.654 Anal lobes and segment 8 of

"Zavrella" sp.

Tanytarsus (Tanytarsus) sp. 5.

"Zavrella" sp.

Figure 27.663 Tip of wing sheath with "Nase" of

Figure 27.655 Segment 4 of Tanytarsus (Tanytarsus)

Figure 27.664 Thoracic horn of Pseudochlronomus

sp. 5.

sp. 1.

Figure 27.656 Anal lobe and segment 8 (left half) of

Figure 27.665 Segment 6 (left half) of

"Stempellinella" sp.

Pseudochlronomus sp. 1.

Figure 27.657 Frontal apotome and anterior thorax

Figure 27.666 Anal lobes and segment 8 of

with thoracic horn of "Stempellinella" sp. 1.

Pseudochlronomus sp. 1.

Figure 27.658 Segment 4 of Stempellinella sp. 1. Figure 27.659 Anal lobes and segment 8

Figure 27.667 Segment 6 of Cladopelma sp. Figure 27.668 Frontal apotome of

Stempellinella sp. 2.

Microchlronomus sp.

Figure 27.660 Frontal apotome and anterior thorax

Figure 27.669 Segment 2 of Microchlronomus sp.

with thoracic horn of "Zavrella" sp.

Figure 27.661 Segment 4 of "Zavrella" sp.

Chapter 27 Chironomidae

21(20).

1219

21'.

Caudolateral margins of segment 8 with a spine or group of spines (e.g., Figs. 27.692-27.694, 27.697, 27.704) Caudolateral margins of segment 8 without spines

22 25

22(21). 22'.

Thoracic horn exceptionally long (Fig. 27.674) Thoracic horn never as long as above (Figs. 27.703, 27.706, 27.720)

Cryptotendipes(in part) 23

23(22').

Tergum 6 with a posterior, median spiniferous process(Fig. 27.667)

Cladopelma

23'.

Tergum 6 with, at most, rows of spines along the posterior margin

24(23'). 24'.

Cephalic tubercles long (Fig. 27.668); PSB II developed (Fig. 27.669) Cephalic tubercles short(Fig. 27.670); PSB 11 absent

25(21').

Hooklets on posterior margin of tergum 2 absent; large tubercles on terga (Fig. 27.673); thoracic horn exceptionally long (Fig. 27.674) Cryptotendipes (in part)

25'.

Not with above characters

26(25'). 26'. 27(26'). 27. 28(27'). 28'.

Caudal region with a forked posterior extension (Fig. 27.676); frontal apotome often with ornate cephalic tubercles (e.g.. Figs. 27.677, 27.678) Cryptochironomus Caudal region without a forked process; cephalic tubercles, when present, never ornate 27 PSB II with spinules (Fig. 27.679) Beckidia PSB II, when present, without spinules 28 Posterior margins of tergites 2-5(6) with a row(s) of long needle-like spines Kloosia Tergites without posterior needle-like spines 29

29(28').

Cephalic tubercles absent

24 Microchironomus Genus 12

26

Chernovskiia

29'.

Cephalic tubercles present

30(29').

Each caudal lobe with more than 100 fringe setae

30

30'.

Each caudal lobe with no more than about 50-60 setae

31(20').

Caudolateral margins of segment 8 with a spine or a group of spines

Genus 13 Harnischia

(e.g., Figs. 27.697-27.699, 27.704, 27.712)

32

31'.

Caudolateral margins of segment 8 without spines

32(31).

33(32').

Thoracic horn with only 2 branches, one of which has numerous short spines distally (Fig. 27.760); terga 2-5(6) with paired transverse patches of spines (Fig. 27.761). .. Lauterborniella If the thoracic horn has only 2 branches there are never distal spines (e.g, Fig. 27.703); terga usually without paired spine patches 33 Cephalic tubercles present (e.g., Figs. 27.684-27.687) 34

33'.

Cephalic tubercles absent

34(33). 34'.

Thoracic horn exceptionally long (Fig. 27.674) Thoracic horn shorter (e.g.. Figs. 27.703, 27.706, 27.720)

35(34').

Terga 2-6, 3-6, or 4-5 with small to large unpaired groups of spines or spiniferous processes (Figs. 27.680, 27.682, 27.683)

36

Terga without such groups of spines or processes

39

32'.

35'.

89

78

Cryptotendipes (in part) 35

36(35).

Terga 2-6 with large spiniferous processes(Fig. 27.680)... .Glyptotendipes (Glyptotendipes)(in part)

36'.

Terga 3-6 or 4-5(6) with smaller spine groups (Figs. 27.682, 27.683)

37(36'). 37'.

Epaulettes on terga 3-6 Small epaulettes on terga 4-5(6)(Fig. 27.683)

37 38 Demijerea

38(37).

Epaulettes on terga 3-6 all generally of about the same size (Fig. 27.682)

38'.

Epaulettes distinctly increasing in size from tergite 3 to tergite 6

Glyptotendipes

Glyptotendipes(Caulochironomus)

1220

Chapter 27 Chironomidae

39(35').

Cephalic tubercles truncate and with a cluster of spinules (Figs. 27.684, 27.764, 27.765)

40

39'.

Cephalic tubercles not truncate and without spinules

43

40(39).

Conjunctives 3-4 and/or 4-5 with a few large dark spines on each side of the midline (Fig. 27.762)

40'.

Conjunctives with, at most, a field(s) of small spinules

41(40').

Anterior conjunctives conspicuously darkened laterally (Fig. 27.763)

41'.

Anterior conjuctives without lateral darkening

42(41').

Frontal setae long, about twice the diameter of the truncated apexes of the cepahlic tubercles (Fig. 27.765)

42'.

Frontal setae at most slightly longer than the truncated apex of the cepahlic tubercles (Fig. 27.765)

43(39').

Hyporhygma 41 Phaenopsectra 42 Endotribelos Sergentia

Cephalic tubercles extremely long and tapering; frontal setae attached near the bases of the tubercles (Fig. 27.685)

Polypedilum (in part)

43'.

If cephalic tubercles very long, not as above (e.g.. Figs. 27.686, 27.687)

44

44(43').

Cephalic tubercles large, heavily sclerotized, and fused (Fig. 27.686)

44'.

Cephalic tubercles not as above (e.g.. Figs. 27.668, 27.670, 27.687)

45(44').

Cephalic tubercles long to extremely long, with frontal setae attached near the tips; and, caudolateral armature of segment 8 with blunt teeth which are weakly serrate at the tips (Fig. 27.766)

Polypedilum (in part) 45

Lipiniella

45'. 46(45').

Cephalic tubercles not so long, or, armature of segment 8 consisting of sharp spines (teeth).... 46 Frontal apotome with large frontal warts and cephalic tubercles (Fig. 27.687), and, caudolateral armature of segment 8 not a compound spur as in Figs. 27.704 and 27.705 Einfeldia (in part)

46'.

Only cephalic tubercles present, or, caudolateral armature of segment 8 a compound spur as in Figs. 27.704 and 27.705

47

47(46').

Sternum 2 and sometimes 1 and 3 with rows of needle-like spines (Figs. 27.688, 27.689), and, caudolateral armature of segment 8 always strong (e.g.. Figs. 27.696, 27.697, 27.767)

48

47'.

Sternum 1-3 without such spines, or, when present, caudolateral armature of segments weak ... 50

48(47).

Needle-like spines on sternum 2 in transverse rows only (e.g., Fig. 27.688)

48'.

Needle-like spines on sternum 2 in transverse and longitudinal rows

(Fig. 27.689)

49 Kiefferulus (Wirthiella)

49(48).

Armature of caudolateral margins of segment 8 usually a simple spine—sometimes with 1 or 2 accessory spines (e.g.. Figs. 27.696, 27.697) Dicrotendipes (in part)

49'.

Armature of segment consisting of 4-6 parallel spines with a common base (e.g., Fig. 27.767) Goeldichironomus (in part)

50(47').

Caudolateral armature of segment 8 usually a single (or double), often sinuate, spine (e.g.. Figs. 27.696, 27.697); rarely the armature of 8 includes a few spines along the entire margin; gonopod sheaths of male never with distal spines; tergites 2-8 without paired spine patches

51

Caudolateral armature of segment 8 a comb of weak to strong spines or a compound spur; if the armature is single, the gonopod sheath of the male has distal spines, or, tergites 2-6 have paired spine patches

52

50'.

51(50).

Caudolateral armature of segment 8 usually a single straight or sinuate spine (e.g.. Figs. 27.696, 27.697); the largest shagreen spinules on tergites 3-6 tend to be in the middle of the tergites or along the posterior margin, or, both, but not with distinct anterior bands on tergites 2-6; conjunctives 3/4—5/6 usually with very small shagreen Dicrotendipes (in part)

Chapter 27 Chironomidae

1221

cephalic tubercles

Figure 27.670 Figure 27.671

Figure 27.673

Figure 27.672

Figure 27.674

Figure 27.675

Figure 27.677

Figure 27.678

Figure 27.676

■/

.

Figure 27.679

Figure 27.680

Figure 27.682 Figure 27.681

frontal tubercles

)iy

Figure 27.684 Figure 27.683

Figure 27.670 Figure 27.671 Figure 27.672

Frontal apotome of Genus 12. Segment 2 of Genus 12. Gaudolateral margin of segment 8 of

Genus 12.

Figure 27.673 Segment 5 of Cryptotendipes sp. Figure 27.674 Ttioracic horn of Cryptotendipes sp. Figure 27.675 Gaudolateral margin of segment 8 (left half) of Cryptotendipes sp. Figure 27.676 Anal lobes and segment 8 of Cryptochironomus sp. 1. Figure 27.677 Frontal apotome of Cryptochironomus sp. 2. Figure 27.678 Frontal apotome of Cryptochironomus sp. 3.

Figure 27.686

Figure 27.685

Figure 27.679 Figure 27.680

Segment 2 of Beckidia sp. Segment 6 of Giyptotendipes

(Giyptotendipes) sp.

Figure 27.681 Gaudolateral margin of segment 8 (left half) of Giyptotendipes (Giyptotendipes) sp. Figure 27.682 Segment 6 of Giyptotendipes (Trichotendipes) sp. Figure 27.683 Segment 5 of Demeijerea sp. Figure 27.684 Frontal apotome of Phaenopsectra sp. Figure 27.685 Frontal apotome of Poiypediium sp. 1. Figure 27.686 Frontal apotome of Poiypediium sp. 2.

1222

Chapter 27 Chironomidae

51'.

Caudolateral armature of segment 8 simple, but sometimes there are additional spines spread along the entire margin; shagreen of tergites 2-6 strongest in anterior transverse bands; only conjunctive 4/5 with shagreen Beardius

52(50').

Comb on segment 8 composed of 4-5 or more strong, nearly parallel spines (Fig. 27.767), and, segment 8 has 5 lateral lamellar setae

53

52'.

If the comb on segment 8 is as above, segment 8 has 4 lateral lamellar setae

55

53(52). 53'.

Conjunctiva 3/4-5/6 without fields of shagreen Conjunctiva 3/4-5/6 with fields of shagreen

54(53').

Most of tergites 2-5 covered with shagreen of nearly uniform size

54'.

Most of tergites 2-5 free of shagreen although shagreen is present it may be confined to relatively small patches Einfeldia (in part) Tergite 6 with the posterior armature stronger than on other tergites Pamchiwnomus(in part)

55(52').

Goeldichironomus (in part) 54 Kieffemlus (Kiefferulus)

55'.

Not as above

56

56(55').

Segment 5 with 1-3 lamellar lateral setae (Figs. 27.690, 27.695)

57

56'. 57(56).

Segment 5 with 4-5 lamellar lateral setae (Fig. 27.707) Segment 5 with 1-2 lamellar lateral setae (Fig. 27.690)

57'.

Segment 5 with 3 lamellar lateral setae (Fig. 27.695)

66 "Pamtendipes"(in part) 58

58(57').

Segment 6 with 3 lamellar lateral setae (Fig. 27.721)

59

58'.

Segment 6 with 4 lamellar lateral setae (Fig. 27.695)

62

59(58).

Terga 2-6 with paired groups of spines (Fig. 27.691); spine(s) on caudolateral margin of segment 8 as in Fig. 27.692

Zavreliella

59'. 60(59'). 60'.

Terga 2-6 without paired groups of spines; spines on segment 8 as in Fig. 27.693 60 Fringe of setae on caudal lobes completely uniserial (Fig. 27.693) Polypedilum (in part) Fringe of setae on caudal lobes at least partially multiserial 61

61(60').

Segment 8 with 3-4 lateral lamellar setae

61'.

Segment 8 with 5 lateral lamellar setae

62(58').

Segment 8 with 3-4 lamellar lateral setae, frontal setae present

62'.

Segment 8 with 5 lamellar lateral setae, if 4, then frontal setae absent

63(62').

Armature on caudolateral margins of segment 8 in the form of a compound spur (e.g., Fig. 27.704)

63'.

Stictochironomus Tribelos

Pamtendipes (in part)

Chironomus (in part) Armature on margins of segment 8 consisting of a single spine (e.g. Fig. 27.694) or groups of spines

64(63'). 64'. 65(64'). 65'.

66(56'). 66'. 67(66).

63

Armature on margins of segment 8 often consisting of single spines (Fig. 27.694); anal lobes with 15 or fewer fringe setae Armature on margins of segment 8 usually consisting of multiple spines; anal lobes usually with many more than 15 finge setae Terga 2-5 or 3-5 with paired groups of dark spines similar to Fig. 27.761; frontal setae present

64

Pagastiella 65 Omisus

Terga without such paired groups, although isolated fields of shagreen may be present; frontal setae absent, but short to rather long cephalic tubercles are present Microtendipes Segment 8 with 4-5 lamellar lateral setae, when 5, caudolateral armature on segment 8 similar to Figs. 27.696, 27.697 67 Segment 8 with 5 lateral setae(5 lamellar or 4 lamellar and 1 hair-like); armature on segment 8 usually not like Figs. 27.696, 27.697 69 Spines on caudolateral margins of segment 8 usually simple as in Figs. 27.696, 27.697 Dicwtendipes (in part)

Chapter 27 Chironomidae

I

( Figure 27.687

Figure 27.688

1223

\

%

Ii

Figure 27.689

Figure 27.690 Figure 27.691

=r Figure 27.694 Figure 27.696

Figure 27.692

Figure 27.695

Figure 27.693

Figure 27.699 Figure 27.700

Figure 27.697

Figure 27.698

Figure 27.702

Figure 27.704 Figure 27.701

Figure 27.687 Frontal apotome of Einfeldia sp. 1. Figure 27.688 Segment 2(ventral view, left tialf) of Dicrotendipes sp. 1 Figure 27.689 Segment 2(ventral view, left half) of Kiefferulus (Wirthiella) sp. Figure 27.690 Segment 5 (right half) of Genus 15. Figure 27.691 Posterior margin of segment 2 and segment 3 of Zavreliella sp. Figure 27.692 Caudolateral margin of segment 8 (right half) of Zavreliella sp. Figure 27.693 Anal lobes and segment 8 of Polypedllum sp. 3 Figure 27.694 Caudolateral margin of segment 8 (left half) of Pagastlella sp. Figure 27.695 Segments 5 and 6 (left half) of Paratendlpes sp. Figure 27.696 Caudolateral margin of segment 8 (left half) of Dicrotendipes sp. 2.

Figure 27.703

Figure 27.697 Caudolateral margin of segment 8 (left half) of Dicrotendipes sp. 3. Figure 27.698 Caudolateral margin of segment 8 (left half) of Paralauterbornlella sp. Figure 27.699 Caudolateral margin of segment 8 (left half) of Cyphomella sp. Figure 27.700 Segment 8(ventral view, left side) of male of Demlcryptochlronomus sp. Figure 27.701 Segment 8(ventral view of posterior margin) of female of Demlcryptochlronomus sp. Figure 27.702 Segment 1 (ventral view) of Pseudochlronomus sp. 2. Figure 27.703 Thoracic horn of Pseudochlronomus sp. 2. Figure 27.704 Caudolateral margin of segment 8 (left half of Chlronomus sp. 1.

1224

Chapter 27 Chironomidae

67'.

Spines on caudolateral margins of segment 8 multiple (e.g., Figs. 27.698, 27.699)

68(67').

Caudolateral margins of segment 8 with 3 or more short straight spines (Fig. 27.698)

68'.

68

Paralauterborniella

Caudolateral margins of segment 8 with a series of closely set curved spines (Fig. 27.699)

Cyphomella

69(66').

Posterior margin of sternum 8 of female with 2 sclerotized spines (e.g. Fig. 27.701)

70

69'.

No such spines present

71

70(69).

Posterior shagreen of tergites 2-3(4) very strong and dark; posterior margin of sternite 8 of male with a row of spines(Fig. 27.700) Demicryptochironomus(Irmakia)

70'.

Posterior shagreen of tergites 2-3(4) not very strong and dark; posterior margin of sternite 8 of male with a row of spines Demicryptochironomus {Demicryptochironomus)

71(69'). 71'.

Sternum I with 2 pairs of spiniferous processes (Fig. 27.702) Sternum I without such processes

72(71').

Spines on caudolateral corners of segment 8 in the form of a compound spur (Figs. 27.704 and 27.705) Chironomus(in part)

72'. 73(72').

Spines on segment 8 not as above (e.g.. Figs. 27.708, 27.712, 27.718, 27.719) Spines on caudolateral margins of segment 8 single, and, gonopod sheaths of

Pseudochironomus (in part) 72

73

male with distal spines

73'. 74(73').

Gillotia

Spines on segment 8 multiple (e.g.. Figs. 27.708, 27.712, 27.718) Spines on caudolateral margins of segment 8 long, yellow, and slightly curved (Fig. 27.708)

74 Einfeldia (in part)

74'.

Spines on segment 8 not as above

75

75(74').

PSB II present(Fig. 27.709)

76

75'.

PSB II absent

77

76(75). 76'.

Segment 4 with no lateral lamellar setae

Segment 4 with 1-2 lateral lamellar setae

Apedilum Parachironomus (in part)

77(75').

Fringe of setae on anal lobes uniserial(Fig. 27.711); armature on segment 8 may be as in Fig. 27.712 Paracladopelma (in part)

IT.

Fringe of setae on anal lobes multiserial

Genus 17

78(33').

Segment 5 with no lamellar lateral setae (Fig. 27.715)

79

78'.

Segment 5 with 3^ lamellar lateral setae (Figs. 27.707, 27.721)

80

79(78).

Anterior regions of terga 2-5 with single or multiple rows of strong spines (Fig. 27.714)

79'.

Polypedilum (impaTt)

Anterior regions of terga 2-6 with multiple rows of smaller spines (Fig. 27.715), a tuft of fringe setae on the apex of each anal lobe

Endochironomus

80(78').

Segment 5 with 3 lateral lamellar setae (Fig. 27.721)

81

80'. 81(80). 81'.

Segment 5 with 4 lateral lamellar setae (Fig. 27.707) Segment 6 with 3 lateral lamellar setae (Fig. 27.721) Segment 6 with 4 lateral lamellar setae (Fig. 27.707)

86 82 85

82(81). 82'.

Fringe of setae on anal lobe extends along inner margin (Fig. 27.716) Fringe of setae does not extend along inner margin of anal lobe

83(82').

Thoracic horn rarely with more than 16 branches; fringe of setae on anal lobes usually uniserial as in Fig. 27.693, but may be completely multiserial in some species Polypedilum (in part)

Genus 18 83

Chapter 27 Chironomidae

83'.

Thoracic horn plumose; fringe of setae on anal lobes distally multiserial(Fig. 27.722)

84(83').

Anal lobes with 2 rows of fringe setae along the distal margins (e.g., Fig. 27.722)

84'.

Anal lobes with 2-4 rows of fringe setae along most of the margins Steimum 1 with 2 pairs of spiniferous processes (Fig. 27.702) Sternum 1 without such processes

85(81'). 85'.

84 "Gmceus"

Synendotendipes Pseudochironomus (in part) Axams

86(80').

Armature on caudolateral margins of segment 8 as in Fig. 27.723, median shagreen patches on terga 7 and 8

86'.

Armature on caudolateral margins of segment 8 not as above

87(86').

Armature on caudolateral margin of segment 8 similar to Fig. 27.724

87'.

Armature on segment 8 not as above Row of booklets on segment 2 narrowly interrupted; frontal apotome with a pair

88(87').

1225

of swollen mounds

Nilothauma 87

Stenochironomus 88 Xestochironomus

Hook row on segment 2 entire; frontal apotome flat

Kribiodomm

89(31').

Cephalic tubercles present (e.g., Figs. 27.668, 21.610, 27.687)

90

89'.

Cephalic tubercles absent

98

90(89).

Terga 2-6 with median spiniferous processes (Fig. 27.680)

Glyptotendipes (Glyptotendipes)(in part) 91

90'.

Terga 2-6 without such processes

91(90').

Terga 2-5 with coarse shagreen arranged in a fenestrated pattern with many spinules located on pigmented spots of cuticle (Fig. 27.725) Xenochironomus (in part)

91'.

Shagreen not as above

92(91').

Frontal tubercles and cephalic tubercles present (Fig. 27.687)

92 Einfeldia (in part)

92'.

Only cephalic tubercles present

93

93(92').

Segment 5 with 0 or 3 lateral lamellar setae (e.g.. Fig. 27.695) Segment 5 with 4 lateral lamellar setae (e.g.. Fig. 27.707)

94 96

93'.

98'.

Segment 5 with no lateral lamellar setae Polypedilum (in part) Segment with 3 lateral lamellar setae 95 Segment 6 with 3 lamellar lateral setae (e.g.. Fig. 27.721) Pamchironomus (in part) Segment 6 with 4 lamellar lateral setae (Fig. 27.695) Pseudochironomus (in part) Tergum 2 with shagreen area about the same size as that of tergum 3 (Fig. 27.726) Saetheria (in part) Tergum 2 without shagreen 97 Rows of spines on posterior margins of terga 3-6 usually with some spines distinctly larger and darker than the others(Fig. 27.727), or with spines on tergum 6 as large or larger than those on tergum 3(Fig. 27.729) Paracladopelma (in part) Rows of spines on posterior margins of terga 3-6 all of about the same size on any one tergum (Fig. 27.730) but those on tergum 6 distinctly smaller than those on tergum 3(Fig. 27.731) Saetheria (in part) Shagreen on terga as in Fig. 27.725 Xenochironomus (in part) Shagreen on terga not as above 99

99(98').

Sternum 2, at least, with rows of needle-like spines similar to Fig. 27.688

99'.

Sterna without needle-like spines

94(93). 94'.

95(94'). 95'.

96(93'). 96'.

97(96').

97'.

98(89').

Rohackia Parachironomus (in part)

1226

Chapter 27 Chironomidae

Figure 27.708

Figure 27.705 Figure 27.707 Figure 27.706

Figure 27.709

'Vi

Figure 27.712

Figure 27.710

f

Figure 27.713

Figure 27.711

\

Figure 27.717

Figure 27.715

Figure 27.714 Figure 27.716

Figure 27.720

Figure 27.718

Figure 27.719

Figure 27.705 Caudolateral margin of segment 8 (left side) of Chironomus sp. 2. Figure 27.706 Thoracic horn of Chironomus sp. 2. Figure 27.707 Segments 5 and 6 (left side) of Chironomus sp. 2. Figure 27.708 Caudolateral margin of segment 8 (left side) of Einfeldia sp. 2. Figure 27.709 Segment 2 of Parachironomus sp. Figure 27.710 Segments 2 and 3 (left side) of Kiefferulus sp. Figure 27.711 Anal lobes and segment 8 of Paracladopelma sp. 1. Figure 27.712 Caudolateral margin of segment 8 (left side) of "Paracladopeima" sp. Figure 27.713 Caudolateral margin of segment 8 (left side) of Genus 17.

Figure 27.714 Segment 5 of Poiypedilum (Asheum)sp. Figure 27.715 Segment 5 (left side) of Endochironomus sp. Figure 27.716 Caudolateral margin of segment 8 and left anal lobe of Genus 18.

Figure 27.717 Conjunctiva 1/2 and 2/3 of Poiypedilum sp. 4.

Figure 27.TiB Caudolateral margin of segment 8 (left side) of Poiypedilum sp. 5. Figure 27.719 Caudolateral margin of segment 8 (ieft side) of Poiypedilum sp. 6. Figure 27.720 Thoracic horn of Poiypedilum sp. 6.

Chapter 27 Chironomidae

1227

Figure 27.721

Figure 27.723 Figure 27.724 Figure 27.722

m m

Figure 27.725

Figure 27.727

Figure 27.728

Figure 27.726

J Figure 27.729

Figure 27.731 Figure 27.730

Figure 27.721 Segments 5 and 6 (left side) of Polypedilum sp. 6. Figure 27.722 Anal lobe and caudolateral margin of segment 8 (left side) of Graceus sp. Figure 27.723 Caudolateral margin of segment 8(right side) of Nilothauma sp. Figure 27.724 Caudolateral margin of segment 8 (left side) of Stenochironomus sp. Figure 27.725 Segment 5(right half) of Xenochironomus sp.

Figure 27.726 Segments 2 and 3 of Saetheria sp. 1 Figure 27.727 Posterior margin of segment 2 and segment 3 of Paracladopelma sp. 2. Figure 27.728 Posterior margin of segment 2 and segment 3 of Paracladopelma sp. 3. Figure 27.729 Segment 6 of Paracladopelma sp. 3 Figure 27.730 Posterior margin of segment 2 and segment 3 of Saetheria sp. 2 Figure 27.731 Segment 6 of Saetheria sp. 2

1228

Chapter 27 Chironomidae

/'j j'/1 '■

1,1

♦ tA * ' /I M /I A f A

Figure 27.733

B

Figure 27.732

Figure 27.734

Figure 27.738

Figure 27.735

Figure 27.736

Figure 27.739

J

/1\

/

Figure 27.737 Figure 27.744

Figure 27.740

Figure 27.741

Figure 27.742

Figure 27.746

Figure 27.745

h Figure 27.748 Figure 27.747

Figure 27.743 Figure 27.749

Figure 27.732 Dense A. simple, B. bifid or C. multibranched shagreen of abdomen found in many Tanypodinae genera, (after Roback 1981). Figure 27.733 Arching shagreen of abdominal tergites of Trissopelopia sp. Figure 27.734 Thoracic horn of Denopelopia sp. (after Roback and Rutter 1988).

Figure 27.735 Thoracic horn of Meropelopia sp. Figure 27.736 Thoracic horn of Helopelopia sp. Figure 27.737 Branched dorsal setae of abdominal tergites of Monopelopia sp. Figure 27.738 Segment 7 of Fittkauimyia sp. Figure 27.739 Segment 7 of Macropelopia (Bethbilbeckia) sp. (after Fittkau and Murray 1986). Figure 27.740 Thoracic horn of Derotanypus sp. (after Fittkau and Murray 1986). Figure 27.741 Thoracic horn of Euryhapsis sp.

Figure 27.742 Segments 7-8 and anal lobes of Vivacricotopus sp. (after Saether 1988). Figure 27.743 Segments 6-8 and anal lobes of Doncricotopus sp. Figure 27.744 Posterior margins of tergites 3 and 7 of Psectrocladius (Mesopsectrocladius) sp. (after Coffman etal. 1986). Figure 27.745 Thoracic horn of Nanocladlus (Plecopteracoluthus) sp. Figure 27.746 Posteriomedial dorsal setae of middle abdominal segments of Rheosmittia sp. 2. Figure 27.747 Anal lobes of Doithrix sp. 1 (after Coffman etal. 1986). Figure 27.748 Anal lobes of Doithrix sp. 2. Figure 27.749 Apophyses (dark transverse markings) on abdominal segments of Epoicociadius sp. 2.

4

''f

Figure 27.750 Figure 27.753 Figure 27.754

Figure 27.752

Figure 27.751

Figure 27.760

Figure 27.759

Figure 27.755

Figure 27.761

Figure 27.758 _^«fc

»a_

Figure 27.762 Figure 27.764

Figure 27.757 Figure 27.756

Figure 27.765 Figure 27.763 Figure 27.767 Figure 27.766

Figure 27.750 Pad of fiooklets on posterior margin of tergite 2 of Acamptocladius sp. (after Coffman ef al. 1986). Figure 27.751 Anal lobes of Antillocladius sp. Figure 27.752 Segments 8-9 and anal lobes of Camptocladius sp. (after Coffman etal. 1986). Figure 27.753 Anal lobes of Gymnometriocnemus (Raphidocladius) sp. (after Coffman etal. 1986). Figure 27.754 Anterior tfiorax and spur of Syndiamesa sp. (after Oliver 1986). Figure 27.755 Frontal apotome witfi cephalic tubercles and frontal warts of Psilometriocnemus sp. Figure 27.756 Armature of tergite 4 of Tanytarsus sp. Figure 27.757 Anal comb of segment 8 of Tanytarsus sp.

Figure 27.758 Thoracic horn of Micropsectra sp. (after Finder and Reiss 1986). Figure 27.759 Shagreen of tergltes 2 and 3 of Zavreiia sp. (after Finder and ReIss 1986).

Figure 27.760 Thoracic horn of Lauterborniella sp. Figure 27.761 Armature of middle tergltes of Lauterborniella sp. Figure 27.762 Large spines of conjunctiva behind tergltes 3 and 4 of Hyporhygma sp. (after Finder and ReIss 1986). Figure 27.763 Darkened lateral areas of conjunctiva of Phaenopsectra sp. Figure 27.764 Cephalic tubercles and frontal setae of Endotribelos sp. (after Grodhaus 1987). Figure 27.765 Cephalic tubercles and frontal setae of Sergentia sp. (after Finder and Reiss 1986). Figure 27.766 Caudolateral armature of segment 8 of Lipiniella sp. (after Finder and ReIss 1986). Figure 27.767 Anal comb of segment of Goeldichlronomus sp. (after Finder and ReIss 1986).

1229

1230

Chapter 27 Chironomidae

KEY TO SUBFAMILY AND TRIBE FOR ADULT MALE CHIRONOMIDAE

1.

Macropterous, wings fully developed with longitudinal veins and at least some crossveins distinct

2

r.

Brachypterous or subapterous, longitudinal veins and crossveins poorly developed, indistinct or nearly absent

2(1).

Coronal suture and coronal triangle of head absent (Fig. 27.769); antepronotal lobes subtriangular and widely separated (Fig. 27.770); coxa of forelegs enlarged (Fig. 27.770); anepisternal suture reduced (Fig. 27.770); endomeres and aedeagus well-developed (Fig. 27.771); marine or intertidal species TELMATOGETONINAE

2'.

Not with the above combination of characters

3(2').

Wings with at least some distinct, lanceolate setae; ninth tergite large and anteriorly membranous, covering gonocoxites in dorsal view; gonostylus with two separate lobes; crossvein M-Cu occurring in basal 1/3 of wing (e.g., Fig. 27.779) BUCHONOM YIINAE

3'.

Not with the above combination of characters

4

4(3').

Crossvein M-Cu of wing absent (Figs. 27.778, 27.780, 27.782, 27.784)

5

4'. 5(4).

Crossvein M-Cu of wing present(Figs. 27.772, 27.775, 27.776, 27.777, 27.779, 27.781, 27.783) 8 Gonostylus freely articulating with gonocoxite and capable of being retracted so that tip of gonostylus points strongly medially or anteriorly when fully retracted (Figs. 27.774, 27.786); first tarsomere of foreleg always shorter than foretibia ORTHOCLADIINAE (in part)

5'.

Gonostylus broadly fused to tip of gonocoxite and not capable of being retracted strongly medially or anteriorly (Figs. 27.789-27.794); first tarsomere of foreleg usually longer than or subequal in length to foretibia, when distinctly shorter then setae hind tibial comb solidly fused at bases (Fig. 27.802) CHIRONOMINAE(in part)....6

6(5').

Crossvein R-M oriented at a distinct angle to long axis of medial vein (Fig. 27.782); wing usually without dense macrotrichia, but when macrotrichia present squama with a distinct fringe of setae

20

3

7

6'.

Crossvein R-M running parallel to long axis of medial vein and usually also parallel to base of longitudinal vein R4+5(Fig. 27.784); squama always bare; wing usually with dense covering of microtrichia TANYTARSINI(in part)

7(6).

Apex of foretibia with a well-developed, black spur (Fig. 27.797) and spurs at apex tibia of middle leg not fused at base (Fig. 27.801); pars ventralis usually present ventrally between bases of gonocoxites(Fig 27.794) PSEUDOCHIRONOMINl

7'.

Apex of foretibia truncate or at most with a shallow, pale, rounded scale (Fig. 27.798) or hyaline elongation (Fig. 27.799); pars ventralis always absent

CHIRONOMINI(inpart)

8(4').

Longitudinal veins R2 and R3 distinct and not interconnected (Eig. 27.783); fourth tarsomere of all legs cordiform (Fig. 27.800) TANYPODINAE (in part), CLINOTANYPODINI

8'.

Longitudinal veins R2 and Rj either fused through all or most of their length forming longitudinal vein R2+3(Fig. 27.776) or longitudinal vein R2+3 absent(Fig. 27.779) 9 Longitudinal vein R2+3 present 12 Longitudinal vein R2+3 absent 10 Longitudinal vein R4+5 intersecting with the costa near tip of wing and

9(8'). 9'. 10(9').

costal extension well-developed (Eig. 27.775)

10'.

PODONOMINAL .... 11

Longitudinal vein R4_|_5 intersecting with the costa distinctly before tip of wing and costal extension very short or not

present

TANYPODINAE (in part), PENTANEURINI(in part)

Chapter 27 Chironomidae

coronal

inner vertical setae

frontal outer vertical setae

- orbital setae

2^

- postocular setae

Figure 27.768

Figure 27.769

anapleural suture

antepronotum

aedeagus forecoxa

Figure 27.771 Figure 27.770

Figure 27.768 Generalized head of male adult Chironomidae showing setal groups. Figure 27.769 Head of adult Telmatogeton sp.

Figure 27.770 Thorax of adult Telmatogeton sp. Figure 27.771 Hypopygium of Telmatogeton sp.

1231

1232

Chapter 27 Chironomidae

subcosta

costa

R

squama

crossvein M-Cu

Figure 27.772 anepisternum

J.

antepronotum

anapleural suture

forecoxa

preepisternum

Figure 27.773 anal point

gonocoxlte

f'

!

./ii

gonostylus

Figure 27.774

Figure 27.772 Wing of Odontomesa sp. Figure 27.773 Generalized thorax of male adult Chironomidae showing scierites and setae.

Figure 27.774 Hypopygium of Orthocladius {Orthocladius) sp.

Chapter 27 Chironomidae

Figure 27.775

ry

Figure 27.776

ry

Figure 27.777 Figure 27.778

Figure 27.779

Figure 27.780

Figure 27.782

Figure 27.781

Figure 27.784 Figure 27.783

Figure 27.775 Figure 27.776 Figure 27.777 Figure 27.778 Figure 27.779 Figure 27.780

Wing of Boreochlus sp. Wing of Diamesa sp. Wing of Djalmabatista sp. Wing of Cricotopus sp. Wing of Labrundinia sp. Wing of Corynoneura sp.

Figure 27.781 Figure 27.782 Figure 27.783 Figure 27.784

Wing of Tanypus sp. Wing of Axarus sp. Wing of Clinotanypus sp. Wing of Tanytarsus sp.

1233

1234

Chapter 27 Chironomidae

Figure 27.785

Figure 27.786

\i

Figure 27.787

Figure 27.785 Hypopygium of Protanypus sp. Figure 27.786 Hypopygium of Pseudosmittia gracilis.

Figure 27.788

Figure 27.787 Hypopygium of Procladius sp. Figure 27.788 Hypopygium of Denopelopia sp.

Chapter 27 Chironomidae

11(10). 11'. 12(9). 12'. 13(12).

1235

Gonostylus bifurcate, forming two well-developed lobes with long axes running parallel to each other (Fig. 27.796) PODONOMINI Gonostylus elongate and simple or with basal swelling forming a lobe that projects at an angle to the long axis of the gonostylus(Fig. 27.795) BOREOCHLINI Longitudinal vein R2+3 usually distinctly forked distally, if not forked then membrane of wing with dense setae TANYPODINAE (in part).... 13 Longitudinal vein R2+3 never forked; setae, when present, only originating from longitudinal veins and never from wing membrane 17 Crossvein M-Cu intersecting with longitudinal vein Cu before it branches (Figs. 27.777, 27.781) 14

13'.

Crossvein M-Cu intersecting at or distal to branch of Cu (Fig. 27.779)

14(13).

Round or oval tubercle present on scutum (e.g.. Figs. 27.803, 27.804); gonostylus slender and curving through its length, without posterior lobe and usually lacking internal basal lobe (e.g.. Fig. 27.809) TANYPODINI Scutum without a tubercle; gonostylus often with a posterior lobe,(Figs. 27.787, 27.807, 27.808) or internal basal lobe (Figs. 27.805, 27.806) PROCLADIINI

14'. 15(13').

15

Costal extension short, less than length of crossvein R-M,or costa not extended

PENTANEURINI

15'.

Length of costal extension at least as long as length of crossvein R-M

16

16(15').

Vertical and postorbital setae uniserial; anepisternal, preepisternal and dorsal postorbital setae all present (e.g.. Figs. 27.768, 27.773); claws spatulate

16'.

Not with the above combination of characters

17(12').

Crossvein M-Cu intersecting with longitudinal vein Cu before it branches (Fig. 27.772)

17'.

Crossvein M-Cu intersecting at or distal to branch of Cu (Fig. 27.776)

at apex; pulvilli absent

NATARSIINI MACROPELOPIINI

18

DIAMESINAE (in part).... 19

18(17).

Antenna with 5-13 flagellomeres and antennal ratio less than 0.25; plume setae of antenna reduced in number and size; vertex of head broad and consisting of two tubercle-like lobes; frontal and vertical setae present and numerous; scutum with strong pubescence DIAMESINAE (in part), BOREOHEPTAGYIINI

18'.

Flagellum always with 13 flagellomeres and plume setae well-developed; vertex of head simple, broadly rounded to pointed; frontal setae absent and verticals uniserial or partially biserial; scutum without strong pubescence PRODIAMESINAE

19(17').

Gonocoxite strongly extended posteriorly beyond point where gonostylus articulates (Fig. 27.785); thorax with setae on anepisternum, epimeron II and preepisternum

PROTANYPODINI

19'.

Not with the above combination of characters

20(1').

Gonostylus broadly fused to tip of gonocoxite and not capable of being retracted strongly medially or anteriorly (Figs. 27.789-27.794); first tarsomere of foreleg usually longer than or subequal in length to foretibia, when distinctly shorter then setae hind tibial comb solidly fused at bases (Fig. 27.802) CHIRONOMINAE (in part).... 21 Gonostylus freely articulating with gonocoxite and capable of being retracted so that tip of gonostylus points strongly medially or anteriorly when fully retracted (Figs. 27.774, 27.786); first tarsomere of foreleg always shorter than foretibia 22 Gonocoxite with at most superior and inferior volsellae, digitus and median volsella always absent (Figs. 27.789-27.791) CHIRONOMINI(in part) Gonocoxite usually with well-developed superior, median and inferior volsellae and digitus (Fig. 27.792) TANYTARSINI(in part)

20'.

21(20). 21'.

DIAMESINI

1236

Chapter 27 Chironomidae

22(20').

Eyes with dense setae between ommatidia that extend to or beyond the apex of individual ommatids; fourth tarsomere cordiform and shorter than fifth tarsomere. ...DIAMESINI(in part)

22'.

Eyes without dense setae, or if with setae then fourth tarsomere not cordiform and usually longer than fifth tarsomere ORTHOCLADIINAE (in part)

KEYS TO TRIBES OF MARINE CHIRONOMIDAE® Larvae(Marine) 1. Larval head with a ventral median mentum, and striate ventromental plates

(Figs. 27.1, 27.3)

CHIRONOMINAE(go to freshwater key)

r.

Larval head with a distinct mentum, but never with striate ventromental plates

2

2(1').

Larval antenna with 5 segments (Figs. 27.810, 27.811)

3

2'.

Larval antenna with 4 segments (Fig. 27.812)

3(2).

Preanal papillae present, but short, with 5-6 terminal setae and 1-2 lateral setae (Fig. 27.813) ORTHOCLADIINAE—"ORTHOCLADIINI"(in part)(Halocladius)

3.

Preanal papillae absent, replaced by 1-3 short setae or a single long seta (Fig. 27.814)

4(3').

Preanal papillae replaced by 1-3 short setae ORTHOCLADIINAE (in part)(Thalassosmittia, tribe uncertain)

4'.

Preanal papillae replaced by a single(may be branched) stout, long seta (Fig. 27.814) ORTHOCLADIINAE—Clunionini

TELMATOGETONINAE

4

Pupae(Marine) I. Anal lobes with at least a partial fringe of long hairs along edge (e.g., Fig. 27.646); thoracic horn multibranched (Figs. 27.706, 27.720) or simple (Figs. 27.613, 27.618); a sclerotized spine or comb of spines usually present on caudolateral edges of segment 8 (Figs. 27.648, 27.653, 27.704); surface of anal lobes in same plane as abdomen CHIRONOMINAE(go to freshwater key) T. Anal lobes with or without a fringe of hairs, when hairs present, they are short and the surface of the anal lobe is in a plane oblique to that of the abdomen (Fig. 27.815); thoracic horn, when present, always simple; no spines on caudolateral edges of segment 8 2 2(1').

Thorax with a thoracic horn; anal lobes with or without a fringe of hairs, in either case, the surface is oblique to the plane of the abdomen (Fig. 27.815) TFLMATOGFTONINAF

2'.

Thorax without a thoracic horn; anal lobes never with a fringe of hairs; surface of lobes in same plane as abdomen

3

3(2').

Anal lobes each with 3 terminal(or subterminal) setae (Fig. 27.816) ORTHOCLADIINAE-"ORTHOCLADIINI"(in part)(Halocladius)

3'.

Anal lobes without terminal setae, but there may be 1-2 spine-like projections from

the margin (Fig. 27.817)

4(3'). 4'.

4

Dorsal surfaces of segments 2-7(at least) heavily and somewhat uniformly shagreened (rough-surfaced)(Fig. 27.817) Dorsal surfaces of segments 2-7 not heavily and uniformly shagreened, at most with

5

anterior and posterior transverse rows of spines or hooks on some segments ORTHOCLADllNAF—"CLUNIONINI"(in part)(a«n/o)

5.

Segment 8 with only very fine dorsal shagreen; caudolateral corners of segment 9 (anal lobe) with 2 small, closely set spines ORTHOCLADIINAE-"CLUNIONINI"(in part)(Tethymyia)

' The first couplet in each of the life stage keys separates the Chironominae, which has many brackish water species, from the truly marine forms. The genus Halocladius(Orthocladiinae: "Orthocladiini") also is known to occur in saline inland waters.

Chapter 27 Chironomidae

Figure 27.789

Figure 27.791

Figure 27.789 Hypopygium of Glyptotendipes sp. Figure 27.790 Hypopygium of Parachironomus sp.

1237

Figure 27.790

Figure 27.792

Figure 27.791 Hypopygium of Cryptochironomus sp. Figure 27.792 Hypopygium of Tanytarsus sp.

1238

Chapter 27 Chironomidae

Figure 27.793

Figure 27.795

Figure 27.793 Hypopygium of Pseudochironomus sp. Figure 27.794 Internal view of hypopygium of Pseudochironomus sp. showing origin of pars ventralis along medial margin of bases of gonocoxites.

Figure 27.794

Figure 27.796

Figure 27.795 Hypopygium of Paraboreochlus sp. Figure 27.796 Hypopygium of Parochlus kiefferi.

Chapter 27 Chironomidae

1239

Figure 27.797 Figure 27.798

Figure 27.799

Figure 27.800

Figure 27.802

Figure 27.801

Figure 27.805

Figure 27.806 Figure 27.807

Figure 27.803

Figure 27.804

Figure 27.809 Figure 27.808

Figure 27.797 Foretibia of Pseudochironomus sp. Figure 27.798 Foretibia of Chironomus sp. Figure 27.799 Foretibia of Lauterborniella sp. Figure 27.800 Cordiform 4th tarsomere of Coelotanypus sp. Figure 27.801 Middle tibia of Pseudochironomus sp. Figure 27.802 Middle tibia of Chironomus sp. Figure 27.803 Lateral view of tubercle on scutum of Coelotanypus sp.

Figure 27.804 Dorsal view of tubercle on scutum of Coelotanypus sp. Figure 27.805 Gonostylus of Djalmabatista sp. Figure 27.806 Gonostylus of Procladius bellus. Figure 27.807 Gonostylus of Procladius sp.

Figure 27.808 Gonostylus of Procladius sp. Figure 27.809 Gonostylus of Tanypus sp.

1240

Chapter 27 Chironomidae

V "■

y

Figure 27.810

Figure 27.811

Figure 27.810 Five-segmented antenna of larva of Halocladius sp. ("Orthocladilni") (redrawn from Hirvenoja [1973]). Figure 27.811 Five-segmented antenna of larva of Tethymyia sp. (Clunionini) (redrawn from Wirth [1949]).

5'.

Figure 27.812

Figure 27.813

Figure 27.812 Four-segmented antenna of larva of Thalassomya sp. (Telmatogetoninae) (redrawn from Wirth [1947]). Figure 27.813 Preanal papilla of larva of Halocladius sp. ("Orthocladilni") (redrawn from Hirvenoja (1973]).

Segment 8 with heavy dorsal shagreen (as heavy as on segments 2-7) (Fig. 27.817); caudolateral corners of anal lobes with an elongated spine-like projection, another located at base of distal third of the anal lobe ORTHOCLADIINAE (in part) (Thalassosmittia, tribe uncertain)

Adults (Marine) 1. Antenna of male with 11-13 flagellomeres, the terminal one longer, or nearly as long,

as all other segments combined (e.g., Fig. 27.818); all tarsal segments cylindrical; gonostylus of male directed rigidly backward from gonocoxite (Fig. 27.789; compare this to Fig. 27.774) CHIRONOMINAE (go to freshwater key) 1'.

Not with above combination of characters

2

2(1').

Antenna of male with 14 flagellomeres, the terminal one nearly as long as segments 1-13 combined (Fig. 27.818); all tarsal segments cylindrical; gonostylus of male directed medially and anteriorly (Fig. 27.774) ORTHOCLADIINAE—ORTHOCLADIINI (in part) (Halocladius)

2'.

Antenna of male with 6-13 flagellomeres, the terminal one never longer than combined lengths of the preceding 4 (Fig. 27.819); 4th tarsal segment often cordiform (heart-shaped) (Fig. 27.800); 5th tarsal segment often trilobed at apex (Fig. 27.820)

3

3(2').

Antenna of male with 8, 12, or 13 flagellomeres; all tarsal segments cylindrical ORTHOCLADIINAE (in part) (Thalassosmittia, tribe uncertain)

3'.

Antenna of male with 6, 7, or 11 flagellomeres (e.g.. Fig. 27.819); 4th tarsal segment

often cordiform (Fig. 27.800); 5th tarsal segment often trilobed at apex (Fig. 27.820) 4(3'). 4'.

4

Fourth and 5th tarsal segments cylindrical ORTHOCLADIINAE-CLUNIONINI Either 4th tarsal segment cordiform (Fig. 27.800), or 5th tarsal segment trilobed at apex (Fig. 27.820) TELMATOGETONINAE

Chapter 27 Chironomidae

1241

J-LL Figure 27.814

Figure 27.816

Figure 27.815

Figure 27.819

\

■t ' \

.'X

P® < liiS/

'. JmUtlflMimst

/

Figure 27.817

Figure 27.820

Figure 27.818

Figure 27.814 Caudal segments of larva of Clunio sp. (Clunlonini) (redrawn from Lenz [1950]). Figure 27.815 A. Lateral view of terminal abdominal segments of pupa of Thalassomya sp. (Telmatogetonlnae). B. Dorsal view of same. Both redrawn from WIrth (1947). Figure 27.816 Dorsal view of caudal lobes of pupa of Halocladius sp. ("Orthocladiini"). Figure 27.817 Dorsal view of segments 7 and 8 and caudal lobes of pupa of Thalassosmittia sp. (Orthocladiinae).

Figure 27.818 Adult male antenna of Cricotopus sp. (very similar to Halocladius)-, scape and pedicel not drawn; only a few of the plume hairs drawn. Figure 27.819 Adult male antenna of Clunio sp. (Clunlonini); scape not drawn (redrawn from Goetghebuer [1950]). Figure 27.820 Ventral view of trilobed fifth tarsal segment of Teimatogeton sp. (Telmatogetonlnae) (redrawn from Wirth [1949]).

1242

Chapter 27 Chironomidae

ADDITIONAL TAXONOMIC REFERENCES® General Johannsen (1905, 1937); Goetghebuer and Lenz(1936-1950); Townes(1945); Sublette and Sublette (1965); Frommer (1967); Hamilton et al.(1969); Bryce and Hobart (1972); Mason (1973); Fittkau et al.(1976); Hashimoto (1976); Finder (1978); Ssther (1979, 1980); Murray (1980); HofTrichter and Reiss (1981); Oliver(1981); Simpson (1982); Wilson and McGill(1982); Ashe (1983); Wiederholm (1983, 1986, 1989); Oliver et a/.(1990); Armitage et al.(1995). Stether et al.(2000); Saether and Ferrington (2003); Spies and Stether(2004); Andersen et fl/. (2013).

Regional faunas California: Sublette (1960,1964b). Canada: Webb (1969); Oliver et al. (1978); Oliver and Roussel (1983). Carolinas: Webb and Brigham (1982); Epler (2001). Connecticut: Johannsen and Townes (1952). Eastern United States: Roback (1976, 1977, 1978, 1980, 1981, 1985, 1986a, 1986b, 1987). Florida: Beck and Beck (1966, 1969); Epler (1992); Epler (1995); Epler (2014). Kansas: Ferrington (1981, 1982, 1983a,b); Goldhammer eta/.(1992). Louisiana: Sublette (1964a). New York: Johannsen (1905); Simpson and Bode (1980). Northeastern United States: Bode (1990). Pennsylvania (Philadelphia area): Roback (1957). Puerto Rico: Ferrington et al.(1992). Southeastern United States: Beck (1968, 1975); Hudson et al.(1990), Caldwell et al (1997)*; Epler (2001). Upper Midwestern United States - Isle Royale National Park: Egan and Ferrington (2015).

Clunionini: Wirth (1949)-A; Goetghebuer and Lenz(1950)-A; Lenz(1950)-L, A. Diamesinae: Stether (1969)-L, P, A; Hansen and Cook (1976)A; Oliver and Roussel (1982)-L; Makarchenko and Makarchenko(2000)-L, P, A. Orthocladiinae: Brundin (1956)-L, P, A; Ssther (1969, 1975, 1976, 1977, 1981, 1982, 1983b, 1985b)-L, P, A; Hirvenoja (1973)L, P, A; Jackson (1977)-L, P, A; Soponis(1977)-L, P, A; LeSage and Harrison (1980)-L, P; Cranston (1982)-L; Oliver (1982, 1985)-L, P; Simpson et al. 1982-L, P; Bode (1983)-L; Boesel (1983)-A; Oliver and Roussel (1983b)-L, P; Stether and Sublette (1983)-L, P; Cranston and Saether (1986)-L, P, A; Ferrington and Ssther (1987)-P, A; Sffither and Ferrington (1993)-L, P, A; Epler and De La Rosa (1995)-L, P, A; Saether (1995)-A; Saether and Wang (1995)L, P, A; Jacobsen (1998)-L, P, A; Epler et al.(2000)-L, P, A; Hestenes and Saether (2000)-L, P, A; Saether (2005)-L, P, A; Spies(2006); Epler (2010)-P, A; Ferrington and Saether (2011)-L,P, A. Podonominae: Brundin (1966)-L, P, A; Saether (1969)-L, P, A; Coffman et al.(1988)-L, P, A. Prodiamesinae: Saether (1985a)-L, P; Cranston et al.(2011)-L. Pseudochironomini: Saether (1977)-L, P; Jacobsen and Perry (2002)-L, P, A. Tanypodinae: Goetghebuer and Lenz(1939)-A; Fittkau(1962)-P, A; Roback (1971)-A.; Roback (1974, 1985, 1987)-L, P; Roback and Ferrington (I983)-L; Murray and Fittkau (1985)-L, P; Epler (1986)-L; Roback and Rutter (1988)-L, P; Caldwell (1993)-L, P, A; Cheng and Wang (2005)-A; Niitsuma and Watson (2009)-L, P, A. Tanytarsini: Bilyj and Davies(1989)-L, P; Oliver and Dillon (1994)-L, P, A; Reiss(1995)-A; Butler (2000)-A; Caldwell (2000a)-A; Kyerematen et at.(2000)-P, A; Ekrem (2006)L, P, A.

Telmatogetoninae: Wirth (1947)-A.

Taxonomic treatments at the subfamily, tribe, and generic levels(L=larvae; P=pupae;A=adults) Chironomini: Roback (1964)-L, P; Stether (1977, 1983a)-L, P; Borkent (1984)-L, P, A; Boesel (1985)-L, P, A; Reiss and Sublette (1985)-L, P; Epler (1987, 1988a,b)-L, P, A; Grodhaus(1987)-L, P, A; Epler and Ferrington (1994)-L; Caldwell (2000b)-L, P, A; Jacobsen and Perry (2000)-L, P, A; Maschwitz and Cook (2000)-L, P, A; Ekrem et al.

(2003)-L, P, A; Michailova and Ferrington (2016)-L.

' This is a very limited list and includes only publications of general interest and those specifically treating the Nearctic fauna. * Caldwell, B. A.,P. L. Hudson, D. R. Lenat, and D.Smith. 1997. A revised annotated checklist of the Chironomidae (Insecta: Diptera)of the southeastern United States. Trans. Am. Ent. Soc. 123: 1-53.

Taxa

Family

Thalassomya(A)

Telmatogeton (A)

Genus

Habit

Trophic Relationships

builders)

marine (rocks of intertldal)

Marine

Clingers (tube

Beach zone—

marshes, estuaries)

Generally marine (tidal pools, salt

North

4313,4141,4313, 6677, 5041, 5083

409

Coasts

Florida Coast

997,4313, 5287, 6677, 407, 408,

2976, 2977, 3694, 4443, 4599, 5046, 3344, 6201, 6677, 238, 862, 944, 3672, 4704, 4705, 1951, 4423, 5829, 210, 3484, 1204

Southeast

West and

Coasts

East and West

Widespread

American Ecological Distribution SE UM M NW MA* References*

CHIRONOMIDAE

{Continued)

TThe number of species per genus has been estimated in terms of probable range A = 1-5 species; B = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective process based on known North American and European diversities and material in the collections of W.P. Coffman and L.C. Ferrington, Jr.

gatherers, scrapers

Collectors—

gatherers

collectors—

macroalgae),

(chewers—

herbivores

shredders—

Scrapers,

Predators (engulfers and piercers)

filterers (2)

Generally of two Essentially all types Generally of aquatic habitats, borrowers (most types:(1) are tube including marine, collectors— builders) springs, tree holes gatherers and

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasls on trophic relationships

Telmatogetoninae

(=Tendipedidae) (midges)

Chironomidae

Diptera - Non-biting Midges

Order

(number of species in parentheses)!

Tolerance Values

J ) ) ) )) ) ) )) ) ) > ) 1 ) 3 > ) ))

Table 27A Summary of ecological and distributional data for Chironomidae (Diptera). (For definition of terms see Tables 6A-6C; tolerance values are taken from Barbour et al.(1999) and represent either the mean (when the range of values within a region was < 3) or median (when the range of values within a region was > 3); table prepared by M. B. Berg, R. W. Merritt, K. W. Cummins, W. P. Coffman, and L.C. Ferrington, Jr.)

))) I J

Tanypodinae (=Pelopiinae)

Family Tribe

Macropelopiini

FittkauimyiinI

Clinotanypodini

Continued

Fittkauimyia (A) Lotic and lentic

Lentic—littoral

Coelotanypus(A)

borrowers

North

Distribution

American

California

Ostracoda, Chironomidae)

Florida, Kansas

Generally predators Widespread (engulfers and piercers)

Cladocera, Chironomidae)

Predator (engulfers; East Oligochaeta,

Eastern United

States, Canada sooth to

Predators

(engulfers; Oligochaeta,

California

sooth to

Generally predators Eastern United States, Canada (engulfers)

6.9

UM

M

7.0

8.0

NW MA*

Tolerance Values

Ecological

5044

440, 4171, 5049

1932,4171,5049

4423

2976, 4443, 4599, 6677, 461, 862,

References**

)1 )D > J

> )) > I ) ) ) ) ) V ) ) ) ) ) J ) ) )

tThe number of species per genus has been estimated in terms of probable range A = 1-5 species; B = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective process based on known North American and European diversities and material in the collections of W.P. Coffman and L.C, Ferrington, Jr.

Borrowers

Borrowers

Generally

builders)

not tube

Generally lentic—

Lentic—littoral, lotic—depositional

Trophic Relationships

Generally Generally predators Widespread (engulfers and sprawlers— swimmers (very piercers) active predators,

Habit

littoral

and lotic habitats

All types of lentic

Habitat

Clinotanypus(A)

Genus

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Empha5is on trophic relationships

Order

species in parentheses)^

(number of

Taxa

Table 27A

Family

Continued

Tribe

Lotic

Lotic

Lotic

Lotic and lentic

Lotic—erosional.

Apsectrotanypus(A)

Bilyjomyia (A) Brundiniella (A)

Derotanypus(A) Macropelopia (A) lentic—littoral

Lotic

Habitat

Alotanypus(A)

Genus

Borrowers—

Sprawlers

Chironomidae)

Ceratopogonidae,

Protozoa, Cladocera, Ostracoda, Crustacea,

(engulfers;

Predators

Chironomidae)

Tardigrada, Hydracarina,

Cladocera, Ostracoda,

Protozoa,

Predators

(engulfers;

Borrowers—

sprawlers

sprawlers

Chironomidae)

Predators

(engulfers;

Borrowers—

sprawlers

sprawlers

Borrowers—

Habit

Trophic Relationships

North

North

North, West

Widespread

Widespread

Widespread

3.8

0.0

Distribution SB

American UM

6.0

6.0

M NW MA*

Tolerance Values

Ecological

2584,5044, 1615

5044

5044

References**

CHIRONOMIDAE

process based on known North American and European diversities and material in the collections of W.P. Coffman and E.G. Ferrington, Jr. {Continued)

tThe number of species per genus has been estimated in terms of probable range A = 1-5 species; B = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective

*SE = Southeast, DM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

species in parentheses)*

(number of

Taxa

Table 27A

:) ) ))) j ) > ) ) ))) ) > ) ) ) ) :i ))) 3 ) )

K)

Family

Procladiini

Continued

Sprawlers

Lentic—profundal

Prodadius(C) (some littoral). lotic—depositional

Sprawlers

Lotic

Lotic

Radotanypus(A)

Sprawlers

Habit

Djalmabatista (A)

Lotic—depositional

Habitat

Psectrotanypus(B)

Genus

North

North) Protozoa,

gatherers (winter and early instars)

collectors—

Gastrotrlcha),

Ephemeroptera, Ceratopogonldae,

microcrustacea.

Widespread (especially

(engulfers;

Predators

Eastern United States

Predators

West

Widespread

Distribution

American

(engulfers)

Chlronomldae)

Trichoptera,

Protozoa, Cladocera, Ostracoda, Crustacea,

(engulfers;

Predators

Trophic Relationships

9.3

9.0

6.5 9.0

9.0

10.0 10.0 8.3 10.0 10.0

Tolerance Values

Ecological

5712

206, 259, 366, 506, 991, 1073, 1077, 1285, 1329, 1357, 1773, 1932, 3044, 3049, 3045, 3534, 3652,4101, 4120,4129,4171, 4795, 5049, 5989, 6001,440,2559, 6020, 6677, 5309, 5399, 6190, 1615,

5044, 2532

1357,2584,5044

References**

process based on known North American and European diversities and material in the collections of W.P. Coffman and L.C. Eerrington, Jr.

tThe number of species per genus has been estimated In terms of probable range A = 1-5 species; B = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

parentheses)^

(number of species in

Taxa

Table 27A

) ) ) ) ) ) ) I I ) ) ) ) ) ) ) ) ) ) ) ) )) ) ) )

a\

Taxa

Family

Pentaneurini

Natarsiini

Tribe

Denopelopia (A)

Conchapelopia (B)

Arctopelopia (A)

Ablabesmyia (C)

Natarsia (A)

ditch

Lentic—drainage

lentic—littoral

Lotic—erosional,

lentic—littoral

Lotic—erosional,

depositional

lotic—erosional and

Lentic—littoral,

ientic—littoral

North

Widespread

Distribution

American

Mountains

Florida

Predators (engulfers Widespread and piercers) (Chironomidae, Trichoptera, Ephemeroptera)

Rocky Predators

(engulfers)

in stars)

gatherers (early

collectors—

Predators (engulfers Widespread and piercers) (Rotifera, microcrustacea, Chironomidae),

Generally predators Widespread (engulfers and piercers)

Ceratopogonidae)

Cladocera, Ostracoda, Copepoda,

(engulfers;

Predators

Trophic Relationships

1.7

6.4

6.0

8.0

4.3 6.0

4.i

6.0

3.0

8.0

UM M NW MA*

10.0 8.0

SE

To erance Va ues

Ecological

5332

1239

206,991, 1329, 3652,440, 1239, 4101, 5049, 5309, 5519, 2532, 3261

1073

References**

CHIRONOMIDAE

(Continued)

tThe number of species per genus has been estimated in terms of probable range A = 1-5 species; B = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective process based on known North American and European diversities and material in the collections of W.P. Coffman and L.C. Ferrington, Jr.

Sprawlers

Sprawlers

Sprawlers

Generally sprawlers

Generally lotic— erosional and

Sprawlers

Habit

Lotic—erosional

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

species in parentheses)f:

(number of

Continued

J J ) > J ))))))) >> ) J 3 )) ) ) )

Table 27A

I )) >

K»

Family

Continued

Tribe

Lotic

Lotic—erosional

Helopelopia(A)

Hudsonimyia (A)

Lotic—erosional.

Lentic—littoral.

Monopelopia (A)

Lentic—littoral

Primarily North

East

East)

North and

Widespread (especially

Widespread

Distribution

Widespread

Widespread

Widespread

Widespread

and detritus)

depositional

Southeast)

4.6

4.0

8.3

5.2

SE

6.0

6.0

6.0

7.0

UM

6.0

4.9 6.0

2.8 6.0

2.7

7.0

6.0

7.0

6.0

NW MA*

4.3 6.0

3.4

3.9

M

Tolerance Values

Ecological

2532

206, 991, 1073, 1329, 1932, 2560, 4129, 5049, 4602,

5332

2532

5049, 2532

References**

tlhe number of species per genus has been estimated in terms of probable range A = 1-5 species; B = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective process based on known North American and European diversities and material in the collections of W.P. Coffman and L.C. Eerrington, Jr.

collectors—

Predators (engulfers Eastern North America and piercers. Chironomidae), (especially

(engulfers)

Predators

(engulfers)

Predators

Cladocera, Ostracoda)

gatherers (diatoms

Sprawlers

Sprawlers

Sprawlers

Sprawlers

North American

Predators (engulfers Widespread and piercers. Oligochaeta,

(engulfers)

Predators

(engulfers)

Predators

Trophic Relationships

detritus), lotic— erosional and

hydrophytes and

(vascular

Lotic—erosional

Nilotanypus(A) Pentaneura (A)

lotic?

Lotic

lentic—littoral

Meropelopia(A)

Larsia (B)

Sprawlers

Lotic—erosional.

Labrundinia (B) lentic—littoral

Sprawlers

Lotic—erosional

Sprawlers

Sprawlers

Habit

Krenopelopia (A)

(hygropetric)

Lentic—littoral

Habitat

Guttipelopla (A)

Genus

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

parentheses)*

(number of species in

Taxa

Table 27A

) ) ) I )) ) ) ) J ) ) ) ) J ) ) ) ) ) ) ) ) J )) )

00

SO

Family Tribe

Tanypodini

Continued

Generally predators (engulfers and piercers)

lentic—littoral

Lentic

Lotic—erosional,

Xenopelopia (A)

lentic—littoral

Lotic—erosional,

lentic—littoral

Lotic—erosional,

rivers)

Zavrelimyia (B)

Trissopelopia (A)

Thienemannimyia(A)

Lotic—erosional

Telopelopia (A)

(sandy bottom

Lotic—erosional

Habitat

Rheopelopia (A)

Genus

Sprawlers

Sprawlers

Sprawlers

Sprawlers

Habit

North M

California

Primarily North

Ostracoda, Chironomidae)

Widespread

Predators (engulfers Widespread and piercers, Oligochaeta,

Predators (engulfers)

Protozoa, Cladocera, Ostracoda, Chironomidae)

Predators (engulfers Widespread and piercers.

4.0

UM

Predators (engulfers Midwest and piercers)

SE

2.9

American Distribution

Predators (engulfers Widespread and piercers)

Trophic Relationships

6.0

6.0

NW MA*

Tolerance Values

Ecological

2584, 5049, 5519

2584

5332

1073, 1077, 3081, 5049, 440, 5519, 6007, 5062, 5309,

References**

CHIRONOMIDAE OJ 00

(Continued)

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships tlhe number of species per genus has been estimated in terms of probable range A = 1-5 species; B = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective process based on known North American and European diversities and material in the collections of W.P. Coffman and E.G. Ferrington, Jr.

Order

parentheses)t

(number of species In

Taxa

Table 27A

N>

Ut

Podonominae

Family Tribe

Podonomini

Boreochlini

Continued

Habitat

Lentic—littoral

Lasiodlamesa (A)

Lotic—erosional

(springs)

Lotic—erosional

Sprawlers

Lotic—erosional

Trichotanypus(A)

Parochlus(A)

Sprawlers

Lotic—erosional

Sprawlers

Sprawlers

Sprawlers

Paraboreochlus(A)

(Including bogs)

Lotic—erosional

Boreochlus(A)

littoral

erosional, lentic—

Generally lotic—

latitudes)

altitudes and

Generally sprawlers

borrowers (tube builders)

depositional and lentic (at high

Generally

Generally lotic—

Sprawlers

Habit

erosional and

Tanypus(B)i=Pelopia) Lentic—littoral

Genus

North

gatherers, scrapers

Collectors—

gatherers, scrapers Primarily North

North

Kansas

Collectors—

Appalachians, gatherers, scrapers

North

Primarily North

Collectors—

gatherers, scrapers

Collectors—

gatherers, scrapers

Collectors—

gatherers, scrapers

collectors—

Generally

gatherers (diatoms, filamentous green algae, detritus)

collectors—

10.0 8.1

6.0

10.0

1078

6677

862,6677

6001

1357. 1773, 3044, 3049, 3045, 3452, 3534, 3437, 5906,

206, 260, 1074,

American Ecological Distribution SE UM M NW MA* References^

Predators (enguifers Widespread and piercers),

Trophic Relationships

Tolerance Values

> ) ) ) ) ) ) ) 1 > ) ) ) ))) J ) ) )

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships tlhe number of species per genus has been estimated in terms of probable range A = 1-5 species; 8 = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective process based on known North American and European diversities and material in the collections of W.P. Coffman and L.C. Ferrington, Jr.

Order

species in parentheses)^

(number of

Taxa

Table 27A

) )))) )

o

Taxa

Diamesini

Boreoheptagylini

Continued

> j J

erosional

Lotic Lotic—erosional

Lotic?

Diamesa (C)

Lappodiamesa(A)

(cold, fast streams)

Lotic—erosional

(oligotrophic lakes)

Arctodiamesa (A)

Kaluginia (A)

Boreoheptagyia (A)

Sprawlers

Sprawlersclingers

some burrowers

(tube builders)

depositional (cold streams), lentic—

erosional and

Habit

Generally— clingers and

Habitat

Generally lotic

North

North and

Primarily North

Undescribed

Ohio

mountains)

and

Collectors— Widespread gatherers, scrapers? (in uplands

Alaska

unknown

im matures

Alaska,

adults from

7.7

8.0

5.0

5.0

463, 2341, 6331, 462, 4602, 4070

2223, 3081, 3080, 3958, 6677

2976, 4443, 4599, 862, 6677

American Ecological Distribution SE UM M NW MA* References**

gatherers, scrapers? mountains

Collectors—

gatherers, scrapers

collectors—

Generally

Trophic Relationships

Tolerance values

/ I J ))3 ) J )))) ) ) ))) )

{Continued)

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships tlhe number of species per genus has been estimated in terms of probable range A = 1-5 species; B = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective process based on known North American and European diversities and material in the collections of W.P. Coffman and LC. Eerrington, Jr.

Diamesinae

Family

parentheses)^

Order

f

Table 27A

t

(number of species in

>

CHIRONOMIDAE

f

'M

Prodiamesinae

Family

Tribe

Protanypodini

Continued

Lentic—littoral

Pseudodiamesa (A)

Lotic—erosional

Compteromesa(A)

Generally sprawlers

Generally lotic—

Lotic—seeps

erosional

Borrowers?

Sprawlers

Sprawlers

Sprawlers

Sprawlers

Sprawlers

Habit

Lentic—profundal

Lotic—erosional

Syndiamesa (A)

Protanypus(A)

Lotic—erosional

springs)

(small mountain

Sympotthastia (A)

Pseudokiefferiella (A)

Lotic—erosional

Potthastia (B)

(erosional)

Lotic

Habitat

Pagastia (A)

Genus

North

Generally gatherers

collectors—

Southeast

North)

Widespread

(primarily

Collectors—

North

Widespread

West, North

Probably primarily North

Widespread

North

Distribution

American

gatherers

gatherers, scrapers?

Collectors—

gatherers?

Collectors—

gatherers

Collectors—

gatherers, scrapers

Collectors—

Trophic Relationships

5.7

4.7

2.2

SE

2.0

1.0

UM

2.0

6.0

4.0

1.0

M NW MA*

Tolerance Values

Ecological

862

383

3080, 6331

1350, 2223

6417

4070

463, 462,4103,

References**

^

1 ) ) ))) ) ) ) 1 ) ) ) ))

) )) )

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships tThe number of species per genus has been estimated in terms of probable range A = 1-5 species; B = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective process based on known North American and European diversities and material in the collections of W.P. Coffman and L.C. Eerrington, Jr.

Order

species in parentheses)^

(number of

Taxa

Table 27A

(=Hydrobaeninae)

Orthocladiinae

Family Tribe

Clunionini

Continued

Eretmoptera (A)

Clunio(A)

gatherers

depositional

borrowers (tube builders)

Generally

sprawlers

Collectors—

North

builders)

macroalgae)

(chewers—

herbivores

Scrapers, shredders—

Clingers (tube

marine (rocks of intertidal)

West Coast

East and West Coasts

North

gatherers, scrapers

Collectors— gatherers, scrapers

Widespread, particularly

Widespread

Widespread

Widespread

Distribution

American

collectors—

Generally

gatherers?

Beach zone—

Beach zone— Clingers (sand marine (rocky shore) tube builders)

Primarily lotic, but with many lentic representatives (especially oligotrophic lakes)

(detritus)

Collectors—

Lotic—erosional and Burrowers-

Prodiamesa(A)

Sprawlers

Lotic—erosional

Collectors—

gatherers? Odontomesa (A)

Sprawlers

Lotic—erosional

Habit

Monodiamesa (A)

Habitat

Trophic Relationships

7.9

5.9

3,0

4.0

M

3.0

4.0

7.0

NW MA*

Tolerance Values

Ecological

6677

6677

997,4313,4438,

979, 1073, 1077, 1190, 2976, 4443, 4599, 6677, 461, 862, 1204

3452, 3652, 5331

1932, 2943, 2944,

4101

References*

CHIRONOMIDAE

(Continued)

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships tlhe number of species per genus has been estimated in terms of probable range A = 1-5 species; 8 = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective process based on known North American and European diversities and material in the collections of W.P. Coffman and L.C. Ferrington, Jr.

Order

(number of species In parentheses)^

Taxa

Table 27A

j j ; i 3 )) ) ))) ) ) > ) ) > )) ) ) )

Ui is-

Family Tribe

Genus

depositional,

Abiskomyia (A)

Lentic

habits

and lentic habitats

"Metriocnemini"

Aagaardia(A)

Wide range of

lentic—littoral

Lotic—erosional and Sprawlers depositional,

planktonic)

(some instar 1

lentic—littoral

Lotic—depositional (aquatic hydrophytes),

lentic—littoral

Sprawlers

Generally sprawlers

Generally ioticercsional and

groups also

other functional

collectors, but

Generally

gatherers

Collectors—

gatherers

Collectors—

gatherers

collectors—

Generally

macroalgae)

(chewers—

herbivores

Scrapers, shredders—

builders)

(marine rocks of intertidai)

Habit

Clingers (tube

Habitat

Trophic Relationships

Beach zone—

Wide range of lotic

Thienemanniella (B)

Corynoneura (B)

Tethymyia(A)

"Orthocladiini" and

"Corynoneurini"

Continued

North

Far North

Canada

Northern

Widespread

Widespread

West Coast

6.0

1.2

6.0

7.0

3.2 6.0

3.0 7.0

6.0

7.0

2532

440, 1073, 1077, 6417, 612, 4602,

1073, 1077, 1329, 1343, 440, 3652, 5045, 6417, 1239, 4602, 2532

6677

American Ecological Distribution SE UM IVI NW MA* References**

Tolerance Values

»

»

1 ) i ) ) ) I )) ) ) ) ) ) ) ))) ) ) ) 1 )

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships tThe number of species per genus has been estimated in terms of probable range A = 1-5 species; B = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective process based on known North American and European diversities and material in the collections of W.P. Coffman and L.C, Ferrington, Jr.

Order

species In parentheses)^

(number of

Taxa

Table 27A

N>

Ol

Family

Continued

Tribe

Flood plain soils—

Lotic—erosionai

Terrestrial (dung)

Bryophaenodadius(B) Camptodadius(A)

miners).

gatherers

collectors—

Brunswick

Widespread?

Widespread

Widespread

5.2

SB

5.0

UM

5.0

5.0

M NW MA*

Tolerance Values

Ecological

4613, 162, 4447, 4295, 5001, 6417, 2277, §

1073, 1077, 1532,

5228

References**

CHIRONOMIDAE

{Continued)

tThe number of species per genus has been estimated in terms of probable range A = 1-5 species; B = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective process based on known North American and European diversities and materiai in the coiiections of W.P. Coffman and LC. Ferrington, Jr. § Unpublished data, R. H. King, G. M. Ward and K. W. Cummins, Keiiogg Bioiogicai Station.

Sprawiers

(in detritus)

wood, sprawiers (chewers and

(miners) in rotted detritivores

Lotic—erosionai and Borrowers

Brillia (A)

depositionai (detritus)

Manitoba, New

Lentic—littoral

East Kansas

North

North

Distribution

Baeoctenus(A)

Shredders—

macroaigae)

(chewers—

Collectors— gatherers

North

American

Tennessee

seeps

Sprawiers

In colonial algae

Habit

Trophic Relationships

Apometriocnemus(A)

Antillodadius(A)

Allodadius(A)

Lotic—erosionai.

Acricotopus(A) lentic—littoral

Lentic—littoral

Habitat

Acamptocladius(A)

Genus

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atiantic **Emphasis on trophic reiationships

Order

parentheses)*

(number of species in

Taxa

Table 27A

o^

Family

Continued

Tribe

Lotic—erosional

Chaetodadius(B)

Diplodadius(A)

7.7

1.7

8.0

7.0

5.0

5.1 7.0

6.0

2.2 5.0

7.0

Ecological

260, 451, 704,

2277

162, 1192, 3080,

1284, 4448, 6677

References*

Lotic—erosional

Sprawlers

borrowers

Collectors—

gatherers?

6331, 6677, 4602, 5244, 6338, 1239, 1457, 2560, 3055, 3661, 3662, 4379, 6321, 6322, 1799, 5309, 6562, 462, 1232, 1524, 1691, 4339, 4070

3958, 4194, 5580, 6667, 440, 463,

Widespread

Widespread

South Carolina

Widespread

6.2

M NW MA*

sediments, detritus), (miners and tube collectors— lotic—erosional and builders) gatherers (detritus depositional (some and algae) instar 1 planktonic)

hydrophytes (and algal mats,

Shredders—

gatherers

Collectors-

Engulfers Widespread (predators of black fly larvae/pupae and Brachycentrus pupae)

Distribution

1285, 1329, 1343, 2976, 3081, 3652,

builders),

Lentic—vascular

North American

herbivores (miners and chewers),

Clingers (tube

Lotic—seeps

Crkotopus(C)

Sprawlers

makers)

clingers (tube

(loose tube construction),

Borrowers

Habit

Compterosmittia (A)

Chasmatonotus(B)

Lotic—erosional

Habitat

Cardiodadius(A)

Qenus

Trophic Relationships

To erance Va ues

»

*

»

r

> )) ) ) ) ) ) ))) ) ) ) )

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships tlhe number of species per genus has been estimated in terms of probable range A = 1-5 species; B = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective process based on known North American and European diversities and material in the collections of W.P. Coffman and L.C. Ferrington, Jr.

Order

species in parentheses)^

(number of

Taxa

Table 27A

Family

Continued

Tribe

Lotic—erosional,

Gymnometriocnemus Semiaquatic (A) (lentic—margins)

Lotic

Lotic—erosional

Georthodadius(B)

lentic—littoral

Sprawlers

Northwest

North

North, East

Western

Widespread

Widespread

Territories

SE

UM

4.8

7.0

M NW MA*

Ecological

1073, 1077, 1399, 2223,463, 3081, 5774, 5989, 3344, 6321, 6417, 462, 4602, 5309, 4070

References**

CHIRONOMIDAE

process based on known North American and European diversities and material in the collections of W.P. Coffman and L.C. Ferrington, Jr.

NJ

(Continued)

tThe number of species per genus has been estimated in terms of probable range A = 1-5 species: B = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective

Sprawlers

and larvae)

gatherers, scrapers. predators (engulfers of chironomid eggs

Collectors—

insects

parasites on lotic

commensals—

Collectors—

gatherers

Lotic—erosional and

depositional

seeps

Euryhapsis(B)

Eukiefferiella (C)

Epoicodadius(B)

Doncricotopus(A)

Michigan Widespread

Lotic?

Distribution

Lotic—springs and

Habit

North American

Diplosmittia (A)

Habitat

Trophic Relationships

Tolerance Values

j ; )))} )) 7 )))

Doithrix(B)

Genus

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

parentheses)*

(number of species In

Taxa

Table 27A

I

00

Family

Continued

Tribe

Lotic—erosional and

Lapposmittia(A)

Lappokiefferiella (A)

Krenosmittia (B)

Hydrosmittia (A)

Fligh latitude pools

(hyporheic)

Lotic—erosional

lentic—littoral

Lotic—erosional,

profundal

lentic—littoral and

Heterotrissocladius(B) Lotic—erosional,

lentic-littoral

Heterotanytarsus(A)

Collectors—

gatherers

America

from North

Not yet known

unknown

pupae

adults; larvae/

America as

in North

Only recorded

Widespread

Widespread

Widespread

Widespread

9.6

5.4

0.0

SE

0.0

UM

1.0

2.0

6.0

M NW MA*

Tolerance Values

Ecological

568, 4602, 6336

4602

979, 1077, 1350,

1350, 3924, 4120

1790, 3081

4313

References**

>

00

i ) ) J > ) )))))) I

process based on known North American and European diversities and material in the collections of W.P. Coffman and LC. Ferrington, Jr.

tThe number of species per genus has been estimated in terms of probable range A = 1-5 species; B = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective

Sprawlers

gatherers

collectors—

Scrapers,

borrowers

Sprawlers

Collectors—

gatherers scrapers?

Sprawlers,

North

North

gatherers, scrapers

Sprawlers

Northeast Coast

Collectors—

Clingers (sand

Distribution

tube builders)

Lotic—erosional

Hydrobaenus(B)

North American

marine (in Fucus)

Habit

Trophic Relationships

Beach zone—

Habitat

Heleniella (A)

Halocladius(A)

Genus

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

species in parentheses)*

(number of

Taxa

Table 27A

Tribe Lentic—littoral

Limnophyes(B)

Oliveridia (A)

Ninelia (A)

Nanocladius(B)

Borrowers,

Sprawlers?

Sprawlers

Sprawlers

Habit

Lentic

lentic—littoral

Lotic—erosional.

Sprawlers

and depositional sprawlers (detritus), lentic— littoral (oligotrophic) (2 pitcher plants pecies)

Lotic

Lotic—erosional

Mesosmittia (B)

Lotic—erosional

Metriocnemus(B)

Mesocricotopus(A)

Lotic (often

Lopescladius(8) hyporheic)

Semiaquatic?

Lipurometriocnemus (A)

(macroalgae). lotic—depositional

Habitat

Genus

gatherers

Collectors—

gatherers. predators (engulfers)

Collectors—

gatherers

Collectors-

gatherers

Collectors—

gatherers?

Collectors—

Trophic Relationships

North

Arctic, Kansas

Widespread

Widespread

Widespread

North

Widespread

Territory

Yukon

Carolina,

South

Widespread

Distribution

American

4.9

2.2

SE

3.0

4.0

6.0

4.5 3.0

6.0

3.1 8.0

3.0

8.0

UM M NW MA*

Tolerance Values

Ecological

1800

844, 1489, 2532

1073, 1077,6417,

1073, 1285, 3021, 3192, 3452, 6505

2532

2175,4602,6336,

162,4194,6331

References**

(Continued)

tThe number of species per genus has been estimated in terms of probable range A = 1-5 species; B = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective process based on known North American and European diversities and material in the collections of W.P. Coffman and LC. Eerrington, Jr.

CHIRONOMIDAE

K(

Family

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

parentheses)*

(number of species in

Taxa

Table 27A

) ) )) ) ) ) 3

K» 0\

Family

Continued

Tribe

(detritus. diatoms. filamentous

profundal, lotic—

Lotic—erosionai

Lotic—erosionai.

Paracricotopus(A) Parakiefferlella (B)

Paralimnophyes(A)

Lentic—littoral

Paradadius(A)

Lentic—pools

lentic—littoral

Lotic—erosionai

Sprawlers

Sprawlers

Sprawlers

Sprawlers

algae)

borrowers

(erosionai) and erosionai

Sprawlers,

Habit

Lentic—littoral

Lotic—erosionai

Parachaetodadius(A)

Orthocladius(C)

Oreadomyia (A)

Onconeura (A)

Oliveiriella (A)

Habitat

North

gatherers

Collectors—

gatherers

Canada

Alaska, Arctic

Widespread

Widespread

Montana

Collectors—

Wyoming, Collectors—

North

Widespread

Alberta

New Mexico

Arizona and

immatures in

recorded as

Provisionally

Mexico

Kansas, Arizona, New

0.0

6.5

6,0

4.8 6.0

6.0

2.9 6.0

4.0

2.0

440, 4602, 6481, 5309, 6336, 2532

1790

1073, 1350, 2223, 3452, 462, 3958, 4194, 5989, 162, 463, 1192,6417, 2341, 2344, 6321, 4602, 6328, 6338

American Ecological Distribution SE UM M NW MA* References**

gatherers

gatherers

Collectors—

gatherers

Collectors—

Trophic Relationships

Tolerance Values

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships tThe number of species per genus has been estimated in terms of probable range A = 1-5 species; B = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective process based on known North American and European diversities and material in the collections of W.P. Coffman and L.C. Eerrington, Jr.

Order

species in parentheses)^

(number of

Taxa

Table 27A

) ) ) ))) ) ))) )) ) ) ) ) ) ) ) ) ) )) ) ) )

o

K>

0\

Family

Continued

Tribe Habitat

Habit

Collectors—

Lentic—littoral, lotic—depositionai (macroalgae)

herbivores

shredders—

gatherers,

Collectors—

Widespread

Columbia

Lentic—shallow

ponds

Propsilocerus(A)

Psectrodadius(C)

Northern British

Lotic—spring seeps and pools

Plhudsonia (A)

Sprawiers, borrowers

Manitoba shredders—

East

Pennsylvania,

gatherers, herbivores

Tennessee, Collectors—

Sprawiers

Platysmittia (A)

Lotic—springs and

Florida

intermittent streams

Southern

Phytoteimatodadius (A)

gatherers

Lotic—erosional

Parorthodadius(A)

Collectors—

Northeast West

Pennsylvania Lotic

Nova Scotia,

Primarily North

Paratrissodadius(A)

gatherers

Widespread

Distribution

Lotic?

Sprawiers

Collectors—

gatherers

North American

Parasmittia (A)

Paraphaenodadius(B) Lotic—erosional and Sprawiers? depositionai

depositionai

Parametriocnemus(B) Lotic—erosional and Sprawiers

Genus

Trophic Relationships

3.8

SE

8.0

UM

5.7 8.0

6.0

5.0

5.0

8.0

4.0

5.0

M NW MA*

Tolerance Values

Ecological

2653

1073, 1329, 1932. 4194, 612, 2341, 6321, 1799, 5309.

2909

6328

462, 1077, 5309, 4602, 2532

References**

CHIRONOMIDAE 00

UT

(Continued)

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasls on trophic relationships tThe number of species per genus has been estimated in terms of probable range A = 1-5 species; B = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective process based on known North American and European diversities and material in the collections of W.P. Coffman and L.C. Ferrington, Jr.

Order

parentheses)^

(number of species in

Taxa

Table 27A

Family

Continued

Tribe

Lotic—sandy

Rheosmittia (A)

Collectors—

6.0

4.3 6.0

6.0

Ecological References**

Widespread

Lentic—ponds Lotic—streams

Stackelbergina(A) Stictodadius(A)

Mexico

Colorado, Idaho, Montana, New

Quebec

Northern

Tennessee

Lotic—springs?

Widespread

Widespread

4194

5571

6417

1073, 1077, 3452,

1790

7.2

NW MA*

1797, 2532

o

M

Primarily North

Semiaquatic—lentic margins

gatherers

o

SE d UM d

o

Widespread

East

Distribution

Smittia (B)

Collectors—

(engulfers)

(chewers— macroalgae), predators

herbivores

shredders—

gatherers,

Collectors—

gatherers

Collectors—

gatherers

North American

Saethenella (A)

Borrowers?

Sprawlers

Lotic—erosional

Rheocricotopus(B)

substrates

Sprawlers

Lotic

Pseudosmittia (B)

Sprawlers

Habit

Psilometriocnemus(A) Lotic—erosional

Lotic—erosional

Habitat

Pseudorthodadius(B)

Genus

Trophic Relationships

d

o

Tolerance Values

d

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships tThe number of species per genus has been estimated in terms of probable range A = 1-5 species; B = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective process based on known North American and European diversities and material in the collections of W.P. Coffman and L.C. Ferrington, Jr.

Order

parentheses)*

(number of species In

Taxa

Table 27A

3 ) ) ) 3 ) > ) ) ) ) ) )) ) ) ) ) ) ) ) ) ) ) ) )

Ki ON K)

ON

Family

Continued

Tribe

Tokunagaia (A)

Lotic—springs in

Thienemannia (A)

Lotic—erosional

moss

Beach zone— marine (rocks of intertidal)

Thalassosmittia (A) {=Saunderia)

Tempisquitoneura (A) Lotic—erosional

Tavastia (A)

Synorthodadius(A)

Sprawlers

macroalgae),

gatherers

Collectors-

gatherers

collectors—

Widespread

Tennessee

coast

herbivores

(chewers—

Northwest

Pacific

Scrapers, shredders—

builders)

Mountains

Arizona, Rocky

North Carolina

Michigan,

Manitoba,

Widespread

Clingers (tube

gatherers, scrapers?

Collectors—

Parasites

South Carolina

On mayfly and stonefly nymphs

Lotic?

Lotic—erosional

Sublettiella (A)

Symblodadius(A)

Habit

Widespread

Habitat

North

4.7

2.0

2.5 2.0

6.0

5041

4141, 4313, 5083,

3652,6417

1041, 4599, 5045, 6677, 844

American Ecological Distribution SE DM M NW MA* References**

Lotic

Stilocladius(A)

Trophic Relationships

Tolerance Values

CHIRONOMIDAE

(Continued)

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships tlhe number of species per genus has been estimated in terms of probable range A = 1-5 species; B = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective process based on known North American and European diversities and material in the collections of W.P. Coffman and L.C. Ferrington, Jr.

Order

species In parenthe5es)t

(number of

Taxa

Table 27A

Chironominae

Family

Tribe

Chironomini

Continued

Apedilum (A)

Acalcarella (A)

(submerged vegetation)

Widespread

Midwest)

North (perhaps

Widespread

Widespread

Widespread

East

Northwest

Southeast)

upper

Lentic and lotic,

3.9

6.6

East (especially 0.0

Widespread

Pennsylvania

Distribution SE

Lotic—sandy areas

gatherers

North American

of rivers

profundal, lotic— depositional(some instar 1 planktonic)

Generally collectors—

borrowers

littoral and

collectors—filterers

Generally

Generally lentic—

erosional

gatherers and

Generally collectors—

Lentic—littoral, lotic

Zaiutschia (B)

Borrowers in

wood

borrowers and

Lotic—depositional

Xylotopus(A)

clingers

Lotic

Vivacricotopus(A)

gatherers

Collectors—

profundal, lotic— depositional and

Lotic

Unniella (A)

Sprawlers

Generally

Lotic

Habit

Lentic—littoral and

Lotic?

Trichochilus(A)

Habitat

Tvetenia (B)

Genus

Trophic Relationships

2.0

5.0

UM

5.0

7.0

2.0

4.0

5.0

M NW MA*

Tolerance Values

Ecological

275

4443, 4599, 6677

6677

4599, 461, 862,

460, 2977, 4443,

1239

3074, 4454

463, 462, 6331, 4602, 6328

References**

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships tlhe number of species per genus has been estimated in terms of probable range A = 1-5 species; B = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective process based on known North American and European diversities and material in the collections of W.P. Coffman and E.G. Eerrington, Jr.

Order

parentheses)t

(number of species in

Taxa

Table 27A

)) ) ))) > ) ) ) ) ))) ) ) ) ) ) ) ) ) ) ) ) )

OV

ON IJl

Family

Continued

North

Burrowers (tube

builders)

Lentic—littoral and

profundal, lotic— depositional

Widespread

9,8

1285, 1325, 1329, 1343, 1773, 1932, 2596, 2674, 2977, 3018, 3048, 3259, 3452, 3533, 3534, 3944, 3957, 4194, 4795, 4830, 4927, 5008, 5088, 5989, 6290, 6294, 567, 1679, 2979, 6677, 3011, 3012, 3015, 3508, 4876, 4668, 462, 3623, 6368, 1799, 5277, 6417, 4437, 6338, 276, 990, 1524, 2233, 2231, 2232, 2532, 2890, 3102, 4719, 5712, 6207, 6780

10.0 8.1 10.0 10.0 451, 654, 1074,

1791, 6420

CHIRONOMIDAE

{Continued)

4The number of species per genus has been estimated in terms of probable range A = 1-5 species; 8 = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective process based on known North American and European diversities and material in the collections of W.P. Coffman and LC. Ferrington, Jr.

herbivores (miners)

shredders—

fiiterers),

gatherers (a few

Widespread

Lotic—sandy areas

Chernovskiia (A)

Chironomus(C) {=Tendipes)

West

Lotic

Texas, Florida

Widespread

Lentic

of rivers

Tolerance Values

American Ecological Distribution SB UM M NW MA* References^

Beardius(A)

Collectors—

Collectors—

gatherers

burrowers

Lotic—depositional, Sprawlers, ientic

Habit

Trophic Relationships

Beckidia (A)

Axarus(A)

Habitat

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

(number of species in parentheses)f:

Taxa

Table 27A

Family

Continued

Tribe

Demijerea (A) Lentic

profundal

Lentic—littoral and

Burrowers

gatherers

Collectors—

Collectors—

gatherers

Collectors—

gatherers

Collectors—

Widespread

piercers of Oligochaeta)

Chironomidae and

Einfeldia (B)

Burrowers

North

Widespread

Distribution

American

Widespread

Widespread

Widespread

Widespread

Widespread

Predators (engulfers Widespread of Protozoa, microcrustacea,

gatherers and filterers, scrapers?

Lentic—littoral

Burrowers

Collectors—

gatherers

(=Limnochironomus) (wide range of microhabitats)

Dicrotendipes(C)

(A)

Demicryptochironomus Lotic—depositional

Burrowers

Lentic and lotic

Cyphomella (A) (sandy rivers)

Sprawiers

Lentic—littoral, lotic—depositional

Cryptotendipes (B)

Burrowers

Sprawiers, burrowers

Lentic—littoral

Oadopelma (B)

Habit

Cryptochironomus(C) Lentic—littoral and profundal, lotic— depositional

Habitat

Genus

Trophic Relationships

9.1

2.1

6.1

6.7

2.5

5.0

6.0

8.0

9.0

7.0

NW MA*

8.0

6.0

3.0

5.9 8.0 8.0

4.2

4.9 8.0 8.0

M

Tolerance Values

Ecological

1324

5309

90, 207, 1073, 1077, 3652, 4120, 4194, 4409, 4795,

440

5309, 6321

206, 1285, 1329, 1611, 3044, 3049, 3534, 3652,4101, 4171, 4194, 5955, 5989, 440, 5331, 6001, 6677, 4437,

4101,4120,6001

References**

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships 4The number of species per genus has been estimated in terms of probable range A = 1-5 species; B = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective process based on known North American and European diversities and material in the collections of W.P. Coffman and L.C. Ferrington, Jr.

Order

parentheses)*

(number of species In

Taxa

Table 27A

) ) ) )))J )) ) )))) > ) ) ) ) ) ) ) ) ) ) )

0\ On

a\

Tribe

Lentic—littoral and

Glyptotendipes(B)

Graceus(A)

Goeldichironomus(B)

Lotic (sandy rivers)

Gillotia (A)

Borrowers

Burrowers

Shredders—

gatherers

Collectors—

and gatherers

collectors—filterers

macroalgae),

and chewers—

herbivores (miners

Shredders—

Burrowers

Lentic—littoral, lotic Sprawlers

stagnant ponds)

Lentic (small

gatherers

Collectors-

(miners and tube herbivores (miners profundal, lotic— depositional(rarely) builders), clingers and chewers— (net spinners) filamentous algae), (some instar 1 planktonic) collectors—filterers and gatherers

Lotic(sandy rivers)

Fissimentum (A)

builders)

Lentic

Clingers (tube

Lentic—littoral

Habit

(algal mats) and profundal (instar 1 planktonic)

Habitat

Endotribelos(A)

Endochironomus(A)

Genus

Trophic Relationships

North

Ontario

Widespread

Widespread

Midwest

Puerto Rico

Carolina,

South

Florida

California,

Widespread

Distribution

American

10.0

3.5

M

10.0 4.7

UM

8.0

10.0

10.0 10.0

NW MA*

Tolerance Values

Ecological

506, 1799

3090

451,803, 1285, 1329, 1343, 1773, 3048, 3143, 3652, 4101, 4120, 4194, 4795, 4437, 4830, 5989, 6294, 567, 4876, 4878, 5309, 6129, 1524, 2005,

2532, 2890

6129

1309, 1343, 2871, 4101, 3055, 3268, 4194, 6294, 4437,

References**

process based on known North American and European diversities and material in the collections of W.P. Coffman and LC. Ferrington, Jr. (Continued)

tlhe number of species per genus has been estimated in terms of probable range A = 1-5 species; B = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective

CHIRONOMIDAE

-j

Family

Continued

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

(number of species in parentheses)^

Taxa

Table 27A

Ki

OS

Family

Continued

Carolina,

profundal

Lentic (sandy littoral) Burrowers Lentic—littoral, lotic—depositional

Microchironomus(A)

Microtendipes(B)

Collectors—filterers Widespread and gatherers

Widespread

Territories

Northwest

Widespread

United States

Southeast

8.0

3.5

6.2

7,0

2.4 6.0

10.0 10.0 5.2

7.5

10.0

6294, 6321

451, 1073, 1077, 4194, 4437, 4795,

1350

4236, 4497, 6294, 6677, 2255

1074, 1773, 3944, 4101,440, 4171, 5309, 5331

process based on known North American and European diversities and material in the collections of W.P. Coffman and LC. Ferrington, Jr.

tlhe number of species per genus has been estimated in terms of probable range A = 1-5 species; B = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective

**Emphasis on trophic relationships

Clingers (net spinners)

Lentic

Lipiniella (A)

tube builders)

(portable sand

Collector— gatherers

gatherers

sprawlers— clingers,

Lentic—littoral and

Lauterborniella (A)

burrowers

Collectors—

Climbers—

Lotic

Kribiodorum (A)

Kansas

South

Widespread

Lotic (sandy

gatherers substrates)

petioles) Collectors—

Kloosia (A)

Burrowers

in stems and

Shredders (miners

Widespread

Lentic

hydrophytes

Burrowers

gatherers, scrapers?

Lentic —vascular

Collectors—

clingers

hydrophytes (submerged zone)

North

Tolerance Values

American Ecological Distribution SE UM M NW MA* References*

Kiefferulus(A)

Hyporhygma (A)

Harnischia (A)

Habit Climbers—

Habitat

Trophic Relationships

Lentic—vascular

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic

Order

species in parentheses)^

(number of

Taxa

Table 27A

) ) ) )))))) ) ) ) )) ) > ) ) ) ) ) ) ))))

00

On

Family

Continued

Tribe

Lentic—littoral, lotic—depositional

Paradadopelma (B)

collectors— fllterers?)

gatherers(and

Scrapers, builders)

Lentic—littoral

Phaenopsectra (B)

Collectors— gatherers

Cllngers (tube

Lotic—depositional. Borrowers (tube lentic—littoral builders)

7.0

7.6

Widespread

7.0

5.3 8.0

5.2

Widespread

Widespread

Cllngers (tube builders on plants)

Collectors— gatherers

Widespread

Sprawlers

gatherers, parasites

collectors—

8.0

4.6 7.0

7.0

5.0 8.0 8.0

1073, 1077, 1329, 3652, 5309, 2532

1789, 1797, 6323, 6324, 2175

4795, 1524

3021, 3493, 3652,

Mollusca)

10.0 5.3

4101,4194, 4437,

8.6

Sprawlers(some Predators parasites in (engulfers),

Paratendipes(B)

hydrophytes

Paralauterborniella (A) Lentic—vascular

Lentic—littoral

Parachironomus(C)

Widespread

1580

profundal

2532

2.6

2532

Widespread

2.0

Widespread

3.1

Ecological References*

Lentic—littoral and

2.0

5.5

Widespread

M NW MA*

Omisus(A)

UM

Distribution

Pagastiella (A)

Habit

Lotic—depositional

Habitat

North American

Nilothauma (A)

Genus

Trophic Relationships

Tolerance Values

CHIRONOMIDAE

{Continued)

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships tlhe number of species per genus has been estimated in terms of probable range A = 1-5 species; B = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective process based on known North American and European diversities and material in the collections of W.P. Coffman and L.C. Ferrlngton, Jr.

Order

parentheses)*

(number of species in

Taxa

Table 27A

Family

Continued

Tribe

Stictochironomus(B)

(miners)

hydrophytes,

Lotic—depositional Borrowers(tube (organic sediments) makers)

lotic—^wood

Borrowers

Lentic Lentic—vascular

Borrowers

Borrowers

Sergentia (A)

(sandy bottom)

Lentic and lotic

(sandy bottom)

Lentic and lotic

Climbers,

clingers

Lentic—^vascular

hydrophytes (floating zone)

Habit

Stenochironomus(B)

Saetheria (A)

Robackia (A)

Polypedilum (C)

Habitat

North

North

Widespread

Widespread

Widespread

Distribution

American

herbivores (miners)

shredders—

gatherers,

Collectors—

Widespread

Collectors— Widespread gatherers shredders (wood gougers)

gatherers

Collectors—

gatherers

Collectors—

(engulfers)

filterers?), predators

gatherers (and

collectors—

herbivores (miners),

Shredders—

Trophic Relationships

6.7

6.4

8.1

3.3

6.7

9.0

5.0

4.0

4.0

3.6

7.0

5.0

6.0

NW MA* 3.1 6.0

M

Tolerance Values

Ecological

3164

1285, 2655, 3144, 4101, 462, 5309,

2890

162, 1532, 4613, 6677, 440, 589, 1796,4409,2532,

4172

4602

440, 5571, 6481,

1285, 3047, 3048, 3944,4120,4121, 4171,4194, 5989, 162, 6001, 6294, 6677, 440, 463, 3237, 5331, 6129, 6321, 6481, 462, 5309, 6328, 2532, 2890, 5712

451, 1073, 1077,

References*

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships 4The number of species per genus has been estimated in terms of probable range A = 1-5 species; B = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective process based on known North American and European diversities and material in the collections of W.P. Coffman and L.C. Ferrington, Jr.

Order

parentheses)*

(number of species In

Taxa

Table 27A

) ) ) ))) ))) ) 1 )3 :) 1 1 1 ) 1 ) ) ) ) )))

o

Tanytarsini (=Calopsectrini)

Pseudochironomini

CHIRONOMIDAE

K)

Family

Continued

Lentic (seasonal pools)

erosional (in algae) and depositional

Caladomyia (A)

Collectors—

and gatherers

borrowers or

clingers (tube builders)

depositional, lentic—littoral Lentic

Generally collectors—filterers

Generally

gatherers

Collectors—

gatherers

Generally lotic—

Borrowers

Borrowers

North

Widespread

Western

Distribution

American

California

Southern

Oklahoma,

Texas,

Widespread

Widespread

Florida

Southern

Widespread

South

Predators (engulfers Widespread of sponges)

erosional and

and depositional

Pseudochironomus(B) Lentic and lotic— erosional (in algae)

Manoa(A)

Lentic

Zavreliella (A) Lentic and lotic—

Lotic and lentic?

Xestochironomus(A)

Borrowers (in

sponges)

Lotic (and lentic) in sponges

(depositional) Xenochironomus(A)

miners)

Lentic and lotic

gatherers

Borrowers(wood Collectors—

Lentic and slow lotic

Habit

Tribelos(A)

Habitat

Trophic Relationships

Synendatendipes(A)

Genus

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships

Order

(number of species in parentheses)!:

Taxa

Table 27A

4.2

7.0

6.6

SE

5.0

0.0

5.0

UM

5.0

NW MA*

4.7 5.0

5.3

M

Tolerance Values

Ecological

6677

(Continued)

4443, 4599, 6226,

1285,4194,4798, 6331, 2229

2911

1285,4194

440

440,6129,2532

References**

Family

Continued

Tribe

Lentic—littoral

Micropsectra(C)

Lotic—erosional

Lotic—erosional,

Paratanytarsus(B) lentic—littoral

Lotic—erosional

(springs) Neozavrelia (A)

Neostempellina (A)

Lentic—littoral

Corynocera (A)

Sprawlers

(Including brackish), sprawlers lotic—depositional

Lotic—erosional

gatherers

Collectors—

Collectors—

gatherers and filterers

Widespread

States

Eastern United

East

Widespread

Canada

Northwest

Mountains),

Wyoming (Rocky

Widespread

Widespread

Distribution

Relationships

Lentic—vascular

Climbers,

Habit

hydrophytes, lotic— depositional

Habitat

Constempellina (A)

Cladotanytarsus(B)

Genus

North

American

Trophic

7.7

1.4

6.0

7.0

4.2 6.0

3.5 7.0

6.0

6.0

7.0

NW MA* 4.4 7.0

M

Tolerance Values

Ecological

6417

1239, 2230, 2560,

4444, 6294, 4602, 6417, 2277

1073, 1077, 1329, 3081, 3958, 4194,

1792

206, 1329, 4194, 4795, 440, 463, 462, 1789

References**

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships tlhe number of species per genus has been estimated in terms of probable range A = 1-5 species; B = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective process based on known North American and European diversities and material in the collections of W.P. Coffman and L.C. Ferrington, Jr.

Order

species in parentheses)^

(number of

Taxa

Table 27A

) ) ) ) )) )) ) ) ) ) ) ) ) ) ) ) > ) ) ) ) ) ) )

N>

N>

N>

Family

Continued

Tribe

Lotic—erosional,

clingers (net spinners)

hydrophytes (floating zone) and profundal, lotic— instar 1 planktonic)

erosional(some

Climbers,

Lotic Lentic—vascular

Sprawlers

Tanytarsus(C) (=Calopsectra)

lentic—littoral

mineral tube

Subletted (A)

Stempellinella (A)

Western

2.0

6.0

1.7

6.7

6.0

6.0

3.5 6.0

2.2 6.0 6.0

2.6 4.0 4.0

2.0

3.3 6.0

UM M NW MA*

5.3 4.0

2.0

Widespread

Widespread

Widespread

Canada

6.4

Distribution SE

Collectors—filterers Widespread and gatherers, a few scrapers

Collectors—

gatherers (detritus, algae)

Climbers—

sprawlers— clingers (portable,

Lotic—erosional, lentic—littoral

Stempellina (B)

builders)

North American

Collectors—filterers Widespread

Lotic?

and net builders) Skutzia (A)

Clingers (tube

Habit

Lotic—erosional

Habitat

Rheotanytarsus(B)

Genus

Trophic Relationships

Tolerance Values

Ecological

2890

206, 260, 451, 949, 1073, 1077, 1285, 1329, 1343, 1932, 3048, 3452, 3533, 3944, 3958, 4120,4194, 4795, 440, 463, 462, 6294, 2655, 6321, 6368, 6417, 1579, 2277, 2532, 2855,

2532

6294

206, 1073, 1077,

440, 1073, 1077, 6294, 463, 462, 3330, 2532, 6492

References**

CHIRONOMIDAE

(Continued)

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships flhe number of species per genus has been estimated in terms of probable range A = 1-5 species: B = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective process based on known North American and European diversities and material in the collections of W.P. Coffman and L.C. Ferrington, Jr.

Order

species in parentheses)^

(number of

Taxa

Table 27A

Family

Continued

Tribe

Zavrelia (A)

Virgatanytarsus(A)

Genus

Lotic

macrophytes) Collectors—

sprawlers— gatherers clingers (portable, mineral tube builders)

Widespread

United States

littoral (on

Southeast

Distribution

Lotic—erosional

Climbers—

Habit

North American

(springs), lentic—

Habitat

Trophic Relationships

2.7

SE

UM

M

8.0

NW MA*

Tolerance Values

Ecological

6294

979, 1073, 1077,

References**

*SE = Southeast, UM = Upper Midwest, M = Midwest, NW = Northwest, MA = Mid-Atlantic **Emphasis on trophic relationships tThe number of species per genus has been estimated in terms of probable range A = 1-5 species; B = 6-19 species; C = 20 or more species. The range for any genus was derived by a subjective process based on known North American and European diversities and material in the collections of W.P. Coffman and L.C. Ferrington, Jr.

Order

parentheses)*

(number of species In

Taxa

Table 27A

) ) ) ) ))) ))) ) ) ) ) ) > ) ) )) ) ) ) ) ) )

to

#2«>'

GLOSSARY OF MORPHOLOGICAL TERMS Brett W. Merritt

Monta^nola (Ticino), Switzerland

A—prefix meaning wanting or without. Ab—prefix meaning off, away from. Abdomen—third or posterior division of insect body. Abdominal—^pertaining to the abdomen. Accessory—secondary, adjoined to any primary structure.

Accessory blade of mandible—blade-like appendage arising on the inner margin of mandible. Accessory glands—any secondary gland of a glandular system. Acuminate—tapering to a long point. Acute angulate—angle smaller than a right angle. Adfrontal area—areas between the ecdysial lines and the adfrontal sutures.

Adfrontal suture—sutures separating adfrontal areas from frontoclypeus. Aedeagus—terminal part of male reproductive organ.

Aenescent—brassy coloring. Air sacs—expanded, thin-walled areas in the tracheae.

Air straps—pair of strap-like appendages derived from abdominal segment VIII. Alula(e)—lobed membranous basal portion of posterior wing margin. Ampbipneustic—open, functional spiracles. Anal—last segment of the abdomen.

Anal angle—distinct angle between inner and outer margins of any wing. Anal area—posterior or anal part of a wing supported by anal veins; axillary area. Anal cells—spaces between anal veins. Anal claw—sclerotized hook at the end of the anal

proleg. Anal crossing—(Ac) where anal vein branches posteriorly.

Anal crossvein—secondary branches of cubitus anterior. Anal fan—fan-like extension of the anal area of the

hind wing in insects. Anal foot—posterior end of larval body modified to serve as a holdfast.

Anal loop—wing area including a few to several cells between branches of anal vein, or between cubitus and first anal vein.

Anal margin—line extending along lower edge of wing. Anal proleg—any process or appendage pertaining or attached to the anus or to the last segment of the abdomen that serves the purpose of a leg. Anal region—posterior area of wing or terminal end of insect.

Anal setae—one or more prominent setae of anal lobes.

Anal spine—enlarged setae, usually blunt and conical.

Anal style—long, thin appendage on the terminal segment of abdomen. Anal triangle—well-marked triangular area bounded anteriorly by vein A' and distally by vein A3. Anal tubercle—tubercle bearing anal organs; anal papilla. Anal tubules—appendages of the anal segment above, between, or at the basal margin of posterior prolegs. Anal vein—(A)longitudinal unbranched veins that extend from base ofinsect wing to outer margin below cubitus.

Anapleurite—sclerotized area above coxa (supracoxal area)in a generalized thoracic pleuron. Anastomose—an opening; coming together; merging or running into each other.

1275

1276

Glossary

Anastomosed crossvein—crisscross-like network of crossveins.

Ancillary—auxiliary; something that helps. Ancipital—flattened and with two opposite edges or angles. Anellus(i)—one or more ring-like structures at base of antenna.

Anepisternum—division of the episternum into two parts by a suture. Angular—forming an angle. Annular—ring-shaped structure. Annulation—areas of reduced sclerotization giving the appearance of concentric rings or annuli on certain antennal segments. Annulus—ring encircling a joint. Anteapical—before apex. Anteclypeus—inferior half of the clypeus. Antecosta—anterior submarginal or marginal ridge on inner surface or a tergal or sternal sclerite corresponding to primary intersegmental fold, and to which longitudinal muscles are typically attached.

Antehumeral—pertaining or relating to space immediately anterior of wing base. Antehumeral stripe—between the middorsal carina and the humeral stripes. Antemedian—pertaining to bristles or conspicuous large setae positioned anterior of middle of a segment or sclerite. Antenna(e)—paired segmental sensory organs, on each side of head.

Antenna(l) groove—groove to receive and conceal antenna.

Antenna(l)tubercle—elevation on the head to which each antenna is attached.

Antennal sheath—structure enclosing an antenna. Antenodal crossvein—crossveins extending between the costa and the subcosta.

Antenodal setae—setae found before or preceding a node.

Antepenultimate—the second before the last. Antepronotum—anterior division of the pronotum. Anterad—contraction of anteriad meaning toward the front; in the direction of the anterior. Anterior—in front, before; pertaining to a structure or color located in front of midline.

Anterior lahral region—anterior portion of the labrum.

Anterior lamina—anterior margin of the genital pocket.

Anterior prologs—locomotory structures originating from ventral surface of first thoracic segment. Anterodorsal—toward the front and dorsum.

Anterolateral—located anteriorly and to the side.

Anteromedian—in front and along the midline. Anteromesal—in the front and along the midline of a body. Anteroventral—^in the front on the lower(or under)side. Antrorse—directed toward front; directed forward or upward.

Anus—at apex of abdomen. Aperture—opening, hole, or gap. Apex—end or tip of insect or appendage. Apical—pertaining td the apex.

Apical dissections—divided distally into several branches or strahds.

Apicolateral—located apically and to the side. Apneustic—tracheal system with no functional spiracles.

Apotome—subdivision divided by a membranous fold. Appendage—structure attached by a joint to a larger structure.

Appendicules—with an appendage or appendages. Appressed—closely applied to. Apterous—without wings; a wingless condition. Apterygote—primitive wingless insects. Arcuate—arched.

Arculus—(a)crossvein giving rise to the media. Arista—reduced flagellomeres of antenna following first flagellomere of antenna. Armature—setae, spines or sclerotized processes on insect.

Article—particular item or object. Articulation—joining or hinging of two or more structures.

Ascending—recurved.

Asperate—small spine-like processes or roughened areas.

Asymmetrical—not of symmetrical form. Atrium—any chamber at the entrance of a body opening. Attenuated—tapering to a point. Auricles—ear-like projection on abdomen. Axillary—placed in the crotch or angle of origin of two bodies; arising from the angle of ramification.

Axillary region—region of the wing base containing the axillary sclerites.

Axillary vein—one or two longitudinal veins toward the inner margin from the anal vein. Band—transverse mark wider than a line.

Basal—at or pertaining to the base or point of attachment or located nearest the body. Basal median cell—cell between media and cubitus

and proximal to basal medial or medial-cubital crossvein.

Base—appendage part nearest the body. Base of wing—same as wing base.

Glossary

Basisternum—anterior of the two sternal skeletal

plates.

Basistyle—male genitalia. Basitarsus—basal tarsomere.

Beak—protruding mouthparts forming the mouth. Bi prefix meaning two. Bicornute—two horned. Bidentate—with two teeth. Bidented—two teeth.

Bifid—forked, cleft, divided into two parts. Biforous—having two openings. Bifurcate—divided into two branches or forked.

Bilamellate—having two plates. Bilobed—with two lobes.

Binodose—two swellings. Biordina!—two alternating lengths in a single row. Biramal—pertaining to two branches. Blade—thin, flat elongate structure. Blood gills—hollow, nontracheated, usually filamentous respiratory processes. Blotch—irregular marking. Blunt—not sharp.

Bothriotricha—long, slender fringe setae. Brace vein—slanting crossvein. Brachypterous—shortened wings. Branched—divided; division.

Breathing tubes—same as siphon. Bridge—secondary longitudinal vein connecting the radial sector with M1+M2.

1277

Cardo—basal division of maxilla.

Carina(e)—elevated ridge or keel. Carinate—having carinae or keeled. Cataclystiform—wing pattern characterized by the presence of marginal and submarginal lines parallel to the termen of the forewing, a postmedial line parallel to the termen from the costal margin to CUA2, bending inward between CuA2 and 1 A,then immediately bending downwards to the inner margin, and a series of submarginal black dots along the termen of the hindwing. Caudad—toward the posterior end of abdomen. Caudal—pertaining to the posterior end of abdomen.

Caudally—turned backwards. Caudolateral armature—spine, comb,compound spur on each caudolateral margin of abdominal segment 8.

Cavernicolous—cave inhabiting. Cephalad—toward the head. Cephalic—belonging to or attached to the head. Cephalic fans—analogous to the labral fans of larval black flies (Simuliidae). Cepbalopharyngeal skeleton—internal sclerotized portion of head, prominent in Diptera larvae. Cepbalotboracic suture—seam or impressed line dividing a united or fused head and thorax. Cephalotborax—fused head and thorax.

Bridge crossvein—one or more crossveins. Bristles—stiff, usually short and blunt setae or hair spinules. Bronzed—yellowish brown or reddish brown in color.

Cercus(i)—terminal paired lateral slender appendage. Cervical—pertaining to the neck area. Cervical gill—gills located between head and thorax

Brushes—flexible outgrowths or projections (e.g, of hairs or setae). Bulge—a rounded swelling which distorts an

Cervix—membranous neck region. Chaetoesema(ta)—sensory organs on head. Chaetotaxy—arrangement and nomenclature of

otherwise flat surface.

Bulla(e)—slightly swollen part of the costal area of the wing toward the tip, with more crossveins, practically equivalent to the stigma. Burrowing books—tips of the front tibiae of some Anisoptera that are more or less flattened and hooked for digging. Calcipala—apical process. Callosity—flattened elevation or hardened hump. Callus(i)—hard lump or swelling of the cuticle. Calypter(es)—two basal lobes at the posterior margin of the wing. Campaniform sensillum—sense organ of the antenna or maxillary palp having the appearance of a circular structure.

Capitate—rounded knob at apex. Cardinal beard—elongate setae arising in definite row underneath ventromental plate.

embedded in cervix.

setae or bristles.

Cbalaza(ae)—elevated pigmented area set with setae. Chelate—pincer-like structure. Chitinous rings—hardened rings. Chloride epithelium—ovoid areas on abdomen. Cilia—thin, long hairs. Ciliate—provided with a parallel row of cilia. Claspers—styli. Claval suture—suture of the forewing. Clavate—clubbed. Claviola—antennal club.

Clavopruina—narrow, white frosted area along the anterior lateral margin of the clavus. Clavus(i)—area behind the claval furrow.

Claws—sharp, hooked structure on the apex of leg. Cleft—split or fork. Closed procoxal cavity—cavity surround by sternal sclerites.

1278

Glossary

Clothing hair—hair covering the body. Club—thickened or enlarged segment at tip of antenna.

Cluster of setae—setae set close together. Clypeus—sclerite at the base of the labrum.

Corrugations—alternate ridges and channels. Costa(e)—first longitudinal vein; vein extending along the anterior margin of the wing from base to the point ofjunction with subcosta.

Coalesced—unite or combine into one mass.

Costal crossvein—those veins that extend between the costa and the subcosta.

Coarctate—concealed into hardened capsule.

Costal fracture—short, transverse line of weakness in

Collar—band at the neck.

the forewing. Costal margin—anterior margin of wing. Costal vein—(C) vein running close to and parallel with the costal margin, extending from base to the margin before the apex. Coxa(e)—basal segment of leg. Coxal—of or pertaining to the coxa. Coxal cavity—opening in the underside of the thorax in which the leg is attached. Crease—line or ridge. Creeping welts—transverse swollen areas that bear

Collophore—ventral tube. Collum—neck or collar.

Comb plate—lateral sclerite with comb scales. Comb scales—specialized spicules. Commissure—junction formed between two structures.

Compound—made up of similar parts. Compound eye—aggregation of separate visual elements known as ommatidia.

Compressed—flattened laterally. Concave—hollowed out; the interior of a sphere as opposed to the outer or convex surface. Concavity—state or quality of being concave. Concolorous—uniform in color.

Confluent—running together or overlapping. Conical—cone-shaped. Coniform—cone-shaped. Conjoined—joined together.

Conjunctiva—weakly chitinized connecting areas between abdominal segments. Connexivum—prominent abdominal margin of Heteroptera, at the junction of the dorsal and ventral plates. Constriction—narrowed area or false joint. Contiguous—touching. Converging—gradually narrowing or tapering. Convex—evenly and broadly rounded; the outer curved surface of a segment of a sphere, opposed to concave. Convolution—coiling, winding or twisting together. Copulatory—of or pertaining to copulation or pairing. Cord—line of transverse joinings, composed of cross-veins and bases of principal forks. Cordiform—heart-shaped. Coriopruina—white frosted area between the anterior apex of the corium and the clavopruina. Corium—flexible membrane between body segments or appendage segments. Corners—acute projection. Corona(e)—crown-like structure. Coronal suture—median unpaired part of epicranial suture.

Corpus bursae—dilated membranous sac at the anterior end of the bursa copulatrix.

setae or spines.

Cremaster—apex of last segment of abdomen. Crenate—margin evenly notched with rounded teeth. Crenulate—small rounded projections. Crenulation—series of rounded projections. Cribiform—sieve-like. Crochets—sclerotized curved hooks.

Crossveins—transverse veins linking the principal longitudinal veins. Cubital—referring or belonging to the cubitus. Cubital vein—cubitus. Cubitoanal crossvein—crossvein between the cubitus and anal vein.

Cubitus—(Cu)sixth longitudinal vein immediately posterior to the media. Cubitus branches—Cuj, Cu2, etc. Cultriform—sickle-like.

Cupule—cup-shaped organ. Cusps—pointed processes at or near apex of teeth or mandible. Cuticle—exoskeleton.

Cylindriform—cylinder-shaped Dactyl—finger-like structures. Declivent—downward slope. Decurrent—closely attached to and running down another body. Decurved—bowed or curved downward.

Deflexed—abruptly bent downward. Demarcate—separate. Dentate—toothed.

Dentations—toothed projections. Dentes(pi.)—tooth or tooth-like process; proximal segment of distal arms of manubrium. Denticles—serrations along inside of the outer cusps. Denticulate—set with little teeth or notches.

Glossary

Dextral—of the right hand.

1279

Differentiated—make different.

Entire—having the margin unbroken. Epandrium—ninth abdominal tergite of the male

Digitate—finger-like process. Digitiform—finger-like. Dilated—widened or expanded. Dimorphic—having both apterous and long-winged

Epaulettes—sclerotized ridge. Epicranial suture—shaped line on the top of the head with two arms diverging anteriorly (frontal

forms.

Dimorphism—difference ofform. Discal cell—large median cell.

Discoidal cell—term applied to some outstanding or major cells of an insect wing. Disk—central part of the pronotum or elytra apart from the margin. Distad—toward wing apex or farthest away from the body. Distal—situated away from the point of origin or attachment.

Dististyle—gonostyle. Divergent—spread out from body. Dorsal—of or belonging to the upper surface. Dorsolateral—at the top and to the side. Dorsomedian—toward the back and near the midline.

Dorsomentum—upper and more distal of the two subdivisions produced when the mentum is completely (in mosquito larvae) or incompletely (in chironomid larvae) divided by a transverse inflection of membrane.

Dorsoventrai—in a line from the upper to lower surface.

Dorsum—upper surface. Ductus bursae—copulatory opening. Ecdysial suture—inverted Y on the dorsal midline of the head.

Elbowed—antenna that is sharply bent. Elevated pit—cup. Elevation—raised places. Elliptical—resembling oval but having two rounded ends equal. Elliptical valves—opening on a Trichoptera case. Elongate—drawn out; lengthened; much longer than wide.

Elytron (elytra)—hard forewing(s). Emarginate—notched at the margin. Emargination—same as emarginated. Embolar—pertaining to the embolium. Embolar fracture—suture or indentation pertaining to the embolium of the heteropterous wing. Embolium—hair-like or pad-like structure on hemelytron. Empodium—any structure between the claws. Endo—prefix meaning within. Endochorion—inner layer of the chorion of the insect egg-

insect.

sutures). Epicuticle—nonchitinous structureless external film

like layer of the cuticle covering the exocuticle. Epidermis—cellular layer of skin, underlying and secreting the cuticula. Epimeron—posterior division of a thoracic pleuron. Epipleuron—deflexed or infiexed portion of the elytron. Epiproct—dorsal part of the eleventh abdominal segment.

Episternum—anterior sclerite on the pleuron. Epithelium(a)—layers of cells which covers a surface or lines a cavity. Equidistant—equally distant. Erect—perpendicular. Evanescent—fading. Exarate—appendages free, distinct, not fused. Excavated—hollowed out.

Excised—with a deep cut or notch. Excision—deep cut. Exo—prefix meaning on the outside of, without. Exochorion—that part of the chorion derived from the ectoderm; the outer layer of the chorion. Exsertile—extrude.

Extratracheal dark pigment—marginal spots on caudal gill. Extrudable—able to be protruded, projected, pressed, squeezed or pushed out. Exuviae—caste skin of insects.

Eye—compound eye. Eye cap—enlarged basal segment of antenna. Eye patch—pigmented area around eye. Eyespot—pigmentation patches of the head capsule. Facet—surface of the numerous small eyes that compose the compound eye. Falcate—sickle-like.

Falciform—same as falcate.

False vein—fold-like thickenings in the wing membrane.

Fan—segment or process flattened and spread triangularly or in a semicircle, appearing fan like.

Fasciculate—clustered.

Fastigium—flattened or depressed area atop the vertex of the head between and in front of the

compound eyes. Feeler—another name for antenna.

1280

Glossary

Felt chamber—internal component of the thoracic horn which is apparent as the continuation of the connection of the tracheal system. Femur (femora)—third leg segment, attached to trochanter and to the tibia.

Fenestrated—with transparent areas or window-like spots.

Fibrils—thread-like filaments at base of gill lamellate.

Filament—long tapering appendages. Filamentous gills—filament- or thread-like gills. Filiform—thread-like.

Fissure—narrow longitudinal opening. Flabellate—folding like a fan. Flabellate antenna—antenna(e) with fan-like processes or projections. Flagellomeres—second through terminal antennal segments.

Flagellum—whip-like process or third part of antenna.

Flange—projecting rim or edge. Flap—broad, flat piece fastened at one end only. Fluting—alternating ridges and valleys in wing. Fold—fold line of wing. Fold lines—line along which wings fold. Foot-sbaped anal loop—cubitoanal loop. Foramen occipitale—occipital foramen. Forceps—pincers or claspers at apex of abdomen. Fore—anterior.

Foreleg—first leg. Foretarsi—tarsus of the foreleg. Foretibia—tibia of the foreleg. Foretrochantin—prothoracic episternum. Forewing—first wing arising from mesothorax. Fossa—deep pit. Fossorial—expanded structure for digging. Foveate—with a deep impression. Foveolae—pair of small, usually elongate-oval depressions on each side of the fastigium near the top-front of the head. Fringe—edging of hairs, scales extending beyond margin. Frons—face.

Frontal—of or pertaining to the frons. Frontal apotome—chitinous region between and posterior to the base of the antenna. Frontal costa (frontal ridge)—raised, broad, vertical ridge along the midline of the front of the face. Frontal suture—suture formed by the epicranial arms.

Frontoclypeal apotome—fused clypeus and frontal apotome.

Frontoclypeal suture—suture between the frons and the clypeus.

Frontoclypeus—combined frons and clypeus when suture is obsolete.

Fumose—smoky.

Funicle—part of antenna between the club and ring segments.

Furcal pits—external pit on thoracic sternum. Furcasternum—sternum bearing the furca. Furcula—forked process. Furrow—marked narrow depression. Fused—united.

Fusiform—spindle-shaped. Galea(e)—outer lobe of maxilla. Gena(e)—cheek, side of head. Geniculate—suddenly bent at an angle.

Genital—of or pertaining to the reproductive organs. Genital opening—external orifice of the genital cavity.

Genital pocket—genital fossa containing male genital. Genital sheaths—usually weakly chitinized sacs that enclose the male and female genitalia. Genitalia—external parts of the reproductive organs. Gill—respiratory organ. Gill remnants—plate-like or filamentous outgrowth of gills.

Gill tuft—group of lateral, mainly filamentous gills. Glabrous—smooth, shiny or without pubescence or punctures.

Gland—cellular organ or structure which secretes certain characteristic products, e.g., was, saliva, silk, hormones, etc.

Globose—sphere-shaped. Globular—spherical. Glossa(e)—tongue or one of a pair of lobes on the inner apex of prementum of labium. Gonapophysis lateralis—lower valves of ovipositor. Gonocoxite—part of the male terminalia. Gonopods—appendage of the genital segment or associated segment, modified for copulation, intromission or oviposition. Gonopore—external opening of the genital duct. Gonostylus (gonostyli)—stylus of genital segment. Granulate—appearing as if covered with small grains.

Granulations—roughening on the surface. Granule—small grain-like elevation. Groove—furrow.

Gula—ventral region between the base of the beak and the collar.

Gular suture—line of division between gula and genae.

Hair—slender flexible filament.

Halteres—modified hind pair of wings, balancing organ.

Glossary

Hammer—tubercle on the posterior part of sternum. Hamules—accessory male genitalia. Harp—largest of multiple, well delimited, radiating cells on wings used for radiating sound. Harpago(nes)—styli (gonostyli) of the ninth segment

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Hypopygium—male genitalia. Hypostoma—region of the subgena behind the mandible.

clasping organs. Haustellum—proboscis.

Hypostomal bridge—mentum or maxillary plate. Immaculate—without a spot or stain. Impressed—lying below the surface. Impression—indentation on the surface. Incision—any cut into a margin or surface.

Head—first or anterior region of the insect body, articulated at its base to the thorax, bearing the

Incisors—cutting tooth or mandible.

of the male insect when modified to form

mouth structures and antennae.

Head capsule—fused sclerites of the head region forming a hard compact case. Head ratio—length of the head capsule divided by the maximum width of the head.

Hemelytra—forewings. Hemicylindrical—half a cylinder. Hemispherical—shaped like a half sphere. Hemolymph—or haemolymph; a fluid, analogous to the blood in vertebrates, that circulates in the

interior of the invertebrate body remaining in direct contact with tissues.

Heteroideous—middle crochets abruptly longer on each proleg. Hind—refers to hind leg.

Hindgut—intestinal canal from the end of chylific ventricle to the anus, including the Malpighian tubules and anal glands. Hindwing—wing arising from the metathorax. Holdfast—organ or structure of attachment. Holoptic—touching. Homoideus—no abrupt change in length. Hook—sclerotized prong with decurved end. Hook plate—paired dorsal sclerites. Horn—pointed sclerotized process on the head. Humeral—near or on the shoulder or humerus.

Humeral crossvein—(h) vein extending between the costa and subcosta close to base. Humerus—shoulder.

Hump—retractile protuberance. Hyaline—clear. Hydrofuge—water repelling. Hydrostatic air-sacs—air sac within the body of an insect.

Hypandrium—ninth abdominal sternite of the male insect; tenth sternite in mayflies, modified into a transverse plate. Hypocostal ridge—ventral surface of embolium at hemelytral base. Hypognathous—mouthparts directed downward. Hypogynial valves—hypovalvae; valvular processes of the eighth abdominal sternite. Hypopharyngeal—relating to the hypopharynx. Hypopharynx—tongue on upper surface of labium.

Incisor lobe—toothed distal lobe on mandible.

Incomplete—not complete. Inferior—below or behind.

Inflexed—bent, turned inward or downward. Infracoxal—subcoxal.

Inner margin—anal margin. Insertion—place of attachment. Integument—outer covering, outer enveloping cell layer or membrane. Intercalary—inserted between. Intercalary medius anterior(IMA)—furrow vein lying between the two ridge branches of MA. Intercalary veins—lie between principal veins. Intercoxal process—median protrusion. Interocular space—area between compound eyes. Interpleural—of or pertaining to the interpleuron(a). Intersegmental—between segments. Intersternum—in front of first abdominal segment. Interval—area between two elytral striae. Irrorate—with minute spots or granules. Juxta—sclerotized plate. Katepisternum—lower portion of divided episternum. Keel—elevated ridge. Labial—on or belonging to the labium. Labium—fleshly lower lip. Labral—of or pertaining to the labrum. Labral fans—pair of head fans or rays on the head of the larval black fly used in feeding to filter particles out of the water current. Labral lamella—one or more smooth to apically pectinate, scale-like or leaf-like lamella. Labral sensilla—simple sense organs, or one of the structural units of a compound sense organ on the labrum.

Labrum—upper lip. Lacinia(e)—inner lobe of each maxilla. Lamella(e)—any thin plate-like structure. Lamellar—having, consisting or arranged in lamellae.

Lamellate—sheet-like, leaf-like or needle-like.

Lanceolate—oblong and tapering to an end. Laterad—toward the side and away from the median line. Laterad—same as lateral.

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Glossary

Lateral—pertaining to the side. Lauterborn organ—compound sensory organ usually originating at the apex of the second antennal segment.

Lauterborn organ stalk—sclerotized structure basal to the lauterborn organ and serving as an attachment to the antenna.

Leg sheatbs—cuticular covering of legs. Legs—paired appendages of thorax. Lid—operculum or cover.

Ligula(e)—collective name for glossae and paraglossae or central portion of the labium. Linear—long narrow line. Lobate—with lobes.

Lobe—^prominent rounded process. Longitudinal—in the direction of the long axis. Longitudinal vein—vein that extends lengthwise. Lower calypter—proximal calypter of wing. Lyre-shaped—with bowed sides and two outward projection lobes. M-appendage—medioventral appendage of prementum.

Medial cells—wing cells anteriorly bounded by the media or its branches. Medial crossvein—the crossvein which extends from

media 2(M2)to media 3(Mj). Mediale—the second axillary sclerite of the insect wing. Median—in or at the middle; of or pertaining to the middle.

Median cell—closed area formed by a line extending from the end of the subcostal to the end of the median veins.

Median terminal filament—same as median gill. Median vein—(M)media. Medio-cubital crossvein—(m-cu)crossvein between the lower first fork of the medial and upper first fork of the cubital.

Mediocubital—of or pertaining to the media and cubitus of the wing. Medius—middle.

Membrane—thin, transparent tissue.

Membranule—small opaque expansion behind first anal vein.

Macrochaeta(e)—large thick setae. Macroplastron—compressible gas gill. Macropterous—long or large winged. Macroseta(e)—setae larger than adjacent setae. Macrotricbia—large microscopic hairs.

Mentum—sclerotized midventral plate of head capsule. Meron—posterior part of basicoxite. Mesal—pertaining to the middle. Mesepimeron—posterior division of the mesothorax

Macula—a spot.

pleura. Mesepisternum—episternum of the mesothorax. Meso—prefix meaning middle. Mesocoxal cavity—middle coxal depression. Mesonotal—pertaining to the notum.

Mala—single lobe of the maxilla. Malpighian tubules—insect urinary system; long and slender blind excretory tubes lying in the haemocoele, variable in number, opening into the commencement of the hind intestine.

Mamillate—nipple-like.

Mandible—first pair ofjaws. Mandibular tusks—tusks of the mandible.

Manubrium—large median base of the furcula bearing the dentes. Margin—border or edge. Marginal vein—portion of single composite vein along wing margin. Margined—with a single border. Maxilla(e)—secondary pair ofjaws, lateral mouth region. Maxillary lacinia(e)—blade-like mesal sclerite attached or belonging to the distal margin of the maxillary stipes. Maxillary palp(i) or palpus—segmented palp originating from the ventral margin of the maxilla. Media—(M)fourth of the longitudinal veins extending from the base through approximately the middle of the wing. Media branches—Mj, M2, etc. Medial—referring to or at the middle.

Mesonotal shield—notum of the mesothorax.

Mesonotum—upper surface of second thoracic segment.

Mesopleural—pertaining to the mesopleuron. Mesopleuron—pleuron of mesothorax. Mesoseries—band of crochets or hooks extending longitudinally on the mesal side of a proleg. Mesosternum—sternum of the mesothorax.

Mesothorax—second segment of thorax bearing the middle legs and forewings. Meta—any posterior (generally third) part of a structure.

Metacephalic rods—internal sclerotized portion of head.

Metanotal shield—notum of metathorax (third thoracic segment). Metanotum—upper surface of third thoracic segment. Metapleural—of or pertaining to the metapleuron(a). Metapleuron—pleuron of the metathorax. Metapneustic—spiracles on the terminal segment only.

Glossary

Metasternum—sternum of metathorax.

Metathoracic—of or pertaining to the metathorax. Metathorax—third segment of the thorax bearing the hind legs and hind wings if present. Metaxyphus—in Hemiptera, spinose or triangular process of the metasternum. Metepisternum—episternum of the metathorax. Micropylar—of or pertaining to the micropyle. Micropyle—minute opening. Microtrichia—fine hairs. Midbasal—median.

Middorsal—of or pertaining to the middle part or median line of the back.

Midgut—midintestine; the chylific ventricle with the caecal glands, tubes or pouches. Midleg—leg of the mesothorax or second segment of thorax.

Minute spine—small spine. Mirror—secondary area on wings which contributes to sound production. Mola—ridged or roughened surface of mandible. Molar—grinding surface of mandibles. Molar plate—projecting basal lobe of a mandible. Moniliform—bead-like.

Mouth brushes—ventral brush-like organs of the labrum.

Mouth hooks—sclerotized hooks protruding ventrally from mouth area. Movable hook—labial hook.

Mucro—small pointed projection, or spine-like ending on a terminus. Multiarticulate—many jointed. Multiordinal—single row, but different lengths, three or more.

Multiserial fringe—arising from three or more lines. Muscle scars—attachment points for muscles of head visible as round spots. Nasale—anterior and median projection from the frons.

Nase—small to moderate tubercle near tip of wing sheath. Nasus—nose. Neck—cervix. Nodal furrow—transverse suture.

Node(us)—costal fracture. Nodose—swelling on each antenna segment. Nodule(us)—small knot or lump. Nota—dorsal plates of the thorax. Notal—of or pertaining to the nota. Notch—indentation on a margin. Notopleural sutures—suture separating the pronotum from the proepisternum. Notuni—dorsal surface or upper sclerite of a thorax.

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Nygma(nygmata)—spots on the wings of certain

insects which have a peculiar dense cuticular structure.

Obconic—form of a reversed cone.

Oblique—any direction between the vertical and horizontal.

Oblique vein—slanting crossvein. Oblong—four-sided figure with one dimension longer than the other. Obsolete—almost or entirely absent. Obtect—appendages fused to body. Obtuse ungulate—angle larger than a right angle. Occipital foramen—opening on posterior surface of head opposed to a similar opening in prothorax. Occiput—upper and posterior portion of head. Ocellus(i)—simple eye consisting of a single lens. Ommatidia—units composing compound eye. Ompbalium—single median scent gland opening. Operculate—having an operculum, lid or cover. Operculum—lid or cover.

Opistbognatbous—^mouthparts directed backward. Orbicular—round and flat.

Ostiole—small openings. Ostium bursae—female copulatory orifice; entrance to the bursa copulatrix. Outer margin—apical margin. Ovipositor—female genitalia. Ovoid, ovoidal, oviform—egg-like; shaped like an egg-

Pad—part between claws. Paddle—flattened joints of the posterior tarsi in aquatic Heteroptera. Pala(e)—tarsus of foreleg. Palatal brushes—brushes on the labrum of larval

mosquitoes that create water movement. Palmate—shaped like a hand.

Palp(s), Palpus(i)—elongated segmented structures on the maxillae and labium.

Palpal—belonging, relating or attached to the palpi. Palpifer—sclerite bearing a palpus. Papilla(e)—soft projections. Papillate, papallatus—with small surface elevations, porous at tip. Para—prefix meaning next to, near by, at the side of.

Paraglossa(e)—paired structure of the labium. Parameres—smaller inner pair of male gonapophyses, closely associated with the aedeagus. Paranal cell—enclosed area to the side of or next to the anus or anal structures.

Paraproctfs)—ventral paired portion of abdominal segment.

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Glossary

Parietal—of or pertaining to the wall of a cavity of the body or of an organ. Patagia—overlapping plates.

Posterior crossvein—vein or veins closing discal cell apically(M and M3). Posterodorsal—of or relating to the posterior part of

Pearl rows—one or more rows of small rounded

projections. Pecten—comb-like structure.

Pecten epipharyngis—three structures behind labral region margin. Pectinate—comb-like or coarsely feather-like. Pedicel—second segment of antenna or a stalk or stem supporting a structure. Pedisulcus—notch. Peduncle—stalk-like structure.

the back.

Posterolateral—lateral and to the rear.

Postgenae—lateral parts of the occipital arch. Postgenal cleft—cleft or longitudinal split situated behind the gena or cheek. Postmentum—mentum plus submentum. Postnodal crossvein—extends from the nodus to the

pterostigma between the costa and radius. Postnodal pruina—frosted area. Postnodal setae—setae found after or following a node.

Pedunculate—set on a stalk.

Peg—short blunt projection. Peltate—shield shape. Penellipse—crochets in a incomplete circle. Penes—penis. Penultimate—next to the last.

Perianal pad—pad-like structure. Petiolate—attached by a stalked. Petiole—stem or stalk.

Phallobase—part of reproductive organ. Phallus—penis. Pharate—of or designating an instar of an insect which is confined within the cuticle of the

previous instar. Pile—hairy or fur-like covering; thick, fine, short, erect hair, giving a surface appearance like velvet.

Pilose—hairy.

Pilosity—fine long hair. Pinaculum (pinacula)—enlarged seta-bearing papilla(e)forming a flat plate. Pinnately branched—similarly branched on each side.

Postnotum—intersegmental plate of the dorsum of the thorax associated with the tergum of the preceding segment, bearing the antecosta and usually a pair of phragmatal lobes. Postocciput—far margin of head. Postocular—behind the eyes.

Postocular space—area in back of or behind the eyes. Postpleurite—pleurite of the prothorax behind the coxa.

Postscutellum—convex area immediately behind the scutellum.

Poucb—large sac. Praecinctorium—unique flap over the tympanal organ.

Pre—prefix meaning before, anterior to. Preanal—next to the last segment of abdomen. Preapical—just before the apex. Precorneal setae—setae close to the base of the thoracic horn.

Prefrons—articular region anterior to the frontal apotome. Prehalter—membranous scale in front of the true halter.

Planate—with flattened surface.

Plastron—air film on the surface of the body. Plate—broad flattened surface.

Prehensile—adapted for grasping. Premandible—paired appendages originating from

Plectrum—rasp.

Pleura—lateral components of each abdominal segment that connect the terga and sterna. Pleural suture—external suture.

Pleurite—any minor sclerite into which the pleural area of a segment is divided. Pleuron—lateral region of any segment. Plica(e)—fold or wrinkle. Plumose—very finely feather-like. Post—prefix meaning after, behind. Postclypeus—posterior part of clypeus. Posterad—contraction of posteriorad meaning toward the rear; in the direction of the hinder or hindmost.

Posterior—the rear.

the ventral surface of the labrum.

Premental—of or pertaining to the prementum. Prementohypopbaryngeal complex—occur on the medioventral aspect of the head complex. Prementum—the stipital region of the labium. Preoral—before the mouth in position. Prescutum—anterior area of the meso- or metanotum in front of the scutum. Presternum—narrow anterior area of the sternum. Pretarsal—anterior to the tarsus.

Primary setae—setae on the head, thorax or abdomen.

Pro

pertaining to the first thoracic segment or its appendages. Proboscis—sucking tube-like mouthpart.

Glossary

Procercus(i)—fleshly tubercle originating from the dorsal surface of the preanal segment. Process(es)—projection(s) or prominent appendage(s). Procoxa—forecoxa.

Proctiger—small papilla (the reduced tenth abdominal segment) bearing the anus. Prognathous—mouthparts directed forward or anteriorly. Projection—outgrowth, protuberance, protrusion. Proleg(s)—usually paired, round, elongate, fleshy retractable processes that bear apical spines or crochets.

Prominent—rising from above the surface or produced beyond the margin. Pronotai—of or pertaining to the upper surface of first thoracic segment(pronotum). Pronotum—of prothorax (pronotai shield or notum). Propleura—pleuron of the prothorax. Propleuron—pleuron of the prothorax. Prosternal horn—membranous projection of the prosternum.

Prosternum—sternum of the prothorax. Prostheca—sclerotized process arising from the mesal margin of the mandible.

Prothoracic—of or pertaining to the prothorax. Prothorax—first segment of the thorax which bears the forelegs. Protrusible—extendable. Protuberance—elevation above the surface.

Protuberant—structure produced above the surface.

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Pygidium—^last visible abdominal segment of the abdomen.

Pygophore—large upper piece of the genitalia in Homoptera.

Quadrangle—four angle cell. Quadrate—four-sided. Radial—of or pertaining to the radius. Radial crossvein—(r) between the radius and first branch of the radial sector.

Radial planate—transverse the interradial area. Radial sector vein—(Rs) divides into two branches.

Radial vein—radius, often the heaviest vein.

Radio-medial crossvein—(r-m) between the lower first fork of the radial sector and upper first fork of median vein.

Radius—(R)third longitudinal vein. Radius branches—R,, R2, etc. Ramus—branch-like subdivision of any structure. Raptorial—grasping. Rastrate—with parallel, longitudinal scratches. Rectum—posterior part of the terminal section of the proctodaeum. Recumbent—lying down or reclining. Recurved booklets—one or more rows of anteriorly directed hooks on the posterior margin of abdominal segments. Repose—at rest. Repugnatorial gland—glands which secrete malodorous or noxious liquids or vapors, as a defense against enemies.

Respiratory organ—typically self-contained

Proximal—part of an appendage nearest the body. Proximal end—nearest the wing base. Proximal part—nearest the body.

respiration. Respiratory tube—caudal breathing tube.

Proximal side—inner end.

Reticulate—meshed or covered with network of fine

Pruinescence—minute dust.

anatomical feature that facilitates external

lines.

Pruinose—frosted.

Reticulate veinlets—network of small, meshed or

Pseudocelli—sense organs resembling ocelli. Pseudopodia—prolegs. Pseudoradula—longitudinal band of fine to minute spinules on dorsal surface of M-appendage. Pterostigma—enlarged cell in wing. Pterothorax—closely fused meso- and metathorax.

netted veins in the wing. Retinaculum—tooth-like process. Retracted—being drawn back or in. Retractile—capable of being drawn back or in. Retractile antenna—antenna(e)capable of being produced and drawn back or retracted. Rhomboid—diamond-shaped. Ridge—raised narrow strip. Ring—circle or annulus, usually margining a discolored spot. Ring joints—proximal segment or segments of

Ptilinal—frontal.

Ptilinal suture—crescent-shaped groove situated on lower part of frons between the bases of antennae and eyes. Pubescent—hairy. Pulvilliform—pad-like. Pulvillus(i)—pad-like structure between base of claws. Punctate—set with punctures. Punctures—small impressions. Puparium—sclerotized exuviae.

claviola.

Rostral—of or pertaining to the rostrum. Rostrum—extended portion of head, beak. Rudimentary—undeveloped. Rugose—wrinkled.

1286

Glossary

Sa 1—medial setal area on mesonotum or metanotum.

Sa 2—mediolateral setal area on mesonotum or metanotum.

Sa 3—lateral setal area on mesonotum or metanotum.

Saddle—sclerite covering dorsal surface. Scale—broad flattened hair.

Scallop—series of concave indentations giving the appearance of evenly rounded depressions. Scape—basal segment or first joint of antenna. Scent gland—glandular structures, sometimes

Siphon—any tubular external process. Snout—prolonged part of head. Somite—body segment of the adult insect. Sonorific—sound producing; applied to stridulating organs.

Sparse—scattered. Spatulate—narrow and flat at the base and enlarged at the apex. Spermatheca—sac or reservoir in the female that

Setal row—row of setae.

receives the sperm during coition. Spicule—small spines. Spine—thorn-like process. Spine groups—groups of spines on abdomen. Spine row—a line of thorn-like processes. Spiniform—in the form or shape of a spine. Spinneret—silk producing structure. Spinule—small spines or scales. Spinule row—a line of small spines or scales. Spiracle—small external breathing opening of tracheal system. Spiracular disk—disk area on caudal segment possessing respiratory openings. Spiracular ring—lightly sclerotized or differentially pigmented margin of spiracle on the dorsum of the antepenultimate abdominal segment. Spur—large modified setae or spine. Spurious—false. Spurious veins—certain folds or thickenings in the wing surface which resemble veins so nearly as to be readily mistaken for them. Squama—scale-like structure on the wing. Stem vein—any vein from which other veins branch. Stemma (stemmata)—simple eye(s). Stemmatic hulla—swelling behind eye. Sterna—ventral plates of the abdominal segment. Sternellum—second sclerite of the ventral part of each thoracic segment. Sternite—ventral sclerotized part of a segment. Sternopleural suture—suture below and nearly parallel with the dorsopleural suture, separating the mesopleura from the sternopleura.

Setal warts—setae bearing warts on dorsum of head

Sternum—sclerite on ventral surface or lower

eversible, sometimes in the form of hair tufts or pencils for diffusing odors that may be repellant or attractive.

Sclerite—any piece of the body hardened wall surrounded by sutures or membranous areas. Sclerotized—hardened and usually darkened integument. Scutellum—posterior shield-like plate of thorax. Scutum—anterior shield-like plate of thorax. Secondary hypocostal ridge—secondary hypocostal lamina.

Secondary setae—setae that occur randomly. Segment—subdivision of the body marked by sutures.

Semioperculate—half or partly covering. Semiprognathous—mouthparts directed partly or in some degree forward or anteriorly. Sense organ—specialized structure for receiving external stimuli.

Sensillum (sensilla)—simple sense organ, or one of the structural units of a compound sense organ. Serrate—saw-like.

Serrulate—finely serrated. Sessile—attached by the base. Seta(e)—sclerotized hair-like projection. Setaceous—set with many setae or bristles. Setal—of or pertaining to the seta. Setal gap—gap between setae patches.

Setula(e)—stiff hairs. Shagreen—complete or partial fields of small spinules on terga, pleura or sterna.

portion of segment. Stigma—outer costal margin of wing. Stigmal vein—short vein extending posteriorly from the marginal vein. Stigmatal bar—spiracle. Stipes—basic sclerite of the maxilla, distad of the

Sheath—structure enclosing others. Shield—sclerotized plate. Shoulder—basal, external angle of each elytron, the

Stria—fine punctures lengthwise on the elytra. Striate—marked with longitudinal impressions or

or thorax.

Setiferous—setae bearing. Setose—hair covered.

humerus.

Sinistral—to the left.

Sinuate—wavy.

cardo.

furrows.

Striations—series of fine, longitudinal impression lines.

Glossary

Stridulatory—connected with or of the nature of stridulation (sound production). Stridulatory file—any structure wherever situated used in stridulating.

Tarsal—of or pertaining to the tarsus. Tarsal claw-—claw at the apex of the tarsus.

Strigil—file.

Tarsal segments—tarsomere.

Strides—rudimentary stria. Style—long thin appendage or terminal segment of

Tarsomere—subdivision of the tarsus.

antenna.

Stylet—small style or stiff process. Styliger—same as subgenital plate. Styliger plate—posterior portion of abdominal sternum 9 of males.

1287

Tarsal formula—number of tarsomeres on the fore, mid and hind tarsi.

Tarsus(i)—jointed foot segment attached to the apex of the tibia.

Teeth—hardened growths on mandibles, maxillae or stomatal walls.

Tegmen (tegmina)—hardened forewing. Tegula(e)—articular sclerites of wing. Telescopic—consisting of one part that slides inside

Stylus(i)—small pointed nonjointed process(es). Sub under, not quite, nearly. Subalar—below the wings.

Tenaculum—minute organ on the ventral surface.

Subanal—ventral of the anus.

Tenent hairs—adhesive setae located on the

Subanal plate—posterior portion of abdominal sternum 9 of females.

Subapical—situated below or near an apex. Subbasal—located just distad of the base. Subconical—nearly or approximately conical. Subcosta vein—(Sc)just behind costal vein. Subcostal—situated below or near the costa.

Subcylindrical—not quite cylindrical. Subequal—similar, but not equal in character. Subgenital plate—posterior portion of abdominal sternum 9 of males. Subhumeral—near the humerus.

Sublateral—just inside lateral margin. Submedian—situated next to a median part or the midline.

Submental—of or pertaining to the submentum. Submentum—proximal division of the postmentum.

Subnodus—additional oblique crossvein, the continuation of the nodus below vein Rj. Subprimary setae—setae found in late instars. Subtriangular—not completely triangular. Sucker—disk to adhere to surfaces.

Suctorial disc—true suction devices on the ventral

body surface. Sulcate—deeply furrowed or grooved. Sulcus—groove or furrow. Superior—above.

Supertriangle—the wing cell just anterior to the triangle. Supra—prefix meaning above or beyond. Supraanal—above the anus. Supracoxal—lying above or over the coxa. Suranal—supraanal. Suture—groove marking the line of fusion of two distinct plates. Syntborax—mesothorax and metathorax fused together.

another.

underside of the tarsi.

Tentorial arms—cuticular invaginations arising from the anterior and posterior tentorial pits. Tentorium (tentoria)—endoskeleton of the insect head, consisting of two or three pairs of apodemes coalescing internally, which gives rigidity and strength to the head, supports the brain and fore-intestine and affords attachment

to many cephalic muscles. Tentoropharyngeal—reclining sclerites on either side of the pharynx. Terete—cylindrical. Terga—dorsal plates of the abdomen.

Tergite—dorsal sclerite or part of a segment, especially when such part consists of a single sclerite.

Tergosternal—of or pertaining to the terga and sternum together. Tergum—dorsal surface or upper sclerite of abdomen.

Terminal—at the tip. Terminalla—terminal segments of parts or structures taken together. Testaceous—brownish yellow. Thoracic—belonging to or attached to the thorax. Thoracic horn—structure of the anterior thorax.

Thorax—first three segments posterior to head. Thyridial cell—cell formed by the first fork of the median vein; the cell behind the thyridium. Tihia(e)—fourth segment of leg. Tibotarsus—terminal leg segment. Tomentum—form of pubescence composed of matted, woolly hair. Tooth—short pointed process. Trachea(ae)—part of respiratory system. Tracheal gill—filamentous extensions of the body wall.

Tracheation—arrangement of tracheae.

1288

Glossary

Tracheole—one of the finer branches of the tracheae.

Transverse—when the longest dimension is across the body. Tri—prefix meaning three. Trichobothria—sensory hair or hair bearing spots. Trichome—lateral pair of specialized setae. Tridentate—having three teeth, processes, or points. Trilobed—having three lobes. Triordinal—single row with three lengths. Tripartite—divided into three parts. Trocbanter—between the coxa and femur of the leg. Trocbanteral organ—expanded distal setae in pit. Trochantin—sclerite in the thoracic wall immediately anterior to the base of the coxa.

Trumpets—paired dorsolateral appendage containing the spiracles. Truncate—cut squarely at tip. Tubercle—elevated fleshy processes. Tuberculate—bearing tubercle. Tubules—gills. Tufts—group of setae the same length arising from the same location.

Turbinate—raised on a stalk-like process. Tusks—large projections of the mandible. Tympana]organs—hearing. Uncus(i)—curved hook directed downward from a triangular dorsal plate in the male, shielding the penis. Undulate—having a wavy form or surface. Unguiculus(i)—small claw. Unguis(es)—claw. Uni—prefix meaning one. Unidentate—with one tooth only. Uniordinal—same length. Uniserial—arise from a single line and the same length. Urogomphus(i)—paired terminal appendage. Vagina(e)—genital opening. Valve—visible potion of the ovipositor and arise from eighth abdominal sternum. Vannal—anal vein.

Vein—structures that extend into the wing. Veinlets—any small vein in the wing. Venations—arrangement of the veins in the wing. Venter—under surface of the abdomen as a whole; the belly. Ventrad—toward the under surface; in the direction of the venter.

Ventral—pertaining to underside of the insect.

Ventral apotome—gular sclerite of the head capsule. Ventral disk—ventral gills. Ventral tube—median lobe projecting ventrally from first abdominal segment. Ventral tubule—fleshy protuberances. Ventrolateral—on the lower surface and to one side of the midline.

Ventromental plate—lateral parts of ventromentum. Ventromentum—lower and more proximal of the two subdivisions produced when the mentum is completely (in mosquito larvae) or incompletely (in chironomid larvae) divided by a transverse inflection of membrane.

Ventromesial margins—lower surface and of the midline.

Vermiculate—marks resembling the tracks of a worm. Verrucae—wart-like elevation bearing setae. Vertex—top of head between the eyes. Vertical fringe—upright setae. Verticillate—whorled setae.

Verticils—one of the whorls of long, fine hair arrange symmetrically on the antennal segments. Vesica spermalis—seminal vesicle; an enlarged portion of the vas deferens used for sperm storage.

Vesicle—dark lobe arising from anterior abdominal segment.

Vestigial—degenerate structure that was better developed or functional at one time. Villopore—ventral patch of stiff hairs at the base of the femur of a foreleg. Vulvar lamina—posterior margin of the abdominal sternum 8 of female.

Whorl—ring of hairs set about a joint or center like the spokes of a wheel. Wing base—proximal part of the wing between the bases of the veins and the body, containing humeral and axillary sclerites. Wing cases—wing cover. Wing cells—areas of the wing enclosed by veins. Wing pads—encased, undeveloped wings. Wing scales—flattened, modified setae on wings; tegula. Wing sheaths—cuticular coverings of the pharate adult wings. Wing(s)—paired membranous appendage(s). Wrinkled—with cracks or creases.

Xyphus—spinous or triangular process of the mesosternum in many Hemiptera.

Y-ridge—mesosternum y-shaped line.

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;c'i'

; V/

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5676. Stark, B. P., and S. W. Szczytko. 1988. Egg morphology and phytogeny in Arcynopterygini (Plecoptera: Perlodidae). J. Kans. Ent. Soc. 61:143-160. 5677. Stark, B. P., B. C. Kondratieff, and C. J. Verdone.

2015. Kathroperla siskiyou, a new stonefly species from California and Oregon, U.S.A.(Plecoptera:

Chloroperlidae). Illiesia 11:92-103. 5678. Stark, B. P., B. C. Kondratieff, J. B. Sandberg, B. A. Gill, C. J. Verdone, and A. B. Harrison. 2015. Sierraperla Jewett, 1954 (Plecoptera: Peltoperlidae), distribution, egg

morphology and description of a new species. Illiesia 11:822.

5679. Stark, B. P., B. C. Kondratieff, R. F. Kirchner, and K. W. Stewart. 2010. Larvae of seven eastern North American

Sweltsa (Plecoptera: Chloroperlidae). Illiesia 6:51-64. 5680. Stark, B. P., B. R. Oblad, and A. R. Gaufin. 1973. An annotated list of the stoneflies(Plecoptera) of Colorado. Ent. News 84:269-277.

5681. Stark, B. P., K. W. Stewart, S. W. Szczytko, and R. W. Baumann. 1998. Common names of stoneflies

(Plecoptera) from the United States and Canada. Ohio Biol. Surv. Notes 1:1-18.

5682. Stark, B. P., K. W. Stewart, S. W. Szczytko, R. W. Baumann,and B. C. Kondratieff. 2012. Scientific and Common Names of Nearctic Stoneflies (Plecoptera), with Corrections and Additions to the List. The Caddis Press, Misc. Contr. 1:1-20.

5683. Stark, B. P., R. W. Baumann, B. C. Kondratieff, and K. W. Stewart. 2013. Larval and egg morphology of Paraperla frontalis(Banks, 1902) and P. wilsoni Ricker, 1965 (Plecoptera: Chloroperlidae). Illiesia 9:101-108. 5684. Stark, B. P., S. W. Szczytko, and B. C. Kondratieff. 1988. The Cultus decisus complex of eastern North America (Plecoptera: Perlodidae). Proc. Ent. Soc. Wash. 90:91-96. 5685. Stark, B. P., S. W. Szczytko, and C. R. Nelson. 1998. American Stoneflies: A Photographic Guide to the Plecoptera. Caddis Press, Columbus, Ohio. 126 pp. 5686. Stark, B. P., T. A. Wolff, and A. R. Gaufin. 1975. New

records of stoneflies(Plecoptera)from New Mexico. Great Basin Nat. 35:97-99.

5687. Stark, B., and D. L. Lentz. 1988. New species of Nearctic Neoperla (Plecoptera: Perlidae). Ann. Ent. Soc. Am. 81:371-376.

5688. Stary, J. 1992. Phytogeny and classification of Tipulomorpha, with special emphasis on the family Limoniidae. Proc. First Internat. Symp. on Tipulomorpha. Ada zool. Cracov 35:11-36. 5689. Stary, J., and F. Brodo. 2009. Arctic species of the

subgenus Symplecta sensu stricto (Diptera: Limoniidae). Can. Ent. 141:1-30.

5690. Statzner, B. 1981. A method to estimate the population size of benthic macroinvertebrates in streams. Oecologia 51:157-161.

5691. Statzner, B., A. G. Hildrew, and V. H. Resh. 2001. Species traits and environmental constraints: entomological research and the history of ecological theory. Ann. Rev. Ent. 46:291-316.

5692. Statzner, B., and B. Higler. 1985. Questions and comments

on the river continuum concept. Can. J. Fish. Aquat. Sci. 42:1038-1044.

5693. Statzner, B., and L. A. Beche.2010. Can biological invertebrate traits resolve effects of multiple stressors on running water ecosystems? Freshwat. Biol. 55:80-119. 5694. Statzner, B., and T. F. Holm. 1982. Morphological adaptations of benthic invertebrates to stream flow—an old question studied by means of a new technique (laser doppler anemometry). Oecologia 53:290-292. 5695. Statzner, B., and T. F. Holm. 1989. Morphological adaptation of shape to flow: microcurrents around lotic macroinvertebrates with known Reynolds numbers at quasi-natural flow conditions. Oecologia 78:145-157. 5696. Statzner, B., J. A. Gore, and V. H. Resh. 1988. Hydraulic stream ecology: observed patterns and potential applications. J. N. Am. Benthol. Soc. 7:307-360. 5697. Steedman, R. J., and N. H. Anderson. 1985. Life history and ecological role of the xylophagous aquatic beetle, Lara avara LeConte (Dryopoidea: Elmidae). Freshwat. Biol. 15:535-546.

5698. Steele, B. D., and D. C. Tarter. 1977. Distribution of the family Perlidae in West Virginia. Ent. News 88:18-22. 5699. Steelman, C. D. 1976. Effects of external and internal arthropod parasites on domestic livestock production. Ann. Rev. Ent. 21:155-178.

5700. Steelman, C. D., and A. R. Colmer. 1970, Some effects of

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5701. Stehr, F. W.(ed.). 1987. Immature Insects. Vol. 1. Kendall/ Hunt Publ. Co., Dubuque, lA. 754 pp. 5702. Stehr, F. W.(ed.). 1991. Immature Insects. Vol. 2. Kendall/ Hunt Publ. Co., Dubuque, lA. 974 pp.

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5716. Stewart, K. W. 2000. Taeniopterygidae (The Willowflies), pp. 55-88. In B. P. Stark and B. J. Armitage (eds.). Stoneflies(Plecoptera) of eastern North America. Vol. 1. Pteronarcyidae, Peltoperlidae, and Taeniopterygidae. Bull. Ohio Biol. Surv. New Ser. Vol. 14. Ohio Biological Survey, Columbus, OH.99 pp. 5717. Stewart, K. W. 2009. New descriptions of North American Taenionema larvae (Plecoptera: Taeniopterygidae). Illiesia 5:128-145.

5718. Stewart, K. W,and B. C. Kondratieff. 2012. Larvae of the Nearctic species of the stonefly genus Megarcys Klapalek (Plecoptera: Perlodidae). Illiesia 8:16-36. 5719. Stewart, K. W,and B. P. Stark. 1977. Reproductive system and mating of Hydroperla crosbyt a newly discovered method of sperm transfer in Insecta. Oikos 28:84-89.

5720. Stewart, K. W,and B. P. Stark. 1988. Nymphs of North American stonefly genera (Plecoptera). Thomas Say Found. Ser. Ent. Soc. Am. 12:1-460.

of western North American Podmosta larvae and

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5726. Stewart, K. W,and J. A. Stanger. 1985. The nymphs, and a new species of North American Setvena lilies (Plecoptera: Perlodidae). Pan-Pac. Ent. 61:237-244. 5727. Stewart, K. W,and J. B. Sandberg. 2003. Life history of a Colorado population of Kugotus modestus (Plecoptera: Perlodidae), pp. 195-200. In E. Gaino (ed.). Research update on Ephemeroptera and Plecoptera. University of Perugia, Italy. 488 pp. 5728. Stewart, K. W,and K. D. Alexander. 1997. Further considerations of mate searching behavior and communication in adult stoneflies (Plecoptera): first report of tremulation in Suwallia (Chloroperlidae), pp. 107-112. In P. Landolt and M. Sartori (eds.). Ephemeroptera and Plecoptera Biology-Ecology-Systematics. Mauron & Tinguley & Lachat SA, Mancor, Fribourg, Switzerland. 569 pp. 5729. Stewart, K. W., and M. Maketon. 1991. Structures used by Nearctic stoneflies (Plecoptera) for drumming, and their relationship to behavioral pattern diversity. Aquat. Insects 13:33-53.

5730. Stewart, K. W., and M. O. Oswood. 2006. The stoneflies (Plecoptera) of Alaska and western Canada. The Caddis Press, Columbus, OH. 5731. Stewart, K. W., and P. P. Harper. 1996. Plecoptera, pp. 217-266. In: R. W. Merritt and K. W. Cummins (Eds.). An Introduction to the Aquatic Insects of North America, 3rd ed. Kendall Hunt, Dubuque, Iowa. 5732. Stewart, K. W., andW. E. Ricker. 1997. Stoneflies (Plecoptera) of the Yukon, pp. 201-222. In H. V. Danks and J. A. Downes (eds.). Insects of the Yukon. Biological Survey of Canada (Terrestrial Arthropods), Ottawa. 1,034 pp. 5733. Stewart, K. W., B. P. Stark, and L. L. Serpa. 2013. Larvae of the two North American species of Calileuctra (Plecoptera: Leuctridae). Illiesia 9:1-13. 5734. Stewart, K. W., B. P. Stark, and T. G. Huggins. 1976. The stoneflies(Plecoptera) of Louisiana. Great Basin Nat. 36:366-384.

5735. Stewart, K. W., B. P. Stark, P. J. Gullan, and P. S. Cranston. 2002. Nymphs of the North American stonefly genera (Plecoptera). The insects: an outline of Entomology. The Caddis Press, Columbus, OH. 5736. Stewart, K. W., G. L. Atmar, and B. M. Solon. 1969. Reproductive morphology and mating behavior of Perksta placida (Plecoptera: Perlidae). Ann. Ent. Soc. Am. 62:1433-1438.

5737. Stewart, K. W., G. P. Friday, and R. E. Rhame. 1973. Food habits of hellgrammite larvae, Corydalus cornutus (Megaloptera: Corydalidae), in the Brazos River, Texas. Ann. Ent. Soc. Am.66:959-963.

5738. Stewart, K. W., L. E. Milliger, and B. M.Solon. 1970.

Dispersal of algae, protozoans, and fungi by aquatic Hemiptera, Trichoptera, and other aquatic insects. Ann. Ent. Soc. Am.63:139-144.

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Additional Coleoptera References Not Included in Bibliography Above 6909. Angus R. B. 2010. Boreonectes gen. n. a new genus for the Stictotarsus griseostriatus(De Geer)(Coleoptera: Dytiscidae), with additional karyosystematic data on the group. Comp. Cytogenet. 4:123-131. 6910. Babin, J., and Y. Alarie. 2004. Taxonomic revision of genus Gyretes Brulle (Coleoptera: Gyrinidae)from America North of Mexico. Coleopt. Bull. 58:538-567. 6911. Baca, S. M., E. F. A. Toussaint, K.B. Miller, and A.E.Z. Short. 2017. Molecular phylogeny of the aquatic beetle family Noteridae (Coleoptera: Adephega) with an

emphasis on data partitioning strategies. Mol. Phylogen. Evol. 107:282-292. 6912.

Balke, M., J. Hajek, and L. Hendrich. 2017. Generic reclassification of species formerly included in Rhantus Dejean (Coleoptera, Dytiscidae, Colymbetinae). Zootaxa, 4258:91-100.

1454

Bibliography

6913. Barr, C. B.2011. Bryelmis Barr(Coleoptera: Elmidae: Elminae), a new genus of riffle beetle with three new species from the Pacific Northwest, U.S.A. Coleopt. Bull. 65:197-212

6914. Bouchard P, Y. Bousquet, A. Davies, M. Alonso-Zarazaga, J. Lawrence, C. Lyal, A. Newton, C. Reid, M.Schmitt, A. Slipinski and A. Smith. 2011. Family-group names in Coleoptera (Insecta). ZooKeys 88:1-972. 6915. Epler, J. H. 2010. The water beetles of Florida: An identification manual for the families Chrysomelidae, Curculionidae, Dryopidae, Dytiscidae, Elmidae, Gryinidae, Elaliplidae, Helophoridae, Hydraenidae, Hydrochidae, Hydrophilidae, Noteridae, Psephenidae, Ptilodactylidae and Scirtidae. Florida Dept. Environ.

6924. Miller, K. B. 2009. On the systematics of Noteridae (Coleoptera: Adephaga: Hydradephaga): Phylogeny, description of a new tribe, genus and species, and survey of female genital morphology. System. Biodivers. 7:191-214. 6925. Miller, K. B., and J. Bergsten. 2012. Phylogeny and classification of whirligig beetles (Coleoptera: Gyrinidae): relaxed-clock model outperforms parsimony and time-free Bayesian analyses. System. Ent. 37:706-746. 6926. Miller, K. B., and J. Bergsten. 2016. Diving Beetles of the World. Johns Hopkins Univ. Press, Baltimore, Maryland. 336 p.

6927. Mousseau,T,and R. E. Roughley. 2007. Taxonomy, classification, reconstructed phylogeny and biogeography

Protect.

6916. Fikacek, M., and D. Vondracek. 2014. A review of Pseudorygmodus(Coleoptera: Flydrophilidae), with notes

6928.

on the classification of the Anacaenini and on distribution

of genera endemic to southern South America. Acta Ent. Musei Nat. Pragae 54:497-514.

6929.

6917. Gustafson, G.T., and K. B. Miller. 2015. The New World

whirligig beetles of the genus Dineutus Macleay, 1825 (Coleoptera, Gyrinidae, Gyrininae, Dineutini). ZooKeys,

6930.

476:1-135

6918. Hansen, M. 1999. World catalogue of insects. Vol. 2. Hydrophiloidea (Coleoptera). Apollo Books, Stenstrup, 416 pp. 6919. Jach M. A., and M. Balke. 2008. Global diversity of water beetles(Coleoptera) in freshwater. Hydrobiol. 595:

6931.

419-442.

6920. Karaite, Y. 2016. Revision of the genus

6932.

Sanderson, 1953, part 3: The O. elegans species group. Koleopterol. Rundsch. 86:205-212. 6921. Maier, C. M., M. A. Ivie, J. B. Johnson, and D. R. Maddison. 2010. A new northern-most record for the

Family Hydroscaphidae (Coleoptera: Myxophaga), with description of a new Nearctic species. Coleopt. Bull.

6933.

64:289-302.

6922. Mclntosh, C. E., and A. E. Z. Short. 2012. New Delaware

records and notes about the endangered Seth Forest Water Scavenger Beetle (Coleoptera: Hydrochidae). Coleopt. Bull. 66:294-296.

6923. Michat, M. C., Y. Alarie, and K. B. Miller. 2017. Higherlevel phytogeny of diving beetles(Coleoptera: Dytiscidae) based on larval characters. System. Ent., doi:10.1111/ syen.12243

6934.

of Nearctic species of Brychius Thomson (Coleoptera: Haliplidae). Coleopt. Bull. 61:351-397. Perkins, P. D. 2012. A revision of Epimetopus Lacordaire, the New World hooded shore beetles(Coleoptera: Epimetopidae). Zootaxa 3531:1-95. Richards, A. B., and W. D. Shepard. 2017. A review of Stenocolus scutellaris LeConte, 1853(Coleoptera: Eulichadidae), with notes on distribution, morphology and life history. Pan-Pac. Ent. 93:127-140. Short, A. E. Z., L. J. Joly, M. Garcia, A. Wild, D. D. Bloom, and D. R. Maddison. 2015. Molecular phylogeny of the Hydroscaphidae (Coleoptera: Myxophaga) with description of a remarkable new lineage from the Guiana Shield. System. Ent. 40:214-229. Short, A. E. Z., and M. Fikacek. 2013. Molecular phylogeny, evolution, and classification of the Hydrophilidae (Coleoptera). System. Ent. 38:723-752. Short, A. E. Z., J. Cole, and E. F. A. Toussaint. 2017a. Phylogeny, classification, and evolution of the water scavenger beetle tribe Hydrobiusini inferred from morphology and molecules(Coleoptera: Hydrophilidae: Hydrophilinae). System. Ent., DOI: 10.11 ll/syen.l2239. Short, A. E. Z., D. Post, and E. F. A. Toussaint. 2017b. Biology, distribution, and phylogenetic placement of the California endemic water scavenger beetle Hydrochara rickseckeri(Horn)(Coleoptera: Hydrophilidae). Coleopt. Bull. 71:461-467. Short, A. E. Z. 2018. Systematics of aquatic beetles: Current state and future directions(Coleoptera). System. Ent. 43:1-18.

6935. Worthington, R.I, J. L. Hellman, and P. K. Lago. 2016. Hydrochidae (Coleoptera) of Mississippi. Trans. Am.Ent. Soc. 142:167-213.

1

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INDEX Acricotopus, 1160, 1162, 1171, 1200, Aagaardia, 1254 Abedus, 206, 527, 528, 556 Ahedus herherti, 75, 86, 102 Abedus indentatus, 528

Abiskomyia, 1159, 1163, 1198, 1199, 1254 Ablahesmyia, 940, 1130,1131, 1137,1189, 1190, 1247 Ahlabesmyia annulata, 1135 Ahlabesmyia mallochi, 1131 Ablahesmyia parajanta, 1131 Ableptemetes, 270, 271 Ahleptemetes distinctus, 333 Acalcarclla, 1149, 1151, 1264 Acalyptris scirpi, 789 Acamptocladius, 1122, 1166,1174, 1202, 1204, 1229, 1255

Acanthagrion, 358, 362 Acanthagrion quadratum, 364,406 Acanthametropodidae, 167, 195, 198, 268, 274, 275,in,301,303, 321

Acanthametropus, 265, 268, 301 Acanthametropus pecatonica, 274, 275, 277, 303, 321 Acanthamola, 328 Acanthocnema, 225, 1018 Acentrella, 269, 280,292, 301, 309, 311,323

Acentrellafergopagus, 269 Acentrella lapponica, 269 Acentrella parvula, 294 Acentrella turbida, 318 Acentria, 765, 769, 770, 771, 774, 776 Acentria ephemerella, 766,769,774, 776,779 Acentropinae, 765-66, 767, 768, 769, 771, 779-83

>

N

s

s

Acentropus, 51, 779 Acerpenna, 282, 292, 309, 323 Acerpenna pygmaea, 83 Achaetella, 91 Achalcus, 993 Acherontiella, 249 Achradocera, 993 Achurum, 423,425 Acidocerinae, 841 Acigona, 784 Aciliini, 818, 829 89, 815, 818, 819, 828, 829, 830, 875 Acneus, 793, 807, 849, 850, 851, 852, 893

1203, 1255 Acrididae, 169,238,411,412, 414,415, 421-24,425 Acridinae, 412, 425 Acroneuria, 203, 432,453,487, 490, 492, 514 Acroneuria arenosa, 436,438,454,463, 465, 468 Acroneuria covelli, 488 Acroneuria evoluta, 85

Acroneuria lycorias, 85 Acroneuriinae, 514-15 Acronictinae, 787 Actaletidae, 169, 250, 254, 260 Actidium, 805

Activity traps, 21, 22, 28,41 Ademon, 922

Adephaga, 89, 182, 209, 210, 792, 794, 795,871-82, 924 Adicrophleps hitchcocki, 614,615,616, 664, 685,686, 748 Adverse outcome pathway, 142 Aedeomyia, 1074, 1079, 1082, 1084, 1086, 1087, 1092, 1096 Aedeomyia squamipennis, 1074 Aedes, 68, 187, 222, 226, 937, 976, 1071, 1073, 1075, 1079,1084, 1087, 1094 Aedes aegypti, 108 Aedes taeniorhynchus, 90 Aegialites, 801, 868, 891 Aegialomyia, 944 Aegilites, 807 Aeshna, 343, 349, 354, 367, 368, 374, 387, 396

Aeshna cyanea, 343 Aeshna grandis, 343 Aeshna interrupta, 388 Aeshnidae, 84, 98-99, 119, 135, 169, 199, 237, 238, 342, 343, 344, 347, 349, 350, 353, 354, 363-68, 369, 370, 371, 374, 388, 396-98 Afghanurus, 269, 270, 284, 285, 307, 312, 328

Afrolimnophila, 1024 Agabetes, 818, 830, 832 Agabetes acuductus, 819, 876 Agabetinae, 830 Agabetini, 818 Agabinae, 818, 820, 829, 830 Agabini, 820

Agabinus, 820, 830,832,876 Agabus, 89, 815, 830, 876 Agahus hipustulatus, 61, 910, 911 Agabus crassipes, 832 Agabus erichsoni, 104 Agapetinae, 687, 734 Agapetus, 87, 217,601,618,619, 687,734

Agapetus bifldus, 92, 102 Agapetusfuscipes, 102 Agapetus illini, 87 Agapetus occidentis, 87 Agapetus walkeri, 688 Agarodes, 587, 604,654,660, 678, 723, 746 Agarodes griseus, 662, 723 Agarodes libalis, 88,654 Agasicles, 793,869 Agasicles hygrophila, 868, 901 Agathon, 227, 932, 972,983 Agathon comstocki, 964 Agathon elegantulus, 935 Agkistrocerus, 1002 Agnetina, 451,487,492,514 Agnetina annulipes, 489 Agnetina capitata, 85, 452 Agraylea, 593,601, 627,629, 667,676, 695, 697, 736 Agraylea multipunctata, 628 Agrenia, 253, 255,259 Agrenia bidenticulata, 255,259 Agriidae, 405 Agrion, 405 Agrionidae, 135,405 Agriotypidae, 106 Agriotypinae, 909 Agriotypus, 96, 106, 909, 912,913 Agrypnia, 188, 219,605, 649, 715,760 Agrypnia colorata, 716 Agrypnia glacialis, 717 Agrypnia improba, 664 Agrypnia vestita, 648,650, 716 Ahautlea, 560

Akephorus, 873 Alabameuhria, 849 Alaskaperla, 455, 482 Alaskaperla ovihovis, 455,458, 482, 520 Aleochara, 887 Aleocharinae, 847 Alisotrichia, 626,695

1455

1456

Index

n

Alisotrichia arizonica, 625,697, 736 Allacma, 256, 257 Allan grab, 26, 39 Allenhypes vescus, 333

Amphinemura wui, 440,463 Amphinemurinae, 509-10 Amphizoa, 209, 796, 797, 803, 805,

Allenhyphes, 287, 289, 290, 305, 315 Allocapnia, 68, 181, 190, 202, 204, 446,466,

Amphizoidae, 49, 167, 209, 794, 796, 797,

470,472,512

Allocapnia granulata, 438, 441,471 Allocapnia nivicola, 85 Allocapnia rickeri, 85 Allocapnia virginiana, 441, 463 Allocladius, 1176, 1213, 1255

Allocosmoecuspartitus, 640,643,644, 703, 751

Allognosta, 939, 942, 1000 Allomyia, 218,605,613,686, 746 Atlomyia cascadis, 683 Allomyia scotti, 612, 657 Allonemohius, 418, 422, 428 Allonemohiusfasciatus, 422

Alloperla, 453,485, 520 Alloperla imhecilla, 454 Allopsectrocladius, 1196, 1199 Allotrichoma, 950, 1010 Allotrichoma simplex, 953, 955 Alluaudomyia, 985 Alnus ruhra, 122

Alotanypus, 1130, 1136, 1186, 1188, 1245 Alotanypus venustus, 1131, 1132 Alternanthera, 793

Alternantheraphiloxeroides, 868 Alysllnae, 909 Ambient surveys, bloassessment, 155 Ambiinl, 765 Amblopusa, 847, 888 Amhlycorypha, 417, 427 Amhlycorypha oblongifolia, 417,420 Amblypsilopus, 993 Ambrysinae, 538 Amhrysus, 206, 538, 541, 558 Ambrysus mormon, 542 Ameletidae, 83, 167, 195, 268, 274, 277, 279, 303, 304, 307, 321 Ameletus, 83, 195, 268, 274, 277, 279, 303, 304, 307,321 Amercaenis, 291, 334 Americahaetis, 280, 292, 310, 311, 323

Americabrya, 250 Americaenis, 315

Ametor, 838, 839, 844, 884

Ametropodldae, 167, 195, 198,274, 275, 301,302, 327

Ametropus, 195, 198, 274, 275, 301, 302, 327

Amiocentrus aspilus, 615, 616, 664,685, 687, 748

Amphiagrion, 358, 361, 406 Amphiagrion saucium, 360, 364 Amphicosmoecus canax, 640,643,659, 702, 703, 751

Amphinemura, 85, 442, 472,510 Amphinemura banksi, 437, 440 Amphinemura delosa, 475 Amphinemura nigritta, 100

809, 871 803, 805, 809, 871

Ampumixis, 860, 861, 863, 864 Ampumixis dispar, 896 Anabolia, 606,640, 709, 751 Anabolia bimaculata, 641, 644, 708 Anacaena, 838, 839, 844, 846, 884 Anacaenini, 844 Anacroneuria, 432,451, 454,487,489, 490, 514 Anacroneuria litura, 489,491

Anafroptilum, 269, 276,279, 281, 303, 323 Anafroptilum conturbatum, 281 Ancifroptilum minor, 294, 295 Anafroptilum ozarkense, 311 Anafroptilum pecatonica, 303 Anagapetus, 618, 619.687, 734 Anagapetus bernea, 87, 92,688 Anagapetus debilis, 668 Anagrus, 910, 913 Anagrus subfuscus, 919 Analetris, 75, 195, 199, 268 Analetris eximia, 21A, 211, 301, 303, 321 Analyzing stream samples, 18-20 Anaphes, 910,913

Anaphes gerrisophagus, 915,919 Anaphes victus, 913 Anatopyniini, 1128 Anax, 199, 343, 367, 397

Anax imperator, 98 Anaxjunius, 58, 84, 370 Anaxipha, 417,422, 428 Anaxipha exigua, 422 Anchycteis, 802, 807, 853, 854 Anchycteis velutina, 899 Anchytarsus, 853, 854 Anchytarsus bicolor, 89, 900 Ancyronx variegatus, 89 Ancyronyx, 806,858, 859, 862, 863 Ancyronyx variegatus, 896 Androprosopa, 223, 228, 992 Anepeorus, 268,269 Atiepeorus rusticus, 283, 310, 316, 328 Angarotipula, 1024, 1025, 1035, 1068 Angarotipula illustris, 1037 Anisocentropus, 590,608 Anisocentropus pyraloides, 616,617, 661, 682,687, 740

Anisodactylus californicus, 873 Anisops, 54 Anisoptera, 53, 54, 55, 81, 84, 342, 343, 344, 345, 346-47, 348, 350, 353, 36391,393-104 Annelida, 177

Annulipalpia, 585, 586, 592, 596, 726 Anodocheilus, 816,823, 825, 876

Anopheles, 227, 937, 1071, 1072, 1073, 1075, 1077, 1079, 1082, 1087, 1088, 1094

Anoplius depressipes, 909,924 Anoplodonta nigrirostris, 1000 Anostostomatidae,411,412 Antarctoperlaria, 181 Anthicidae, 793, 795, 800, 801, 809, 868

o

rs

Anthicus, 801

Anthomyiidae, 1016, 1017 Anthopotamus, 197, 271, 272, 273, 297, 300, 337 Antichaeta, 949, 1019 Antichaeta borealis, 951 Antichaeta testacea, 951 Antillocladius, 1174, 1175, 1202, 1229, 1255 Antocha, 223, 925, 1023, 1025,1027, 1028, 1029,1033, 1038, 1066

Antocha opalizans, 1058 Antocha saxicola, 90, 965, 1033 Anurkla, 247, 250, 251, 258

Anurida ashbyae, 245, 246

rs

Anurida calcarata, 246 Anurida mara, 246

Anurida maritima, 245, 248 Anuridella, 250, 258 Anuridella calcarata, 258

Anurophorus, 252 Apanisagrion, 361

Apanisagrion lais, 358, 364, 366, 407 Apatania, 605, 613,665,674,677,679, 686, 747

Apatania arizona, 612 Apataniafimbriata, 115 Apatania zonella, 683 Apataniidae, 167, 218, 220, 586, 589,605, 607, 611-13, 657,663,665,670,674, 675, 677, 679,681,683,686, 746-47 Apatolestes, 943, 1002 Apedilum, 1154, 1156, 1158, 1224, 1264 Apedilum elachistus, 1120 Aphelocheirus, 48, 51 Aphelocheirus aestivalis, 45,61 Aphrosylus, 996 Aphylla, 75, 343, 372, 376, 394 Aphylla williamsoni, 341, 374, 388 Apobaetis, 274, 278, 281, 309, 323 Apodesmia, 922 Apometriocnemus, 1255 Aprostocetus, 913,919 Apsectrotanypus, 1130, 1131, 1136,1186,

r

1188, 1245 Apsilops, 912,923

Apsilops hirtifrons, 909,917 Aptenopedes, 421, 425 Apteraliplus, 812, 813, 815 Apteraliplus parvulus, 874 Aquarium rearing method,40 Aquarius, 523, 535, 536, 537, 548, 554 Aquarius remigis, 537 Aquatic net, 18, 19, 21, 22 Arachnomorpha, 177 Araeopidius, 214, 853 Archanara, 785

Archeognatha, 177 Archilestes, 355,405

r

Index

Archilestes grandis, 98, 357, 360 Archisotoma, 253, 259 Archisotoma hesselsi, 245 Archisotoma interstitialis, 246

Archisotoma polaris, 246 Archistoma hesselsi, 255 Arcolamalloi, 767

Arctoconopa, 1023, 1051, 1062 Arctoconopa carbonipes, 1053 Arctocorisa, 532, 534, 559 Arctocorisa chanceae, 533 Arctocorisa sutilis, 533

Asynarchus rossi, 711 Asyndetus, 994 Atelocerata, 177 Atelopodella, 1174, 1175 Athericidae, 91, 168, 224, 229, 925, 926, 927, 928, 929, 930, 941, 942, 963, 967, 973,977, 981,993 Atherix, 224, 229, 941, 942, 977, 981,993 Atherix lantha, 91

Atherix pachypus, 967 Athyroglossa, 1010

1457

Baetodes, 216, 309,311,324 Baetopus, 302 Baetopus trishae, 278, 279, 324 Bag sampler, 19, 20 Bagous, 211, 869, 903 Bagous americanus, 869 Bagous longirostris, 869 Banksiola, 588,649, 674,678, 715, 761 Banksiola crotchi, 87, 717 Banksiola dossuaria, 648,650 Barbaetis, 302

Arctodiamesa, 1251

Atissa, 1013

Arctodiamesa appendiculata,

Atopsyche, 217,601,603,620,622,666,

Barhaetis benfleldi, 218, 292, 324 Barcode sequencing, 156 BasiaeschnaJanata, 363, 368, 369,

667,668, 676, 690, 735 ,4 tractelmis, 860, 863

Basket-type artificial substrate sampler,

1177, 1179

Arctopelopia, 1133, 1134, 1138,1140, 1189, 1190, 1247

Arctoperlaria, 181 Arctopora, 590, 640, 641, 709, 751 Arctopora trimaculata, 708 Arctopsyche, 78, 216,622,691, 726 Arctopsyche grandis, 87,692,693 Arctopsyche irrorata, 87, 621 Arctopsychinae, 691,726 Arctotipula {Tipula subgenera), 1023, 1024, 1031, 1035, 1041 Arctotipula salicetorum, 1032 Arcynopteryx, 457, 495 Arcynopteryx dichroa, 495, 496, 516 Argia, 200, 358, 361,407 Argia extranea, 360 Argia moesta, 347, 365 Argyra, 994 Argyractini, 765, 766 Argyractis, 765, 769, 770, 771, 773 Argyractis drumalis, 767, 776, 779 Argyractis subornata, 765, 767, 768 Argyrotaenia ivana, 788 Arigomphus, 373, 377, 394 Arigomphus pallidus, 375 Arigomphus villosipes. 381 Arrhopalites, 245, 256 Arsapnia, 446,470, 472, 513 Arsapnia decepta, 473 Arthoplea, 265 Arthroplea, 195, 268 Arthroplea bipunctata, 281, 282, 310, 331 Arthropleidae, 167, 195, 282, 302, 310, 331

Arthropoda, 177 Articulata, 177 Artificial stream rearing design,40 Artificial substrate, 28 Artificial substrate samplers, 18, 20, 21 Arzama, 785 Aschiza, 929 Ascllus, 134 Asheum, 1152, 1226 Asheum beckae, 1153 Asioplax, 270, 271 Asobara, 922

Aspidogyrus, 912 Asynarchus, 638, 640, 711, 752 Asynarchus montanus, 644, 710

A tractelmis wawona, 896

Atrichomelina puhera, 1019 Atrichopogon, 939, 940, 984 Attaneuria, 451, 487, 492 Attaneuria ruralis, 451,454,487,492, 493, 494,514 Attenella, 289, 315, 331 Atylotus, 944, 1002 Atyphohelea macroneura, 985 Auleutes, 903 Austrotinodes, 216, 600, 604, 670,678 Austrotinodes texensis, 616, 617,672, 687,731 Autotrophic/Heterotrophic Index, 131, 132 Axarus, 1154, 1157, 1225, 1233, 1265 Axarusfestivus, 1155 Axelsonia, 253, 255 Axeisonia tubifera, 259 Axymyia, 221, 226, 227, 934 Axymyiafurcata, 935, 967, 976,983 Axymyiidae, 168, 221, 226, 227, 925, 928, 930, 932, 935, 963, 967, 973, 976,983 Axysta, 1014 Azola caroliniana, 868

B

371, 397 20, 39 Bathythrix, 910, 923 Batrachideinae,426 Baumannella, 459, 499, 502 Baumannella alameda, 459,468,499, 501, 502, 503, 504, 516 Beardius, 1154, 1156, 1222, 1265 Beckidia, 1122,1154, 1219, 1221, 1265 Beckidia tethys, 1153 Behningiidae, 75, 84,167, 195, 267, 272, 273,291,299,300,337 Belgica antarctica, 1119

Bellardina {Tipula subgenera), 1024, 1038, 1043 Bellura, 765, 767, 768,769, 770, 785 Bellura densa, 767

Bellura gortynoides, 88, 767, 768 Bellura obliqua, 114, 776 Beloneuria, 85, 432,453, 487,490,515 Beloneuria georgiana, 454,467,491 Belostoma, 206, 231, 522, 527, 528, 556 Belostoma americanus, 231 Belostoma bakeri, 526

Belostoma malkini, 86 Belostomatidae, 49, 51, 75, 86, 96, 102, 114, 136, 170,206,237, 240, 522, 525, 526, 527, 528, 555-56 Benacus, 527

Bactra, 788

Baeoctenus, 1161, 1202, 1204, 1255 Baeoctenus bicolor, 1163, 1166 Baetidae, 81, 83, 95, 96, 98, 134, 166, 167, 195, 198, 240, 264, 267, 268, 269, 274, 275, 277, 279, 281, 292, 294, 295, 301,302, 304, 307,311,318, 323-26

Baetis, 67, 75, 79, 83, 98, 111, 115, 190, 264, 265, 275, 277, 278, 280, 281, 292, 293, 304, 307, 310, 324 Baetis bicaudatus, 96, 110 Baetis phoebus, 269 Baetis rhodani, 97, 99, 188 Baetis tricaudatus, 264

Baetisca, 195, 198, 271, 273, 302, 335 Baetisca rogersi, 84, 264, 301 Baetiscidae, 75, 84, 167, 195, 198, 271, 273, 301,302, 335

Benacus griseus, 556 Benhingidae, 97 Beothukus complicatus, 648,649,659, 715, 716,761 Beraea, 218, 587, 610,611,613,612,673, 680,686, 739 Beraea gorteba, 614, 664 Beraeidae, 168, 218, 586, 587, 589, 599, 610,611,613,614,664,672,673, 680, 684, 686, 739 Beringotipula {Tipula subgenera), 1024, 1036

Berlese-Tullgren funnel, 40 Berosini, 841 Berosus, 213, 793, 837, 838, 841, 843, 885 Bethbilbeckia, 1134,1136,1137, 1188, 1228 Bezzia, 939, 940, 985

1458

Index

Bezzia varicolor, 966

Biaraxis depressa, 888 Bihiocephala, 221, 932 Bihiocephala grandis, 935,983 Bidessini, 817, 823 Bidessonotus, 816, 822, 824, 825, 876 Bilyjomyia, 1139, 1245 Bioanalytical screens, 142 Bioassessment, 141-64 causal assessment, 161 community-based, 162-63 data, quality assurance for, 155-56 defined, 141 history, 142 indices, 144-49 overview, 141 scales, 142-44 study design, 152-55 with molecular methods, 156-60 Bioinformatics, 156 BLsancora, 453,485, 520

Bisancora rutriformis, 456, 488 Bittacomorpha, 222, 226, 934, 971,992 Bittacomorpha clavipes, 937 Bittacomorphella, 934,992 Blaesoxipha, 1017 Blattodea, 180 Bledius, 847, 848, 888

Blepharicera, 932, 983 Blepharicera tenuipes, 935 Blepharicera williamsae, 90 Blephariceridae, 49, 55,71,90, 119, 167, 190, 221, 227, 243, 925, 926, 927, 928, 930, 932, 935, 957, 964, 969, 972, 983, 1028

Blephariceromorpha, 185 Biera, 1021 Blissia, 252

Bolhodimyia alrata, 1002 Bolotoperla, 442, 464 Bolotoperla rossi, 445, 464,468, 509 Bolshecapnia, 446, 464, 472,513 Bolshecapnia sasquatchi, 475 Bolshecapnia spenceri, 447,449 Bonetograstrura, 249 Bonetrura, 253, 257 Booneacris, 423,426 Borenectes, 817, 827 Boreochlini, 1177,1235,1250 Boreochlus, 1124, 1177, 1181,1183, 1184, 1233. 1250

Brachycentrus, 74, 77, 87, 110, 130, 218, 588, 589, 609,613,614,674,679, 686, 748

Brachycentrus americanus, 662,685 Brachycentrus chelatus, 662 Brachycentrus echo, 614 Brachycentrus etowahensis, 662 Brachycentrus numerosus, 662 Brachycentrus spinae, 87, 111 Brachycera, 928, 931, 939, 969 Brachycercus. 269, 289, 311,317, 334

Brachycercus nitidus, 295 Brachydeutera, 954, 1007 Brachydeutera argentata, 956, 970 Brachymeria podagrica, 913 Brachymesia, 383, 390,401 Brachymesia gravida, 381 Brachypogon, 985 Brachypremna, 1024 Brachypremna dispellens, 1031, 1032, 1033, 1036, 1068

Brachypterainae, 509 Brachystomella, 249, 251 Brachystomella honda, 258 Brachyvatus, 817, 823,825 Brachyvatus apicatus, 876 Bracon, 909,922 Braconidae, 170, 243, 913, 914, 917, 922-23 Braconinae, 909 Brechmorhoga, 383, 390, 401 Brechmorhoga mendax, 381 Brennania, 1002 Brevitentoria, 739-46 Brillia, 76, 1164, 1166, 1170, 1194, 1195, 1196, 1197, 1255

Broscus cephalotes, 872 Brundiniella, 1131, 1134, 1137, 1187, 1189, 1245

Brychius, 812,813,815,874 Brychius hungerfordi, 812 Bryelmis, 856, 860, 863, 897 Bryobiota, 888 Bryophaenocladius, 1161, 1164, 1172, 1200, 1203, 1255 Bryothinusa, 847 Bryothinusa catalinae, 888 Buchonomyiinae, 1230 Bucrates, 417, 420,427 Buenoa, 54, 73, 74, 538, 543, 562 Buenoa scimitra, 543

Boreoheptagyia, 1176, 1178, 1202, 1205.1251

Box-type sampler, 18 Boyeria, 363, 367, 397 Boyeria vinosa, 369

Caelifera, 412 Caenidae, 75, 83,119, 134, 167,195, 198, 237, 264, 267,272, 275, 295,296, 303, 304, 305,311,334-35 Caenis, 53, 83, 195, 198, 264,275, 291, 304, 311,315, 334

Brachinus, 796

Caenis arnica, 83

Brachybamus electus, 904 Brachycentridae, 77, 87, 167, 218, 586,588, 589, 596, 599, 603, 609,613-16,662, 664,674,675, 679, 685,686-87,747-48

Cafius, 888 Calacanthia, 540, 543 Calacanthia trybomi, 565 Caladomyia, 1271

Boreoheptagyiini, 1176, 1235, 1251 Boreonectes, 816, 817 Bourletiella. 247, 256, 257, 261

Calamoceratidae, 88, 104, 167, 218, 586, 589, 590,607, 608,616,617,661,671, 675,680, 682,684,687, 740 Calileuctra, 448,476,479, 512

Calileuctra dobryi, 465, 481, 483 Calileuctra ephemera, 452 Calineuria, 451,487,489,490 Calineuria californica, 85,451, 487,490, 491,493,515 Callibaetis, 67, 198, 265, 277, 278, 279, 281, 303, 324 Callibaetisfloridanus, 83,98 Callicera, 1021 Callicorixa, 74, 532, 533, 559 Callicorixa audeni, 534 Callicorixa vulnerata, 533

Callinapaea, 1007 Callineuria, 204

Calliperla, 457, 497,502 Calliperla luctuosa, 457,460,497, 500, 502, 503,518 Calocoenia, 1007

Caloparyphus, 941, 1000 Caloparyphus major, 942 Calopsectra, 1273 Calopsectrini, 1271 Calopterygidae, 169, 189, 190,201, 342, 348, 349, 351,355,357,405 Calopteryx, 200, 201, 349, 355,405 Calopteryx aequabilis, 342 Calopteryx angustipennis, 351, 357 Calopteryx maculata, 357 Calopteryx virgo, 61 Camelobaetidius, 265, 276, 279, 306, 307, 325

Campopleginae,909 Campsicnemus, 994 Campsurus, 265, 291, 317, 339 Camptocladius, 1174,1202, 1229, 1255 Canace, 946, 1004 Canace macateei, 948

Canaceoides, 946, 980, 1004 Canaceoides nudata, 948

Canacidae,49, 167, 926, 928, 930, 946,948, 974, 980, 1004 Canada balsam, 1125 Canister sampler, 18 Cannacria, 401

Cannaphila insularis, 382, 389, 391,401 Cantharidae, 793

Cantopelopia, 1139 Capnia, 446,466,472, 513 Capnia californica, 473 Capnia gracilaria, 469 Capnia lacustra, 65,429,470 Capnia lacustris, 234 Capnia vernalis, 441,449 Capniidae, 67,68,85, 134, 169, 202, 204,

r

240, 429,430,431,438, 439,440,441, 447, 449,462,463,469, 471,473,475, 512-13

Capnura, 446,466,470, 472, 513 Capnura elevata, 473 Capnura venosa, 449

r

r r /■

Index

Capnura wanica, 473 Capsula, 785 Carabidae, 167,182, 209, 793, 794, 795, 796, 797, 802, 803, 805, 872-73 Caraphractus, 910,913 Caraphractus cinctus, 910, 911, 913, 919 Cardiocladius, 1165, 1167, 1200,1203, 1209, 1210, 1256

Chaetarthriinae, 841 Chaetharthriinae, 844 Chaetharthriini, 841 Chaetocladius, 1168, 1170, 1202, 1205, 1211, 1256 Chalcididae, 170,910,913,914,919 Chalcidoidea, 909, 910, 913, 919-21 Chalcis, 910,913,919

Caricea, 1016

Chalcosyrphus, 1021 Chaoboridae, 49, 55, 73, 74, 90, 119, 150, 167, 221, 226, 925, 926, 927, 928, 929, 934, 938, 962, 965, 973, 976,988 Chaoborus, 73, 74, 188, 221, 226, 934, 976,988 Chaoborus americanus, 14, 90, 938,965 Chaoborusflavicans, 74

Carpelimus, 847, 888 Cascadoperla, 457,497, 502 Cascadoperla trictura, 457, 460, 500, 502, 503, 518 Catatasina, 941

Caudatella, 286, 314, 332 Cautochironomus, 1219

Caurinella, 268,293 Caurinella idahoensis, 271, 288, 315, 332 Causal Analysis/Diagnosis Decision Information System (CADDIS), 161 Causal assessment, 161

CDC light trap,40 Cecidomyiidae, 1028

Chaoborus punctipennis, 74, 90 Chaoborus trivittatus, 90 Chara, 429 Chasmatonotus, 1174, 1213, 1256 Chathamiidae, 96, 104 Chauliodes, 571, 574, 575, 577, 578, 583

Chauliodes pectinicornis, 573

Celina, 209, 814, 816, 822, 829, 876 Celithemis, 383, 390, 401 Celithemis elisa, 84, 381 Celithemisfasciata, 84

Chauliodinae, 583-84 Cheilotrichia, 1051, 1062 Cheiltrichia cinerescens, 1055 Chelifera, 946, 947,997

Celithemis verna, 386

Chelipoda, 997 Chematopsyche pettiti, 81 Chernokrilus, 459,492, 516 Chernokrilus misnomus, 459, 461,492, 496 Chemovskiia, 1145, 1219, 1265 Cheumatopsyche, 75, 102, 621,623,624,

Cell assays, 142 Cenocorixa, 529, 532, 533, 560 Cenocorixa kuiterti, 533 Centrarchidae, 343

Centrohiopsis odonatae, 920 Centrocorisa, 530, 531

Centrocorisa nigripemis, 560 Centroptilum, 190, 269 Centroptilum triangulifer, 83 Ceraclea, 104, 187, 218, 587, 589, 608, 630, 631,632,664,698,741 Ceraclea ancylus, 190 Ceraclea cancellata, 699 Ceraclea excisa, 88 Ceraclea maculata, 631 Ceraclea transversa, 88

Ceratocombidae, 170, 521, 553 Ceratocomhus, 553 Ceratophyllum, 766, 774, 812

Ceratophysella, 194, 249 Ceratopogon, 939, 986 Ceratopogonidae, 37, 49, 136, 167,221, 227, 344, 926, 927, 928, 929, 936, 940, 963, 966, 973, 977,984 Ceratopogoniidae, 90 Ceratopogoninae, 939, 985-87 Cercobrachys, 269, 291, 296, 315, 335 Cercobrachys etowah, 296, 305 Cercyon, 836, 885 Ceriana, 1021

Ceriodaphnia dubia, 151 Cernotina, 652,656, 718, 731 Cernotina spicata, 651 Ceropsilopa, 1005 Chaenusa, 922 Chaetarthria, 837, 841, 885

625, 691,727

Cheumatopsyche analis, 87,693 Cheumatopsyche pasella, 87 Chilo, 767, 784 Chilostigma itasca, 707 Chilostigma itascae, 635,637, 706 Chilostigmodes, 633 Chilostigmodes areolatus, 633, 705,706, 752

Chiloxanthinae, 540 Chiloxanthus, 540, 545, 565 Chimarra, 86,604, 646,647, 712, 730 Chimarra aterrima, 86 Chimarra ohscura, 714 Chionea, 1028 Chioneinae, 1023, 1026, 1027, 1062-64 Chironomidae, 2, 37, 49, 53, 55, 73, 75, 76, 77, 80, 81, 82, 90-91, 92, 95, 108, 112, 115, 125, 127, 150, 166, 167, 186, 221, 227, 234, 243, 925, 926, 927, 928, 929, 931, 936, 940, 962, 963, 966, 973, 977,1119-1274 Chironominae, 119, 187, 1126, 1213-29, 1230, 1235, 1236, 1240, 1264-74

Chironomini, 77, 119, 134, 977, 1122, 1123, 1124, 1127, 1145-59,1230, 1235, 1264-71

1459

Chironomus plumosus, 57, 90, 108 Chironomus riparius, 90 Chironomus tenuistylus, 90 Chironomus thummi, 57 Chlaenius, 209, 797

Chloroperlidae, 86,169, 203, 204, 429, 431, 438,439,440,443, 454,456, 458, 462,467,484,486, 488, 519 Chloroperlinae, 520 Chlorotabanus, 943

Chlorotabanus crepuscularis, 967,1002 Chorebus aquaticus, 917,922 Choristoneura, 787

Choroterpes, 111, 284, 308, 313, 335 Chorthippus, 425 Christobella, 250

Chromagrion, 345 Chromagrion conditum, 358, 363, 407 Chrysendeton, 766, 769, 771, 780 Chrysendeton medicinalis, IK) Chrysobasis, 362 Chrysobasis lucifer, 359, 362 Chrysogaster, Ti, 946, 948, 1021 Chrysomelidae,49, 73, 89, 167, 189,211, 792, 793, 794, 795, 798, 801, 807, 809, 868-69, 901-3 Chrysops, 73, 91, 229, 943, 967, 978,1002 Chrysops cincticornis, 945 Chrysops excitans, 945 Chrysops furcatus, 945 Chrysotimus, 994 Chrysotus, 994 Chyranda centralis, 634,636,657, 708, 709, 752 Cicadellidae, 524, 526 Cinygma, 283, 285, 307, 310, 328 Cinygmula, 79, 283,285, 307, 312, 329

Cinygmula ramaleyi, 83 Cinygmula reticulata, 83 Cirrula, 950, 1007 Cirrula austrina, 955 Cirrula gigantea, 955 Claassenia, 451, 485,490 Claassenia sahulosa, 85, 451,452, 485, 489, 490, 493,514 Cladopelma, 1147, 1150, 1218, 1219, 1266

Cladotanytarsus, 1142, 1143, 1144, 1216, 1217, 1272

Cladotanytarsus mancus, 90 Clanoneurum americanum, 1005 Clasiopella uncinata, 1005 Cleptelmis, 860, 861, 863,864

Cleptelmis addenda, 897 Climacia, 208, 569, 578, 579, 580, 584 Climacia areolaris, 86, 580 Climacia californica, 578 Clinocera, 946, 947, 968, 998 Clinocera stagnalis, 968 Clinocerinae, 944,946 Clinohelea, 986

Chironomus, 54, 73, 82, 90, 112, 227, 940,

Clinotanypodini, 1127

966, 1120, 1155, 1158, 1159,1222, 1223, 1224, 1226, 1239, 1265 Chironomus attenuatus, 58

Clinotanypus, 1127, 1129, 1187, 1189, 1233, 1244

Clioperla, 203,497, 499

1460

Index

Clioperla ctio, 85, 443,497,499, 500, 503,518 Clistoronia, 638, 711, 752 Clistoronia maculata, 710

Clistoronia magnifica, 88, 104,639,658 Cloeodes, 280,310,311,325 Cloeon, 70, 265,281,306

Cloeon dipterum, 46, 53, 265,278, 295, 325 Clogmia, 936, 990 Clostoeca disjuncta, 635, 636,637,665, 707, 709, 752 Clunio, 66, 1236, 1241, 1253

Conocephalus, 205,413,415,427 Conocephalus attenuatus, 419 Conocephalus spartinae, 413 Constempellina, 1139,1142,1215, 1216, 1217, 1272 Contacyphon, 849 Contiger, 766, 769, 771 Contiger vittatalis, 780 Copelatinae, 818, 829 Copelatus, 818, 819, 828, 829, 830, 877 Copelatus glyphicus, 89

Clunionini, 1236, 1240, 1241, 1253-54

Copiocerinae, 411 Coptotominae, 817, 830 Coptotomus, 89, 805, 815, 817, 819, 822,

Cnephia, 976, 1104, 1106, 1113, 1116 Cnephia dacotensis, 1103, 1108, 1114

Coquillettidia. 50, 73, 926, 1073, 1075,

Clunio marinus, 191

Cnodocentron, 722

Cnodocentron yavapai, 666, 723, 733 Coelamhus, 826

Coelomomycetaceae, 1120 Coelopa, 1004 Coelopa frigida, 960 Coelopidae, 168, 926, 954, 960, 1004 Coelopina anomala, 1004 Coelotanypodini, 1244 Coelotanypus, 1127, 1129, 1189, 1239, 1244 Coelotanypus concinnus, 1129 Coenagrion, 359, 362,407 Coenagrion interrogatum, 362 Coenagrion puella, 53 Coenagrionidae, 53, 84, 135, 169, 200, 240, 343, 344, 345, 347, 348, 350, 351, 352, 355, 358-63, 360, 364, 365, 366, 387,

828, 830, 832,877 1077, 1079, 1087, 1095

Coquillettidia perturbans, 1075 Cordilura, 91,961,980,1018 Cordiluridae, 240, 1017 Cordulegaster, 199, 201, 354, 379, 399 Cordulegaster boltonii, 61 Cordulegaster maculata, 356, 388 Cordulegastridae, 135, 169, 188, 199, 201, 342, 343, 350, 353, 354, 356, 379, 388, 399 Cordulia shurtleffi, 380,400 Corduliidae, 84, 100, 169, 199, 201, 342, 343, 350, 353, 354, 356, 379-80, 381, 387, 399-400 Core sampler, 20, 21,23

Core sampler with pole, 38

Coenia curvicauda, 959

Corethrella, 222, 227, 934, 976, 989 Corethrella brakeleyi, 938, 965 Corethrellidae, 168, 222, 227, 928, 929, 934, 962, 965, 973, 976,989 Corisella, 529, 530, 531, 532, 560

Coleophora, 788 Coleophoridae, 170, 767, 788 Coleoptera, 2, 3-6, 13, 37, 49, 51, 53, 65, 67, 71, 72, 73, 76, 80, 82,89, 97, 104-

Corixidae, 49, 55, 86, 100, 119, 133, 166, 170, 206, 237, 240, 521, 522, 523, 524, 525, 527, 528, 529-32, 533-34, 549,

406-9

Coenagrioninae, 355, 359 Coenia, 954, 1008

6, 113, 115, 119, 120, 125, 126, 127, 166, 167-68, 174, 176, 181, 182, 188, 189, 190, 209-14, 232, 234, 236, 238, 239,242,243,791-908,910

Collembola, 2, 37, 71,120, 166, 169, 176, 194,232, 233, 241,242,245-61 CoHophora, 256 Collops, 801

Corisella decolor, 533

559-61

Corixidea major, 553 Corixinae, 527 Corixini, 530

Cornops aquaticum, 411 Corydalidae, 55, 86, 95,102, 135, 170, 190,

Cosmopterygidae, 215 Cossidae, 767 Cosumnoperla, 457, 497, 505, 519 Cosumnoperla hypocrena, 460, 500, 504 Cosumnoperla sequoia, 504 Counting, 34 Crambidae, 88, 106, 136, 170, 215, 238, 241, 243, 765, 767, 770, 772, 773, 774, 775, 776, 777, 779 Crambinae, 767, 770, 777, 784 Crambus, 767 Cremastinae, 909 Crenitis, 837, 844, 885 Crenitulus, 837, 844, 885 Crenitulus suturalis, 844 Crenitus, 840

Crenophylax sperryi, 642, 710, 711, 753 Cressonomyia, 1006 Cricotopus, 90, 221,1162,1164, 1168, 1169, 1172, 1174, 1198, 1201, 1206, 1207, 1211, 1214, 1233, 1241, 1256 Cricotopus bicinctus, 90, 1172

Cricotopus triannulatus, 90 Cricotopus trifascia, 90, 1168, 1169 Crinodessus, 816, 824

Crinodessus amyae, 877 Crocothemis servilia, 384, 389, 401 Crustacea, 66, 177

Cryphocricinae, 538 Cryphocricos, 523, 538, 541 Cryphocricos hungerfordi, 86, 558 Cryptinae, 909 Cryptochia, 633, 698, 753 Cryptochia pilo,sa, 636, 663, 701 Cryptochironomus, 1147, 1150, 1219, 1221, 1237, 1266

Cryptochironomus Marina, 1150 Cryptolabis, 75, 1023, 1027, 1034, 1045, 1047, 1062

Cryptopygus, 253 Cryptostemma, 553 Cryptostemmatidae, 553 Cryptotendipes, 1147,1150, 1219, 1221, 1266

Ctenophora, 1031 Ctenophorinae, 1024 Culex, 187, 1071, 1073, 1079, 1080, 1084, 1086, 1087, 1095 Culex nigripalpus, 90 Culex pipiens, 57,67, 108

Colobaea americana, 1019

208, 237, 240, 241, 569, 570, 571, 572, 573, 574-78, 579,582 Corydalinae, 582-83

Colohurella, 252

Corydalus, 54, 208, 570, 571, 572, 573, 574,

Culex restuaizs, 90

Colymheles, 89, 815,820, 830, 832,876 Colymhetes sculptilis, 106 Colymbetinae, 820, 830

575, 577, 579, 583 Corydalus cornutus, 86, 114, 573

Culicidae, 37, 48,49, 50,68, 71, 73, 76, 80,

Comaldessus, 816, 823 Comakkssus stygius, 876 Community-based bioassessment, 162-63 Comparator sites, 161 Compteromesa, 1180, 1252 Compterosmittia, 1256 Conchapelopia, 1129, 1133, 1134, 1135, 1138, 1140, 1186, 1188, 1247 Condicinae, 786 Conocephalinae,427

Corynocera, 1139, 1142, 1144, 1216, 1217, 1272

Corynoneura, 90, 1159, 1163, 1190, 1191, 1233, 1254 Corynoneurini, 1254 Corynothrix, 252, 254 Coryphaeschna, 363, 368, 397 Coryphaeschna adnexa, 367 Coryphaeschna ingens, 369, 371 Cosmopterigidae, 170, 767, 788-89 Co.smopterix, 789

81,90, 108, 115, 119, 167,222,226, 227, 234, 238, 241, 925, 926, 927, 928, 929, 930, 934, 937, 962, 973, 976, 1028, 1071-96 Culicoides, 227, 939, 940,986 Culicoides variipennis, 90 Culicoides venustus, 966

Culicomorpha, 185, 186 Culiseta, 1071, 1073, 1079, 1080, 1084, 1086, 1087,1096

Culoptila, 618, 690, 735 Culoptila hamata, 688

Index

Culoptila moselyi, 619 Culoptila thoracica, 688 Cultus. 459,497, 499, 502, 505, 516 Cultus aestivalis, 461, 501, 503 Cultus verticalis, 503 Curculionidae, 49, 73, 89, 166,167, 189, 211,242, 243, 792,793, 794, 795, 798, 800, 801, 802, 805, 869, 903-8 Curicta, 538, 542, 557 Curictini, 538 Cushing-Mundie drift sampler, 19 Cybister, 805, 814, 815, 818, 819, 828, 829, 877 Cybister fimbriolatus, 89 Cybistrini, 818,827 Cyclorrhapha, 928,929 Cyclorrhaphous, 939

Demicryptochironomus, 1147, 1150, 1223, 1224, 1266 Demijerea, 1219, 1266 Denisiella, 256

Denopelopia, 1134, 1137, 1138, 1185, 1228, 1234, 1247 Dentatella, 271 Dentatella coxalis, 271

Derallus, 837, 838, 840, 841, 843 Derallus altus, 885 Deronectes, 880, 881

Derotanypus, 1128, 1135, 1188, 1228,1245 Derotanypus alaskensis, 1131, 1132 Derovatellus, 814, 816, 823 Derovatellus lentusfloridanus, 877 Desmona, 634, 706, 753 Desmona bethula, 88, 636,639, 657, 705

Cylindrical-type artificial substrate sam pler, 20 Cylindrotomidae, 168, 223, 1023, 1026,

Desmopachria, 814, 816, 824, 825,

1027, 1028, 1032, 1057, 1061 Cylindrotominae, 1061 Cylloepus, 858, 859, 865, 866, 897 Cymatia, 527, 528 Cymatia americana, 560 Cymatiinae, 527 Cymbiodyta, 838, 839, 841, 846, 885 Cynipoidea, 909, 910, 914, 921 Cyphoderinae, 250 Cyphoderus, 250 Cyphomella, 1149, 1150, 1223, 1224, 1266 Cyphon, 802,848, 849, 851, 895 Cyphon bicolor, 849 Cyphon coUaris, 849 Cyphon comptus, 849 Cyphon concinnus, 849 Cyphon confinis, 849 Cyphon confusus, 849 Cyphon drymophila, 849 Cyphon ruficollis, 849 Cyrnellusfraternus, 651,652, 718, 719,731 Cyrtacanthacridinae,425 Cyrtobagous salvinae, 904

Despaxia, 439, 448,476,479 Despaxia augusta, 448, 450, 452, 476, 479,

828, 877 Desoria, 253

480, 483,512

Deuterophlebia, 90, 222, 226, 227, 932, 972, 981, 989

Deuterophlebia nielsoni, 935, 964 Deuterophlebiidae, 49,90, 168, 222, 226, 227, 925, 926, 927, 928, 929, 930, 932, 935, 957, 964, 969, 972, 981, 989 Deutonura, 250 Diachlorus, 943

Diamesa, 221, 1177, 1178, 1206, 1207, 1233, 1251 Diamesa incallida, 90, 112 Diamesa kohshimai, 69 Diamesa mendotae, 1119 Diamesa nivoriunda, 90

Diamesinae, 1127, 1176-77, 1180, 1182, 1191-1213, 1235, 1251-52 Diamesini, 1235, 1236, 1251 Dianous, 214

Diaphorus, 994 Diapria conica, 922 Diapriidae, 170, 909, 910, 912, 914, 915, 921-22

Diaprioidea, 921-22 Diaspididae, 524 D-frame aquatic net, 38 D-vac vacuum sampler,22 Dacnusa, 922

Dactylolabinae, 1023, 1026, 1064 Dactylolabis, 1024, 1026, 1030, 1046, 1064 Dactylolahis montana, 1048 Dagamaea, 253 Dannella, 271, 289, 315, 332 Dannella simplex, 311 Dasycorixa, 529, 530, 560 Dasyhelea, 939, 940, 984 Dasyhelea traverae, 966 Dasyheleinae, 984 Deinocerites, 1078,1079, 1080, 1083, 1084, 1087, 1091, 1095 Delphacidae, 524 Demeijerea, 1221

Diaulota, 847, 888 Dibolocelus, 837, 840, 841,843 Dibusa, 592 Dibusa angata, 87, 626,627,629,665,694, 697, 736 Dicercomyzon, 190 Dichaeta, 91, 108 Dichaeta caudata, 953, 959

Dicranomyia modesta, 1030, 1033 Dicranomyia simidans, 1047 Dicranophragma, 1050, 1064 Dicranophragmafuscovarium, 1053 Dicranopselaphus, 849, 850, 851, 852 Dicranopselaphus variegatus, 893 Dicranota. 90, 933, 1023, 1025, 1029, 1033, 1042, 1044, 1067 Dicranota bimaculata, 90 Dicranota subtilis, 1044

Dicrotendipes, 90, 1155, 1158, 1159, 1220, 1222,1223, 1266 Dictya, 226, 229, 949, 961, 979, 1019 Dictya expansa, 951 Dictya pictipes, 951 Diclyacium, 1019 Dicyrtoma, 256 Didymops, 356, 378, 379, 399 Diglotta, 847, 888 Dimecoenia, 950, 954, 1008 Dimecoeniafuscifemur, 960 Dimecoenia spinosa, 955 Dineutes, 89 Dineutes hornii, 72 Dineuticida dineutes, 920

Dineutus, 104, 210, 810, 811, 872 Diostracus, 994 Diphetor, 311

Diphetor hageni, 280, 293, 309, 325 Diphuia nitida, 1010 Diptax, 404 Diplectrona, 622,691, 727 Diplectrona modesta, 87,621,692 Diplectroninae, 586,691, 727 Diplocladius, 1160, 1171, 1208, 1209, 1256 Diploperla, 459, 497,499, 516 Diploperla duplicata, 461 Diploperla robusta, 500 Diplosmittia, 1257 Dipseudopsidae, 168, 216, 586, 591, 603, 604, 616, 617, 670,672,678,684, 687, 730

Dipsocoridae, 170, 521, 553 Dipsocoromorpha, 521 Diptera, 2, 13, 15, 37,49, 51, 53,67,68, 69,71,72, 73,74, 76, 77, 80, 82, 90-91,96, 97, 106-9, 113, 115, 119, 120, 125, 137, 166, 167-68, 174, 176, 185-86, 187, 188, 189, 190, 192, 22129, 232,233, 234, 236, 238, 239, 241, 242, 243, 791, 792, 909, 925-1022, 1071,1093-96, 1116-18 Discocerina, 950, 1011 Discocerina obscurella, 91, 953, 955 Discomyza, 1006

Dichromorpha, 414,423,425 Diclasiopa lacteipennis, 1010 Dicosmoecus, 113, 130, 639,640,643,644,

Discomyzinae, 1005 Disonycha, 868,902 Ditrichophora, 1011

703, 754 Dicosmoecus atripes, 665, 702 Dicosmoecus gilvipes, 88 Dicranomyia, 48, 1024, 1030, 1033,

Diura, 430, 459,497, 499, 502, 516 Diura bicaudata, 188 Diura knowltoni, 461, 500, 501 Diura washingtoniana, 501 Dixa, 936, 937, 975,989 Dixa aliciae, 966

1045, 1066

Dicranomyiafrontalis, 1047

1461

1462

Index

Dixella, 111. 934, 937, 966,989 Dixidae, 37, 167, 222, 925, 926, 927, 928, 929, 930, 934, 937, 962, 966, 973, 975,989

Djalmahatista, 1128, 1183, 1184, 1233, 1239, 1246

Djalmahatista pulcher, 1131, 1132, 1135 Doddsia, 439, 464 Doddsia occidentalis, 439, 444,464, 469, 509

Dohrniphora cornuta, 979, 1017 Doithrix, 1170, 1175, 1198,1228, 1257 Dolaitia, 75, 97, 195, 267 Dolania americana, 84, 265, 111, 273,291, 299, 300, 337

Dolichocephala, 946, 998 Dolichopeza, 1024, 1031, 1032, 1068 Dolichopeza americana, 1032 Dolichopeza walleyi, 1036 Dolichopezinae, 1068 Dolichopodidae, 49, 167, 224, 241,926, 927, 928, 929, 930, 944, 947, 961, 963, 968,974, 978, 993-97, 1027 Dolichopus, 961, 978,994 Dolomedes, 909

Dolophilodes, 591,646,647, 669,676, 712,730

Dolophilodes distincta, 86, 714 Dolophilodes novusamericana, 598, 714 Dolophiloides, 190 Donacaula, 767, 783 Donacia, 73, 89, 189, 211, 801, 807, 833, 868, 869, 902 Donaciella cinerea, 89 Donaciinae, 869

Doncricotopus, 1170, 1173, 1193, 1228, 1257 Dorocordulia, 380,400 Doroneuria, 451,487, 490, 515 Doroneuria baumanni, 454

Doroneuria theodora, 489,491, 493 Draeculacephala, 524 Drag-type samplers, 20 Drift net, 38 Dromogomphus, 373, 374, 376, 394

Dromogomphus spinosus, 374, 378 Drop trap, 21 Drosera, 767

Drosophila, 42,53 Drunella, 71, 83, 196, 275, 286, 287, 314,332 Drunella cornutella, 308 Drunella doddsi, 190

Drunella grandis, 279 Dryomyzidae, 168, 926, 928, 949, 974, 979, 1005

Dryopidae,49, 55, 65, 77, 80, 167, 182, 190, 211,234, 791, 792, 793,794, 800, 805, 806, 808, 853, 854, 855, 857, 894 Dryops, 854, 855, 894 Dryotribus mimrticus, 904 Dubiraphia, 856, 859, 863,864,897 Dubiraphia quadrinotata, 89 Dyschiriodes, 873

Dysmicohermes, 571, 574, 583 Dysmicohermes disjunctus, 576, 577 Dysmicohermes ingens, 575, 576 Dythemis, 383, 390,401 Dytiscidae, 45,49, 50, 51, 73, 77, 89, 104-6, 114, 119, 135,166, 167, 190, 209,238, 243, 791, 792, 794, 796, 802, 803, 805, 814,815-33,819, 836, 875-81, 909,911

Dytiscidae, 190 Dytiscinae, 817,827 Dytiscini, 818, 827 Dytiscus, 50, 89,209,815, 818, 827,877 Dytiscus verticalis, 58

Ecclisocosmoecus scylla, 635,637, 644, 701, 703, 754

Elophila, 215, 765, 766,768, 769, 770, 771, 774, 776, 780 Elophila gyralis, 766 Elophila icciusalis, 766, 768, 776 Elophila obliteralis, 766, 773, 775, 776 Elophila occidentalis, 111, 11A Emergence traps, 22, 24, 39 Empeda, 1055 Empedomorpha, 1024 Emphyastes, 805 Emphyastesfuciola, 904 Empididae, 49, 67, 91, 167, 224, 229, 926, 927, 928, 929, 944, 947, 963, 968, 974, 979, 997-99

Enallagma, 200, 359, 361, 362, 408 Enallagma aspersum, 84, 342 Enallagma civile, 351, 360 Enallagma divagans, 364 Enallagma hageni, 84

Ecclisomyia, 633, 636,657,677, 703, 754 Ecclisomyia maculosa, 701 Eccoptura, 451,487,490 Eccoptura xanthenes, 85, 434, 435, 441, 451,452,487,489,490, 494,515 Ecdyonurus, 70, 97, 269, 270

1226, 1267 Endonura, 250

Ecdyonurus insignis, 61

Endopterygota, 113, 181,232,233

Ecnomidae, 168, 216, 586,600,604,616, 617,670, 672,678, 684,687, 731

Endotribelos, 1156, 1158, 1159, 1220, 1229, 1267

Enallagma pollutum, 341 Enallagma vesperum, 341 Endeodes, 801, 802, 890-91 Endochironomus, 1151,1154, 1157, 1224,

Ecological tables, 165-74

Energetics, 137-39

Ectemnia, 1100, 1101, 1104, 1106, 1113, 1116 Ectemnia invenusta, 1103, 1105, 1107,1114 Ectemnia taeniatifrons, 1111, 1112 Ectopria, 214, 807, 849, 850, 851, 852, 894

Enicocerus, 845,846 Enlinia, 994 Enochrinae, 841 Enochrus, 213, 793, 837, 838, 841, 846,885 Ensifera, 412

Edmundsius agilis, 274, 303, 322

Entomobrya, 252, 260 Entomobrya arula, 246 Entomobrya laguna, 246, 254 Entomobryidae, 169, 194, 241, 250, 251,

Eichhornia, 767, 869 Einfeldia, 1154, 1158, 1159, 1220, 1222, 1223, 1224, 1225, 1226, 1266 Einfeldia natchitocheae, 1155

Einfeldia pagana, 90 Einfeldia synchrona, 90 Ekman grab, 17, 19,20, 22,23,24, 39 Elassoptes marinu.s, 904 Electroshocking, 18 Eleocharis, 767

Elgiva, 949, 1019 Elgiva rufa, 91 Elgiva solicita, 91,951,952 Ellipes, 421,427 Ellipes minutus, 416 Elliptera, 1024, 1046, 1048, 1066 Ellipteroides, 1024, 1051, 1062 Ellipteroides slossonae, 1055 Ellis-Rutter stream sampler, 18, 38 Elmidae, 49, 51,65, 81, 82, 89, 106, 133, 134, 166, 167, 182, 190, 211, 238, 243, 791, 792, 794, 799, 802, 806, 808, 853, 855, 856-68,896-99

Elodea, 118, 766 Elodes, 848, 849, 851, 895 Elueophila, 1023, 1045, 1064 Eloeophila obliteralis, 768 Eloeophila persalsa, 1030, 1047 Eloephila trimaculata, 1034

254, 260

Entomobryinae, 252 Environmental DNA(eDNA), 156, 160 Eobrachycentrus gelidae, 615, 616,664, 686, 748 Eocosmoecus, 88, 642, 703, 754 Eocosmoecus frontalis, 643,659, 704 Eocosmoecus schmidi, 643, 659

Eoparargyractis, 769. 770, 771, 780 Eoparargyractisplevie, 88, 767, 768, 773 Eotettix, 421,426

Epantius obscurus, 868, 891 Epeorus, 70, 190, 269, 275, 283, 307, 312, 329

Epeorus frisoni, 312 Epeorus pleuralis, 98 Ephemera, 73, 273,291, 300, 317, 337 Ephemera simulans, 84 Ephemerella, 83, 98,196, 266, 271, 287, 288,298,299, 301,314, 332 Ephemerella alleni, 288 Ephemerella aurivillii, 271, 308 Ephemerella dorothea, 319 Ephemerella invaria, 318 Ephemerella maculata, 308 Ephemerella needhami, 287, 290, 308

Index

Ephemerella nuda, 288 Ephemerella suhvaria, 318 Ephemerella tibialis, 275, 287, 288, 319 Ephemerella verruca, 288 Ephemerellidae, 83, 98, 134, 166, 167, 190, 196, 237, 240, 266,268,272,275, 279, 287, 290, 293,298, 299, 301, 302, 308,311,318,319, 331-33 Epheraeridae, 13,84,98,119, 167,196,198, 240, 273,291, 300, 311,317,337-38 Ephemeroidea, 268 Ephemeroptera, 2, 13,28, 37,49,53, 55, 67, 68, 76, 77, 81, 82, 83-84, 92, 96, 97, 98, 112,113, 115,119, 120, 125, 145, 150, 166, 167-68, 174, 176, 178, 179, 180, 186, 189,190, 195-98, 232, 233, 235, 236, 237,238, 239, 240, 263-339, 791 Ephoron, 53, 197, 264, 265,273, 291, 300, 317,339 Ephoron album, 67, 84 Ephoron leukon, 84 Ephydra, 225, 229, 950, 953, 1008 Ephydra cinerea, 956 Ephydra cineria, 66 Ephydra riparia, 956 Ephydra subopaca, 956, 970 Ephydrella breviseta, 67 Ephydrella marshalli, 67 Ephydridae,49, 73, 91, 108, 166, 167, 186, 225, 226, 229, 909, 925, 926, 927, 928, 929, 930, 950, 953, 955, 956, 958, 959, 960, 970, 974, 979, 981, 1005

Ephydrinae, 1007 Epiaeschna heros, 367, 397 Epicordulia, 380, 400 Epicordulia princeps, 381 Epimetopidae, 168, 798, 804, 838, 841, 844-45, 884

Epimetopus, 798, 804, 838, 842,844, 884 Epiphragma, 228, 925, 1024, 1046, 1050 Epiphragma solatrix, 1056 Epitheca, 84, 201, 341, 356, 400 Epitheca cynosura, 387 Epitheca princeps, 380 Epoicocladius, 1161, 1164, 1166, 1202, 1204,1228, 1257

Erebomyia exalloptera, 994 Ereboporus, 816, 826 Ereboporus naturaconservatus, 877 Eretes, 818, 828, 829 Eretes occidentalis, 877 Eretini, 818, 829 Eretmoptera, 1253 Eriocera, 1049

Eriocerafuliginosa, 1030, 1034 Eriococcidae, 524 Eriococcus, 524

Erioptera, 1024, 1034,1042, 1051, 1062 Erioptera distincta, 1054 Erioptera divisa, 1034 Erioptera dyari, 1054 Erioptera lutea, 1030

Erioptera straminea, 1044 Erioptera vespertina, 1054 Eriopterinae, 1023 Eriopterini, 1023 Eristalinus aeneus, 1022

Eristalis, 50, 187,225, 229, 946, 961, 970, 978, 1022 Eristalis tenax, 948

Erpetogomphus, 349, 372, 376, 394 Erpetogomphus designatus, 375, 378 Erythemis, 3S2, 391,401 Erythemis simplicicollis, 354, 381 Erythrodiplax, 383, 384, 391,402 Euarthropoda, 177 Eubrianax, 802, 807, 850, 851, 852 Eubrianax edwardsii, 893 Eubriinae, 893-94

Eucapnopsis, 446, 466, 472 Eucapnopsis brevicauda, 446, 449,466, 471,472,513 Encorethra, 221, 934 Eucorethra underwoodi, 938, 988

Eudactylocladius, 1169, 1203 Eudicranota (Dicranota subgenera), 1042 Euholognatha,429 Euhrychiopsis lecontei, 904 Eukiefferiella, 1124, 1162, 1172, 1174, 1200,1202, 1204, 1205, 1206, 1208, 1209, 1210, 1257 Eukiefferiella brevinervis, 91 Eukiefferiella devonica, 91,1167 Eukiefferiella potthasti, 1167 Eukiefferiella pseudomontana, 1167 Eulalia, 1001 Eulichadidae, 167, 212, 800, 808, 853, 854, 900 Eulimnichus, 214, 853

Eulophidae, 170, 238, 910, 911, 913, 914, 915,919 Eunemobius, 418,422,428 Euorthocladius, 1168, 1169, 1198, 1199, 1200, 1201, 1203, 1206 Euparyphus, 941, 942, 1000 Euparyphus greylockensis, 967 Euphylidorea, 1024, 1050,1052,1064 Eupteromalus, 920 Euryhapsis, 1170, 1173, 1191,1228, 1257 Eurylophella, 271, 287, 289, 315, 333 Eurylophella coxalis, 271 Eurylophella doris, 83 Eurylophellafuneralis, 83 Eurylophella lutulenta, 308 Eurylophella oviruptis, 264 Eurylophella temporalis, 83 Eutaenionotum guttipenne, 1008 Euthycera arcuata, 1019 Euthyplucia, 196, 273, 299 Euthyplocia hecuba, 112, 272, 297, 338 Euthyplociidae, 167, 196, 272, 273,297, 299, 338

Eutonia, 1024,1050,1065 Eutonia marchandi, 1052 Exneria, 849

Exopterygota, 113

1463

Fabria, 588 Fahria inornata, 648,649,650, 715, 717, 761 Fallceon, 269, 282, 309, 325 Fallceon quilleri, 83, 307 Fallceon thermophilos, 309 Farrodes, 270, 286, 293, 314, 335 Farrodes texanus, 293, 316 Farula, 589, 655,660,663,680, 723, 764 FarulaJewetti, 655 Fattigia, 219, 604,670 Fattigia pete, 88, 654,660,662, 720, 723, 746 Ferrissia, 133

Figitidae, 170,914, 921 Filtering Collector Index, 131, 132 Fine particulate organic matter(FPOM), 174

Fissimentum, 1145, 1158, 1267 Fittkauimyia, 1128, 1136, 1188,1228, 1244

Fittkauimyiini, 1128,1244 Flavohelodes, 895

Fletcherimyia, 225, 1017 Fletcherimyiafletcheri, 949, 953 Floating cages, 40 Folsomia, 252, 254 Folsomides, 253 Folsomina, 252 Fonscolombia, 397 Fontinalis, 118

Forcipomyia, 221, 939,985 Forcipomyia brevipennis, 940 Forcipomyiajohannseni, 966 Forcipomyiinae, 939, 984-85 Fraxinus, 767

Freeze-core samplers, 18 Frenesia, 638, 706, 755 Frenesia missa, 639,658, 705 Friesea, 249,251,259

Frieseafara, 246 Friesea rothi, 246 Frisonia, 457, 495

Frisonia picticeps, 457,460,494,495, 496, 516 Fumonta major, 646, 712, 714, 730 Funnel trap, 22

Galerucella, 73, 793, 902

Galerucella nymphaeae, 89, 869,902 Gallerucella nymphaea, 110 Gallerucella nymphaeae, 111 Gammarus, 134

Gasterophilidae, 53 Gastrops, 1011 Gelastocoridae, 170, 206, 523, 524, 525, 532, 534, 566-67 Gelastocorinae, 532 Gelastocoris, 532, 566 Gelastocoris oculatus, 534 Gelastocorus, 206 Gelechiidae, 767

r 1464

Index

r Georissidae, 167,212, 793, 797, 798, 804, 806, 845, 883 Georissus, 212, 797, 798, 804, 806, 845,883 Georthocladius, 1162,1163, 1166, 1172, 1174, 1175, 1198, 1201, 1257

Geranomyia, 1024, 1027,1045, 1066 Geranomyia canadensis, 1047 Gerking sampler, 21, 39 Gerridae, 66, 71,86, 100, 136, 170, 181, 207,240, 521, 522, 523, 527, 532, 535, 536-37,548,553 Gerrinae, 554 Genis, 86, 100, 523, 535, 548, 554

Gerromorpha, 66, 181, 207, 521, 522, 523 Gesonula punctifrom, 411 Gigantodax, 1100, 1104, 1106, 1109, 1116 Gigantodax adleri, 1105, 1108 Gillotia, 1147, 1224, 1267 Giulianium, 889

Glaenocoris propinqua, 560 Glamocorisa, 528, 529, 530 Glaenocorisini, 530 Glenanthe, 1011

Glossoma nigrior, 94 Glossosoma, 94, 110,126, 190, 220, 593, 601,618,661,667, 676,687, 734 Glossosoma intermedium, 87,688

Glossosoma nigrior, 32, 87, 111, 114,129, 138-39, 172,619

Glossosoma penitum, 87, 92 Glossosomatidae, 32, 87, 94, 102,126, 130, 133, 167, 217, 220, 586, 589, 592, 593, 599, 601, 603, 616-18, 619,661, 666, 667, 668, 676, 681, 687-90,688, 733 Glossosomatinae,687, 734 Glutops, 224, 941, 999 Glutops rossi, 945, 967 Glyphopsyche, 638, 706, 755 Glyphopsyche irrorata, 639, 658, 704 Glyptotendipes, 1154, 1155, 1219, 1221, 1225, 1237, 1267

Glyptotendipes paripes, 91 Goeldichironomus, 1124, 1152, 1155, 1220, 1222, 1229, 1267 Goeldichironomus devineyae, 1155 Goera, 94, 190, 588, 609,618,690, 749 Goera calcarata, 94, 663,689 Goera fuscula, 620, 689 Goeracea, 218, 588, 609, 618,674, 690, 749

Goeracea genota, 620, 663,689 Goeracea oregona, 689 Goereilla, 606, 671,680 Goereilla baumanni, 654,656,659, 720, 721,763 Goeridae, 87, 94, 133, 167, 218, 586, 588, 589,607,609,618, 620, 663, 674, 677, 679,681,684,689,690,749 Goerita, 618,690, 749 Goerita betteni, 663

Goeritaflinti, 663 Goerita semata, 87,620, 689 Gomphaeschna, 367, 368, 397 Gomphaeschnafurcillata, 369, 371

Gomphidae, 13, 75, 80, 84, 135, 166, 169, 190, 199, 201, 341, 342, 343, 344, 347, 349, 350, 352, 353, 354, 356, 372-79, 374, 378,381,388, 394-96 Goraphocerinae, 412,425

Gomphurus, 201, 356, 373, 377, 394 Gomphurus dilatatus, 356 Gonempeda, 1023, 1051, 1062 Gonempeda burra, 1056 Gonielmis, 860, 861, 865, 866 Gonielmis dietrichi, 897

Gonomyia, 1024, 1051, 1062 Gonomyia aciculifer, 1055 Gonomyia cinerea, 1055 Gonomyiaflavibasis, 1055 Gonomyia sulphurella, 1055 Gonomyodes, 1042, 1062 Gonomyodes tacoma, 1047 Gononotus angulicollis, 904 Grab samplers, 20 Graceus, 1225, 1227, 1267 Graded sieves, 18, 19, 21,40 Grammotaulius, 640, 641,644, 706, 755 Grammotaulius lorretae, 707 Granisotoma, 253

Graphoderus, 73, 89, 818,829, 830, 878 Graphoderus liberus, 74 Graptocorixa, 529, 530, 560 Graptocorixa californica, 531 Graptocorixa serrulata, 86 Graptocorixini, 527 Greniera, 1104, 1106, 1109, 1113, 1116 Greniera humeralis, 1110, 1114 Grensiapraeterita, 635, 637, 658, 706, 707, 755 Griseosilvius, 944

Gryllacrididae, 240, 412 Gryllidae, 169,411, 413,415,417-18, 422, 428 Gryllotalpidae, 169, 205, 237, 413, 417, 428

Gryllotalpinae, 428 Gryllotalpus, 205 Gumaga, 587, 660, 720, 746 Gumaga nigricola, 654 Gumaga nigricula, 662 Guttipelopia. 1133,1187, 1189,1248 Guttipsilopa, 1006 Gymnochthebius, 846, 847,892 Gymnoclasiopa, 1011 Gymnometriocnemus, 1161, 1164, 1172, 1200, 1206, 1229, 1257

Gyranomyzinae, 1010 Gymnopais, 926, 1099, 1100, 1101, 1106, 1109, 1116

Gymnopais dichopticodes, 97 Gymnopais holopticus, 1102, 1110, 1111 Gymnopternus, 994 Gymnoscirtetes, 423, 426 Gymnoscirtetes pusillus, 412 Gynacantha, 363, 368, 398 Gynacantha nervosa, 341, 369, 371 07^^,802,810,811,872

Gyrinidae,49, 55,71,72, 89, 104, 114, 135,

r

167, 210, 243, 791, 792, 794, 796, 800, 802, 809-11,871

Gyrinophagus, 920 Gyrinophagus aper, 920 Gyrinus, 792,810,811,872

r

r H Habitat Stability Index, 131, 132 Habrophlebia, 53, 277, 284, 313 Habrophlebia vibrans, 84, 308, 335 Habrophlebiodes, 286, 313, 336 Habrophlehiodes americana, 285, 287, 308 Hadrotes crassus, 889

Haemagogus, 1074,1079, 1081, 1084, 1086, 1087, 1090, 1095 Haematopota, 943, 1003 Hagenella, 649 Hagenella canadensis, 648, 649,664, 715,761 Hagenius, 344, 346 Hagenius brevistylus, 342, 343, 372, 375, 376, 378, 394

Haideoporus, 792, 816, 826 Haideoporus texanus, 878 Halesochila taylori, 638,639, 704, 706, 756 Haliaspis spartinae, 524 Haliplidae, 49, 89, 104, 167, 210, 238, 791, 792, 794, 796, 803, 805, 812-13, 873-75

Haliplus, 104, 210, 805, 812, 813, 815, 874 Haliplus immaculicollis, 89 Halisotoma, 253

Halobates, 66, 181, 522, 523, 535, 537, 555 Halobatinae, 535, 555 Halobrecta algophila, 889 Halocladius, 1160, 1171, 1202, 1205, 1236, 1240, 1241, 1258 Halocoryza arenia, 873 Haloscatella, 1008

Halticoptera, 920 Hamatabanus carolinensis, 967 Hand collection, 18

Hand dipper, 22, 38 Hand screen collector, 18, 19, 38 Hansonoperla, 451,487,515 Hansonoperla appalachia, 491 Haploperla, 455, 482, 520 Haploperla brevis, 86, 458 Haploperla chukcho, 486

Hardy plankton indicator type sampler, 19,20 Harlomillsia, 250

Harmstonia, 994 Harnischia, 1122,1149, 1219, 1268 Harpalus, 805

Hayesomyia, 1134, 1140,1141, 1189, 1190 Hebridae, 170, 207, 527, 535, 539, 564 Hebrus, 207, 535, 539, 564 Hebrus sobrinus, 539 Hecamede, 1011

Hecamedoides glaucellus, 1011 Hedria, 949

r

Index

Hedria mixta, 952, 1019

Hedriodisciis, 229, 941, 1000

Hedriodiscus truquii, 67 Helaeomyia, 950 Helaeomyia petrolei, 1006 Helcomyza, 957 Helcomyza mirahilis, 1015 Helcomyzidae, 168, 926, 957, 1015 Helmiella, 91, 1165, 1208, 1209, 1258 Heleniella thienemanni, 1163, 1166 Heleodromia pullata, 998 Helichus, 211, 793, 805, 854, 855, 857,894 Helicopsyche, 75, 126, 218, 587, 600, 610, 618, 670,672, 678,690, 741 Helicopsyche horealis, 88, 119, 620, 661 Helicopsyche limnella, 88 Helicopsyche mexicana, 88 Helicopsychidae, 88, 119, 126, 130, 133, 168,218, 586, 587,589, 596,600, 610,618-21,620, 661,670, 672,678, 684,690, 740-41 Helius, 1024, 1027, 1033, 1050, 1053, 1067 Heliiis mainensis, 1030, 1053 Helius pallirostris, 1033 Helohata, 839, 844,846

Heptagenia solitaria, 284 Heptageniidae, 55, 70, 83, 97, 98,119, 122, 166, 167,196,198,238, 268, 269, 270, 275, 282,285, 302, 304, 307,310,316, 328-31 Heptageniidae Drunella, 133 Hercostomus, 995 Hermione, 1001 Herthania, 849

Hesperagrion, 361, 362 Hesperagrion heterodoxum, 358, 360, 365, 408

Hesperiidae, 767

Hesperoconopa, 1023, 1025, 1042, 1062 Hesperoconopa dolichophallus, 1044 Hesperocorixa, 74, 528, 530, 531, 560 Hesperocorixa interrupta, 86 Hesperocorixa vulgaris, 531 Hesperoperla, 451, 487,492, 515 Hesperoperla pacifica, 452, 465, 467, 489, 494 Hesperophylax, 642,643, 659, 709, 756 Hesperophylax magnus, 708 Hess sampler, 18, 21, 22, 26,27, 28 Hetaerina, 349, 355,405

Helohata larvalis, 886

Hetaerina titia, 357

Helochara communis, 524

Heterelmis, 858, 859, 865, 866, 897 Heterlimnius, 860, 861, 866,867, 897 Heteroceridae, 212, 791, 793, 795, 800, 801,807, 809, 853

Helochares, 838, 839, 844, 886 Helocomhus, 839, 841

Helocombus bifldus. 846,886 Helocordulia, 380, 400 Helodidae, 49, 89 Helodun, 1100, 1104, 1106, 1109, 1117 Helodon alpestris, 1112 Helodon decemarticulatus, 1111 Helodon onychodactylus, 1105 Helopelopia, 1133, 1134,1138, 1188,1228, 1248

Helophilus, 1022 Helophoridae, 168, 212, 798, 804, 838, 842, 845, 884 Helophorus, 212, 798, 804, 838, 842, 845, 884

Hehpicus. 85,459,495, 516 Helopicus bogaloosa, 498 Hehpicus nalatus, 461 Helopicus subvarians, 498 Helotrephidae, 52, 521, 523 Hemerodromia, 944, 947, 968,998 Hemerodromia einpiformis, 91 Hemerodromiinae,944 Hemiacridinae, 411 Hemimetabolous, 233 Hemiosus, 837, 841 Hemiosus exilis, 886

Hemipachnobia, 786 Heniiptera, 3-6, 13, 37, 49, 56, 66, 71, 72, 73, 76, 82, 86, 97, 100-102, 113, 115, 119, 120, 166, 170, 174, 176, 181, 187, 188, 232, 233,234, 236,237, 238, 239, 240, 241, 242, 521-67, 523, 792, 794, 910 Heptagenia, 284, 285, 304, 312, 329 Heptagenia culacantha, 284

Heterocerus, 212 Heterocheila, 957 Heterocheila hannai, 1015

Heterocheilidae, 168, 926, 957, 1015 Heterocloeon, 195, 265, 280, 306, 325 Heterocloeon amplum, 294 Heterocloeon curiosum, 83, 294, 318 Heterocloeon grande, 280 Heteromurus, 250

Heteroplectron, 76, 104, 218, 589, 590, 616, 671,680, 687, 740 Heteroplectron americanum, 617, 661,687 Heteroplectron californicum, 88 Heteroptera, 65, 75, 80, 150, 206-7, 521, 522, 523, 524 Heterosternuta, 817, 825, 827, 878

Heterotanytarsus, 1159, 1194, 1258 Heterotanytarsus perennis, 1163, 1166 Heterotrissocladius, 73, 1161, 1163, 1164, 1194, 1197,1258 Heterotrissocladius maeaeri, 1167

Heterotrissocladius marcidus, 1167 Hexacola, 914, 921 Hexacola hexatoma, 910 Hexacylloepus, 858, 859, 866 Hexacylloepusferrugineus, 897 Hexagenia, 73, 98, 190, 196, 198, 273, 291, 311,317, 337 Hexagenia limbata, 84, 231, 264, 311

Hexagenia munda, 84 Hexagenia rigida, 58 Hexapoda, 177 Hexatoma. 1023, 1024, 1025, 1026, 1027, 1030, 1034, 1045, 1046, 1065

1465

Hexatoma californica, 1049 Hexatominae, 1023 Hexatomini, 1023 Heyterota hlumbea, 889 Himalopsyche, 217 Himalopsyche phryganea, 653, 656, 718, 738

Histeridae, 167, 793, 795, 804, 805, 845, 883

Holocentropus, 651,652, 656, 718, 731 Holocentropus interruptus, 719 Holometabola, 180, 181, 232, 233 Holometabolous, 233, 1071, 1120 Holorusia, 1024

Holorusia hespera, 1038, 1043, 1069 Holotanypus, 1187, 1188, 1189 Homoeoneuria, 197, 282, 310, 327 Homoleptohyphes, 270, 271

Homoleptohyphes dimorphus, 83 Homophoheria, 786 Homophylax, 606,634, 636,698, 756 Homophylax andax, 657 Homophylaxflavipennis, 701 Homoplectra, 621, 622,691, 727 Homoplectra doringa, 693 Homoptera, 523, 524, 525, 526 Hoperius, 820,833 Hoperius planatus, 878 Hoplitimyia, 1000 Hoplodictya, 1020 Hoplolabis, 1023, 1024, 1051, 1054, 1063 Horismenus mexicanus, 915,919

Hudsonimyia, 1133, 1188, 1248 Hudsonimyia karelena, 1135 Huleechius, 858, 865, 866 Huleechius marroni, 897

Husseyella, 527, 544, 547 Husseyella tumalis, 552 Hyadina, 1014 Hyallela, 133 Hyallela azteca, 151 Hybomitra, 224, 944, 1003 Hybomitra epistates, 967 Hydaticini, 818, 829 Hydaticus, 89, 815, 818, 828, 829, 878 Hydatophylax, 634,636,657, 709, 756 Hydatophylax argus, 634, 707

Hydatophylax Hesperus, 634,637 Hydatostega, 995 Hydracarina, 190, 344 Hydraena, 212, 805, 846, 847, 892 Hydraenidae,49, 167, 183, 212, 234, 792, 793, 794, 797, 799, 804, 805, 845-47, 891-92

Hydrellia, 73, 909, 910, 950, 1013 Hydrellia williamsi, 955 Hydrelliinae, 1013-14 Hydrilla, 766 Hydrobaeninae, 1253 Hydrobaenus, 1162, 1165, 1167, 1192, 1193, 1258

Hydrohaticus, 844 Hydrobiomorpha, 839, 840, 841, 842 Hydrobiomorpha casta, 886

1466

Index

Hydrobiosidae, 168,217, 586, 589, 592, 601,603, 620,622,666,667, 668, 676, 681,690, 735

Hydrohius, 836, 839, 840, 844 Hydrohiusfuscipes, 839, 844, 886 Hydrobiusini, 844 Hydrocanthus, 210, 833, 834, 835, 882 Hydrochara, 213,806, 837, 840, 841,

Hydroptila arctia, 81 Hydroptilidae, 81, 87, 127, 136, 166, 167, 217, 241, 586, 589, 592, 593, 599, 600, 601, 625-30,665,666, 676,677,681, 691-97, 735-38

Hydroptilidae, 220 Hydropyrus, 1007 Hydroscapha, 209, 792, 799, 801, 802, 803, 835, 836, 882

842, 886

Hydrochara rickseckeri, 836 Hydrochusma, 1011 Hydrochidae, 168, 213, 798, 804, 841, 845,884

Hydrochus, 213, 798, 804, 842,845,884 Hydrochus spangleri, 845 Hydrocolus, 816, 827, 878 Hydrocyrius, 50 Hydrodytes, 816, 829 Hydrodytes dodgei, 878 Hydrodytinae, 829 Hydrolsotoma, 253 Hydroisotoma xchaefferi, 247, 255 Hydroistoma schaefferi, 259 Hydrometra, 207, 527, 541, 551 Hydrometra amtralis, 541 Hydrometra martini, 86 Hydrometridae, 86, 170, 207, 527, 541, 551 Hydromyza confluens, 1018 Hydroperla, 459, 495, 517 Hydroperla croxbyi, 85, 100, 461,498 Hydroperla rickeri, 498 Hydrophiilini, 844 Hydrophilidae, 49,89, 106, 114, 134, 135, 167,213, 238, 243, 791, 792,793, 794, 795,796, 798, 806, 836-44, 846, 884-87

Hydrophilinae, 839, 841, 844 Hydrophilini, 839 Hydrophilius, 843 Hydrophiloidea, 795 Hydrophilus, 836, 837, 839, 840, 841, 843, 886 Hydrophilus triangularis, 106 Hydrophorus, 995 Hydrophorus oceanus. 968 Hydrophylita aguivolans, 915, 921 Hydroporinae,814, 816, 820 Hydroporini, 817, 826 Hydroporninae,829 Hydroporus, 805, 814, 815,817,825, 826, 828,878 Hydropsyche, 87, 102, 103, 220, 586, 591, 602,622,670, 678,691,728 Hydropsyche betteni, 621, 623,624 Hydropsyche orris, 76 Hydropsyche slossonae, 87, 110, 111, 114, 692,693

Hydropsychidae, 77, 78, 81, 87, 102, 119, 135, 167, 190, 216, 220, 586, 591, 599, 600,602,621, 622-25,666,670,678, 682,690-91,692,693, 726

Hydropsychinae, 586,690 Hydropsychoidea, 726-27 Hydroptila, 220,626,627,628,694, 736

Hydroscapha natans, 835,836 Hydroscapha redfordi, 835 Hydroscaphidae,49, 51, 167, 209, 791, 792, 799, 801,802,803, 835-36, 882 Hydrosmilodon, 270, 284,285, 308, 313 Hydrosmilodon primanus, 336 Hydrosmittia, 1176, 1211, 1258 Hydrothassa, 902 Hydrotrupes, 814, 818, 829, 830 Hydrotrupes papalis, 878 Hydrotrupini, 818, 829 Hydrous, 51 Hydrovatini, 823 Hydrovatus, 816, 822, 823, 878 Hygrobiidae, 794 Hygronemobius, 418,428 Hygrophila, 767 Hygrotus, 817,826, 828, 879 Hyiogomphus, 373, 377, 395 Hymenoptera, 2, 13, 37, 106, 113, 120, 166, 170, 174, 176, 185, 232, 234, 236, 238, 239, 242, 243, 344, 909-24 Hyphydrini,824 Hypocharassus, 995 Hypogastruidae, 248 Hypogastrura, 249,251 Hypogastrura littoralis, 246 Hypogastruridae, 169, 247, 251, 258-59 Hypogastrurinae, 247 Hyponeura, 407 Hyporheic canister sampler,41 Hyporheic standpipe corer, 41 Hyporhygma, 1149, 1153, 1220, 1229, 1268 Hyposmocoma, 215, 767, 789

loscytus politus, 545 Iris versicolor, 342 Irmakia, 1224

Iron, 53,71,269, 329 Ironodes, 283, 307, 312, 329 Ironopsis, 269, 329

Ironoquia, 642,643, 644,665, 703, 756 Ironoquia punctatissima, 702 Ischnura, 359,361,362,408 Ischnura posita, 364 Isocapnia, 77, 429,446,466,470, 513 Isocapnia grandis, 469,471 Isocapnia Integra, 447 Isocladius, 1164, 1169 Isocladius elegans, 1168 Isocytus, 565 Isoetes, 767

Isogenoides, 430,457, 495, 517 Isogenoides olivaceus, 85 Isogenoides varians, 498 Isogenoides zionensis, 460 Isonychia, 70, 77, 83, 122, 196, 198, 265, 268, 282, 301, 327

Isonychia bicolor, 58, 83,93 Isonychiidae, 77, 83, 93, 167, 196, 198, 268, 282, 297, 301,327

Isoperla, 80,85,204,432, 455,457,459, 497,499, 502, 505,519

Isoperla bilineata, 438,460 Isoperlafulva, 85, 501, 503 Isoperla signata, 85 Isoperlinae, 518-19 Isoptera, 180 Isotoma, 252, 253,255, 260

Isotoma dispar, 246 Isotomidae, 169,194,241, 250, 251, 254, 255, 257, 259-60 Isotomiella, 253 Isotomodes, 253 Isotomurus, 253, 255 Isotomurus tricolor, 255

Issidae, 524 Iswaeon, 280, 306, 325 Iswaeon anoka, 294

I

Ithytrichia, 593,626,628,629,630,695,

Ichneumonidae, 170, 238, 914, 917, 923-24 Ichneumonoidea, 909, 913, 922-23

697, 736 Ithytrichia clavata, 665

Idiataphe cubensis, 383, 390,402 Idiocera, 1024, 1051, 1063

Idiocera gothicana, 1055 Idiocoris, 52 Idioptera, 1024, 1050, 1052, 1065

Idioptera pulchella, 1029 Ilisia, 1051, 1063 Ilisia venusta, 1054

Ilybiosoma, 820,830 Ilybius, 89, 815,820, 830, 832, 879 llyocoris cimicoides, 61 Ilytheinae, 1014-15

r\

Isotuma marisca, 246

Janetschekbrya, 254 Jenkinshelea, 986 Johannsenomyia, 977,986

Juga, 133 Juncus, 413

K Kaluginia, 1251

Inscudderia, 417 Insudderia, 428

Kambaitipsychidae, 586 Kathroperla, 77, 453,482, 519 Kathroperla perdita, 454,456

Integripalpia, 585, 586-92, 594, 733-39

Keirosoma slossonae, 995

loscytus, 544, 546

Kellen grab, 21, 22, 38

ry

Index

Kick sampling, 18, 26 Kiefferulus, 1154, 1220, 1222, 1223, 1226, 1268

Kirmaushenkreena, 268, 302 Kirmaushenkreena zarankoae, 268, 282, 293, 325 Kkidotoma, 914 Kleidotoma parydrae, 910, 921 Kloosia, 1158, 1159, 1219, 1268 Knowltonella, 249

Kogotus. 455, 499, 505, 517 Kogotus modestus, 501, 504 Kogotus nonus, 458 Krenopelopia, 1133, 1135, 1138, 1139, 1186, 1188, 1248 Krenosmittia, 1160, 1166, 1172, 1200, 1204, 1258 Kribiodorum, 1154, 1156, 1225, 1268 Krizousacorixa, 560

Lathromeroidea gerriphaga, 912, 921 Latineosus, 291 Latineosus cibola, 335 Latineous, 269

Lauterborniella, 1148, 1154, 1157, 1219, 1229, 1239,1268

Leaf pack sampler, 39 Leaf packs, 18 Lednia, 446, 474,476,510 Lednia tumana, 447,477,478 Leiodidae, 793

Leiponeura, 1055 Lemanea, 592

Lemmatophora typa, 181 Lemmatophoridae, 180 Lemna, 118

Lemnaphila, 950 Lemnaphila scotiandae, 1013 Lenarchus, 642,643, 711, 757 Lenarchus brevipennis, 710 Lenarchus vastus, 659 Lenziella, 1142, 1143

Labiobaetis, 268, 278, 292, 307, 309, 326 Labiohaetis longipalpus, 278 Labostigmina, 1001 Labrundinia, 1132, 1133, 1135, 1187, 1 189, 1190, 1233,1248 Laccobiiini, 844 Laccobius, 837, 838, 843, 844, 886 Laccomimus, 818, 820 Laccomimus pumilio, 879

Laccophilinae, 818, 820 Laccophilini, 818 Laccophilus, 802, 815, 818, 819, 820, 822, 879

Laccophilus maculosus, 89 Laccornini, 826 Laccornis, 814, 817, 825, 826,879 Lachlania, 273, 282, 299, 310, 327 Ladona, 201,382, 389,402 Ladona deplanata, 84 Lambourn sampler, 19, 21 Lampracanthia, 540 Lampracanthia crassicornis, 546, 565 Lamprochromus, 995 Lamproclasiopa, 1011 Lamproscatella, 1008 Lampyridae, 213, 793, 795, 797, 798, 804, 868

Langessa, 770, 771 Langessa nomophilalis, 766, 780 Lanthus, 373, 377, 395 Lanthus vernalis, 84

Lappodiamesa, 1177,1179, 1206,1251 Lappokiejferiella, 1258 Lapposmittia, 1165,1168, 1202, 1204, 1258

Lara. 76,211, 856, 857, 860, 862,897 Lara avara, 82, 89, 112 Larsia, 1133, 1135, 1184, 1185, 1248 Lasiodiamesa, 1177,1181, 1182, 1183, 1250

Lasiomastix, 1050, 1053 Lathromeroidea, 920

Lepania, 679 Lepania cascada, 618,620,663,690, 749 Lepidocyrtus, 194, 250, 254 Lepidoptera, 13, 37,49, 53, 55, 73, 88, 106, 113, 120, 127, 166, 170, 176, 183, 189, 190, 192,215,232, 233,234, 235, 236, 238,239, 241, 242, 243, 765-89,909

Lepidostoma, 88, 219, 588, 609,630,631, 671,682, 697, 750 Lepidostoma bryanti, 88 Lepidostomajlinti, 662 Lepidostoma quercinum, 698 Lepidostomatidae, 88, 134, 167, 219, 586, 588, 589, 596,607,609,630, 631, 662, 664,671, 675,682,686,697, 698, 749

Lepidostomatinae, 750 Leptanthicus, 868 Lepthemis, 401 Leptobasis, 359, 362,408 Leptoba.sis melinogaster, 387 Leptoceridae, 72, 88, 104, 134, 167, 190, 218, 220, 586, 587, 589, 595,600, 608, 630-33, 664, 665,672, 673,680, 682, 684,697-98, 699, 700, 741-43 Leptocerus, 587 Leptocerus americanus, 630, 631,632,664, 698, 700, 742

Leptoconopinae, 939,987 Leptoconops, 939,987 Leptohyphes, 75, 111, 289, 290, 304, 305, 315,334

Leptohyphidae, 75, 81, 83, 134, 167, 196, 264, 267, 270, 272,273, 287, 290, 302, 304, 305, 333-34 Leptohyphinae, 270

Leptonannus latipennis, 553 Leptonema, 624 Leptonema albovirens, 625, 690,692, 729 Leptophlebia, 130, 192, 277, 286, 307, 313, 336

1467

Leptophlebia bradleyi, 277 Leptophlebia cupida, 98 Leptophlebiidae, 84, 98, 119, 134, 166, 167, 196, 265,268,270, 273, 274, 275,277, 285,287,293, 301,302, 308,316, 335-37 Leptophylax, 633 Leptophylax gracilis, 633, 704, 709, 757 Leptopodidae, 521, 522

Leptopodomorpha, 207, 521, 522 Leptopsilopa, 1006 Leptotarsus, 1023, 1028,1069 Leptotarsus testaceus, 1036 Leptyisminae, 412 Leptyminae,411 Leptysma, 421, 425 Leptysma marginicollis, 412,414 Leptysminae, 411,425 Lestes, 76, 84, 200, 201, 342, 345, 349, 351,355,406 Lestes congener, 98 Lestes disjunctus, 52, 53 Lestes inaequalis, 357 Lestes sponsa, 188 Lestes vigilax, 352, 360 Lestidae, 55, 76, 84, 98, 135, 169, 200, 201, 342, 348, 349, 350, 351, 352, 355, 357, 360,405-6 Lestremiinae, 1028 Lethemurus, 250 Lethocerinae, 527 Lethocerus, 231, 522, 527, 528, 556 Lethocerus americanus, 86,526 Lethocerus maximus, 86

Leucorrhinia, 383, 384, 390,402 Leucotabanus, 944, 1003 Leucotabanus annulatus, 945

Leucotrichia, 593,625,626,629,696, 697, 736 Leucotrichiini, 592 Leucrocuta, 83, 269,270, 284, 312, 329 Leuctra, 85, 202, 429,448,450, 476,479, 512

Leuctra biloba, 480

Leuctraferruginea, 79, 114 Leuctra grandis, 463,465,483 Leuctra moha, 480

Leuctra sibleyi, 438 Leuctridae, 79, 85, 114, 134,169, 202, 204, 237, 429,430, 431,438, 439, 44!, 450,452,462,463,465,478,480, 481,483,511 Leuctrinae, 511-12 Liancalus, 995 Libellula, 84, 382, 389, 402 Libellula incesta, 381

Libellula quadrimaculata, 341, 385 Libellula semifasciata, 389 Libellulidae, 84,119, 135, 169, 188,190, 199, 201, 240, 342, 343, 345, 350, 352, 353, 354, 381, 382-91,401^ Limnebius, 212, 846, 847, 892 Limnellia, 1008

1468

Index

Limnephilidae, 68, 73, 88, 102, 133, 134, 137, 166, 167,219,220,240,241,586, 589, 594, 595, 596, 597, 599, 606, 607, 633-42, 643, 644, 657, 658,659,663, 665,677, 681, 682,698-711, 750-60

Lissorhopterus simplex, 89 Lissorhoptrus, 73, 189, 905 Lissorhoptrus oryzophilus, 73 Lissorhoptrus simplex, 869 Listronotus, 905

Malenka coloradensis, 475

Lobelia, 767

Malirekus, 459, 497,499, 517 Malirekus hastatus, 458, 501, 503 Mallochohelea, 986 Mallota, 1022 Mandibulata, 177 Manoa, 1127, 1271

Lixellus, 905

Limnia, 1020

Lophognathella, 247 Lophotrochozoa, 177 Loxocera cylindrica, 960

Limnocorinae, 538 Limnocoris, 538, 541, 558

Limnocoris moapensis, 542 Limnogonus, 537 Limnogonusfranciscanus, 554 Limnohydrohius, 839, 844, 887 Limnophila, 933, 1023, 1026, 1034, 1042,

Longurio, 1028, 1036 Lopescladiu,s, 1159, 1163, 1182, 1211, 1212, 1259

Lurdia, 1044 Lutrochidae, 168, 182, 214, 799, 806, 808, 853-55,900-901 Lutrochus, 214, 806, 808, 853, 854, 901

Lymantria dispar, 1025 Lymnaecia phragmitella, 789 Lype diversa, 87,653,656, 718, 720, 732 Lysathia ludoviciana, 868 Lythrum, 793,868 Lytogaster, 954, 1015 Lytogaster excavata, 958

1064-66

Limnophora, 926, 960, 961, 980, 1016 Limnophora riparia, 91, 970 Limnophyes. 1170, 1207, 1209, 1259 Limnoporus, 207, 523, 535, 536, 554 Limnoporus canaliculatus, 86 Limnoporus notahilis, 86 Limoniidae, 167, 223,228, 1023, 1024, 1025, 1027, 1028, 1029, 1030, 1033, 1034, 1044, 1047, 1048, 1049, 1052, 1053, 1054,1055, 1056, 1057, 1058, 1059, 1061 Limoniinae, 1026, 1027 Limoniini, 1023 Limoninae, 1023 Liodessus, 816, 817, 824, 825, 879

Liogma nodicornis, 933 Lioporeus, 816, 827,828, 879 Liparocephalus, 847, 848, 889 Lipiniella, 1154, 1156, 1157, 1220, 1229, 1268 Lipochaeta slossonae, 1012 Lipogomphus, 535, 539 Lipogomphus hrevis, 564 Lipsothrix, 76, 82, 90, 112, 925, 1024, 1046, 1067 Lipsothrix hynesiana, 1048 Lipsothrix nigrilinea, 106 Lipsothrix Sylvia, 90, 1058 Lipurometriocnemus, 1259 Lispe, 225, 229, 1016 Lispocephala, 1016 Lispoides aequifrons, 1016 Lissorhopterus oryzophilus, 89

Manophylax, 613,677, 686, 747 Manophylax annulatus, 612, 683 Manophylax butleri, 670 Mansonia, 50, 73, 1071,1072, 1075, 1077, 1079, 1082, 1087, 1090, 1095 Mansonia dyari, 1075 Mansonia titillans, 1075

Mantodea, 180 MareIlia remipes, 411 Marilia, 645,646, 711, 744 Marilia flexuosa, 713 Marilia nohsca, 662

Martarega, 538

1044, 1050, 1065

Limnophila macrocera, 1053 Limnophilinae, 1023, 1025, 1026, 1027,

1037, 1069

Llythea, 1014

640,641,682,709,711,757 Limnephilus externus, 658 Limnephilusfumosus, 710 Limnephilus indivisus, 88,102 Limnephilus lunatus, 73, 102 Limnephilus rhomhicus, 708 Limnephilus samoedus, 701, 703

Limnius volckmari, 61 Limnochironomus, 1266

Maekistocera longipennis, 1032, 1035,

Lixus, 905

Litohrancha recurvata, 291, 311, 317, 337

Limnichites, 853 Limnichoderus, 853

Maekistocera, 1024

Malaise trap, 40 Malenka, 442, 472, 510 Malenka californica. 440

Limnephilus, 68, 590,606, 636,638,639,

Limnichidae, 168, 214, 793, 799, 808, 853,900

Macrovelia hornii, 539, 551 Macroveliidae, 170, 523, 527, 535, 539, 551

Martarega mexicana, 543, 562

M Macan sampler, 21, 39 Maccaffertium, 269, 283 Maccaffertium modestum, 83 Maccaffertium vicarium, 83 Macdunnoa, 283, 285, 307, 313, 330

Maruina, 222, 228, 926, 936, 938,990 Maruina californiensis, 964 Maruini,990 Matinae, 818, 830 Matriella, 271

Mackenziella, 256

Matriella teresa, 287, 288, 314, 319, 333 Matus, 818,819, 830, 879

Mackenziella psocoides, 257

Matus bicarinatus, 89

Mackenziellidae, 256

Mayatrichia, 628,630,695,697, 736 Mayatrichia ayama, 628,629 Medophron, 910,923

Macrancylus linearis, 905 Macratria, 868

Macrelmis, 858, 859, 863, 864, 898 Macrodiplax halteata, 383, 390,402 Macromia, 200, 201, 343, 354, 378, 379, 399 Macromiidae, 169, 200, 201, 344, 350, 353, 354, 356, 378, 379, 399 Macromiinae, 75 Macronema, 52 Macronematinae, 586,690, 729

Macronychus, 860,861, 862, 863 Macronychus glabratus, 89, 898 Macropelopia, 1131, 1132, 1134, 1136, 1137, 1139, 1186, 1187,1188,1189, 1228, 1245

Macropelopia decedens, 1132 Macropelopiini, 1128, 1235, 1244 Macrophotography, 193-230 Macroplea, 189 Macrorhyncolus littoralis, 905 Macrostemum, 78, 591,622, 625, 690, 729 Macrostemum Carolina, 87, 621,624 Macrostemum zebratum, 87,692 Macrothemis, 383, 389, 403 Macrothemis celeno, 381 Macrovelia, 522, 535, 539

Megadytes, 818, 827 Megadytes fraternus, 880 Megaleuctra, 448, 476, 511 Megaleuctra complicata, 441, 463 Megaleuctra flinti, 480 Megaleuctra kincaidi, 450,478 Megaleuctrinae, 511 Megaloptera, 13, 37,49, 53, 54,65, 73, 86, 95, 97, 102, 112, 113, 120, 166, 170, 174, 176, 181-82, 189, 190, 232,234, 236, 237, 239, 240, 241, 242, 569-84 Megamelus davisi, 524 Megarcys, 455,492, 517 Megarcys signata, 458,494 Megasetia, 948, 1017 Melanderia, 995

Melanoplinae, 412,425-26 Melanoplus, 423, 426 Melittobia, 913

Melyridae, 168, 793, 794, 795, 800, 801, 802, 803, 847, 890-91 Meridiorhantus, 820, 833 Meridiorhantus calidus, 880

Meringodixa, 936

Index

Meringodixa chalonensis, 937, 989

Micrasema rusticum, 614

Morulina, 250, 251

Mermiria, 423,425

Micrasema wataga, 614, 615, 685 Micrathyria, 384, 390, 403 Micrisotoma, 253

Morulodes, 250

Meropelopia, 1133, 1139, 1141, 1188, 1228, 1248

Meropleon diversicolor, 786 Merragata, 535, 539, 564 Merycomyia, 73, 943, 1003 Merycomyia whitneyi, 945 Mesachorutes, 249 Mesites, 905 Mesocapnia, 448, 466,470,472, 513 Mesocapnia arizonensis, 475

Mesocapnia bergi, 470 Mesocapnia frisoni, 449, 463,471 Mesocricotopus, 1194, 1196, 1197, 1259 Mesocyphona (Erioptera subgenera), 1024, 1051

Mesoleptus, 923 Mesonoterus, 835

Mesonoterus addendus, 882 Mesopsectrocladius, 1196, 1228 Mesosmittia, 1259 Mesothemis, 401

Mesovelia, 207, 522, 523, 527, 539, 564 Mesoveliidae, 170, 207, 523, 527, 539, 564 Mestocharis, 913

Mestocharis bimacularis, 919

Metabarcode sequencing, 156, 159 Metachela, 946, 947, 998 Metacnephia, 1100,1104, 1106, 1113, 1117

Metacnephia borealis, 1108 Metacnephia saskatchewana, 1111 Metacnephia sommermanae, 1102, 1112 Metacnephia villosa, 1107 Metaleptea, 423, 425 Metaleptea brevicornis, 412 Metamorphosis, 113-15, 1071 Methlini, 829 Metisotoma, 253, 254

Metretopodidae, 167, 197, 274, 275, 285, 302, 304, 322-23

Metretopus, 284, 304, 313, 322 Metretopus alter, 285 Metrichia, 627, 694, 696,697, 737 Metrichia nigritta, 626 Metrics/multimetric bioassessment indi

ces, 144, 145-47 Metriocnemini, 1254 Metriocnemus, 1169, 1198, 1201, 1206, 1259 Metriocnemusfuscipes, 1168 Metriocnemus knabi, 1168

Metrioptera, 428 Metrobates, 207, 535, 536, 555 Metrobates trux, 536 Miathyria marcella, 352, 383, 386, 390, 403 Micracanthia, 544, 565 Micracanthia quadrimaculata, 545 Micralymma, 847, 889 Micranurida, 250

Micranurophorus, 252 Micrasema, 588, 609,616, 687, 748 Micrasema gelidum, 87

Microbledius, 889 Microcara, 848

Microcara explanata, 895 Microchara, 849

Microchironomus, 1147, 1150, 1218, 1219, 1268

Microchrysa, 961 Microcylloepus, 858, 859, 865, 866,898 Microgastrinae, 909 Microgastrura, 249, 251 Microisotoma, 257 Micromorphus, 995

Micronaspisfloridana, 868 Micronecta, 538, 549 Micronecta ludibunda, 549, 561 Micronectidae, 170, 525, 529, 538, 561 Micropsectra, 91, 1143, 1144, 1145, 1146, 1213, 1214, 1216, 1217, 1229, 1272

Micropterus salmoides, 72 Microsporidae, 836 Microsporus, 836 Microtendipes, 1148, 1154, 1158, 1222, 1268

Microvelia, 207, 522, 544, 547, 552 Microvelia beameri, 547

Microveliinae, 544 Mimapsilopa cressoni, 1006 Minto sampler, 21 Mitchellania, 249 Mochlonyx, 934,938,988 Modified air-lift sampler, 41

Modified Gerking sampler, 21, 39 Modified Hess sampler, 18, 38 Modified KUG sampler, 21 Molanna. 190, 219, 220, 587, 608, 642, 673,680,711,743 Molanna angustata, 53 Molanna blenda, 661 Molannaflavicornis, 88, 712 Molanna tryphena, 645 Molannidae, 88, 167, 219, 220, 586, 587, 589, 599,608,611,642,645,661, 673,675,680,684, 711, 712, 743-44

1469

Moselia, 448, 476,479, 512 Moselia infuscata, 450,480, 483 Moselyana, 220,605,679 Moselyana comosa, 611,612,657,683, 686, 747 Mosillus, 1012 Motschulskium sinuatocolle, 845, 883

Multiple core sampler, 24, 38 Multiple-plate artificial substrate sampler, 39

Multiple-plate sampler, 20 Mundie pyramid trap, 19 Munroessa, 765, 780 Muscidae, 91, 167, 225, 229, 243, 925, 928,957, 960, 961,970, 974, 980, 1016

Musotiminae, 765, 769, 783 Mycetophiloidea, 1028 Mymaridae, 170, 910, 911, 913, 914, 915, 919-20

Myolepta, 1022 Myriapoda, 177 Myriophyllum, 766 Myriophyllum aquaticum, 868 Mystacides, 631,632, 633,698, 742 Mystacides interjectus, 88,699, 700 Myxophaga, 209, 791, 792, 795, 882-83 Myxosargus, 941, 1001 Myxosargus nigricornis, 942

N Namamyia plutonis, 645,646, 647, 662, 711,713,744 Nannochoristidae, 235 Nannocoris arenarius, 553 Nannothemis bella, 384, 389,403 Nanocladius, 1165, 1168, 1172, 1191, 1193, 1194, 1195, 1196, 1228, 1259

Nanomyina barbata, 995 Nanonemoura, 442, 472 Nanonemoura wahkeena, 440, 442,447, 472,510 Narpus, 856, 859, 863, 864, 898 Nartus, 820, 833

Molannodes, 608

Nasiaeschna pentacantha, 367, 370, 398

Molannodes tinctus, 642,645, 661, 711,

Nasiternella, 1024, 1042

712, 744 Mollusca, 177 Molophilus, 1023, 1024, 1034, 1050, 1053,

Nasiternella hyperborea, 1067 Natarsia, 1124, 1128, 1131, 1186, 1188,

1063

Molophilus nitidus, 1058 Monodiamesa, 1178, 1180, 1196, 1199, 1253

Monohele, 986

Monopelopia, 1133, 1134, 1137, 1138, 1139, 1188, 1228, 1248 Monophylax mono, 634, 705, 706, 757 Monopsectrocladius, 1194, 1195 Montezumina, 417, 428 Moribaetis, 282, 293, 309, 311, 326 Morphocorixa, 530, 531, 560

1247

Natarsiini, 1128, 1235, 1247 Naucoridae,49, 51, 55, 86, 170, 206, 522, 523,525,538,541,542,558 Naucorinae, 538 Neanura, 250, 251 Neanurinae, 247 Neargyractis, 769, 770, 771 Neargyractis slossonalis, 767, 768, 777, 781

Neaviperla, 453, 482 Neaviperlaforcipata, 486, 520 Nebrioporus, 816, 817, 827, 880

1470

Index

Nectopsyche, 218, 587,631,632, 633, 698, 742

Nectopsyche alhida, 88,699 Nectopsyche tavara, 664 Neelidae, 256 Neelus, 256 Nehalennia, 358, 359, 361,408 Nehalennia irene, 365

Nehalennia minuta, 364 Nematocera,927, 928-29, 931, 969, 1071

Nematoproctus, 995 Nemobiinae,428

Nemocapnia, 446, 466, 470 Nemocapnia Carolina, 447, 466, 470, 473,513 Nemotaulius hostilis, 638,639,658, 705, 706,757

Neoleptophlebia, 270, 286, 313, 336 Neoleptophlebia adoptive, 319 Neoleptophlebia mollis, 319 Neolipophleps, 1055 Neomusotima, 783

Neonemobius, 418,422,428 Neoneura, 359, 363,409 Neoneura aaroni, 366 Neopachylophus, 804, 805, 845

Neopachylophus sidcifrons, 845,883 Neoperla, 432, 451,485,489,490, 514 Neoperla clymene, 85,452,491 Neoperla coosa, 488 Neoperla stewarti, 491 Neophemeridae, 299 Neophylax, 68,88, 94, 126, 589, 590,605, 655,660, 723,763

Neurocordulia alabamensis, 354 Neurocordulia molesta, 84, 354 Neurocordulia obsoleta, 381

Neurocordulia yamaskanensis, 381 Neuroptera, 37, 49, 53,65,86, 97,113, 120, 166, 170, 174, 176,181-82, 189, 232, 234, 236, 237, 239,240, 241, 242, 569-84

Neuropteroids, 208 Neurorthidae, 181, 234

New Jersey light trap, 40 Nigronia, 126,174, 208, 570, 571, 574, 578, 579, 583

Nigroniafasciatus, 576 Nigronia .serricornis, 86, 575, 576 Nilobezzia, 987

Nilotanypus, 1129, 1132, 1133, 1135, 1184,

Neophylax consimilis, 88 Neophylaxfuscus, 114 Neophylax mitchelli, 88 Neophylax oligius, 94, 721

1185, 1248 Nilothauma, 1149, 1150, 1225, 1227, 1269 Ninelia, 1259 Niphograpta albiguttalis, 767, 785

Nippotipula (Tipula subgenera), 1023, 1025,

Neobagoidus carlsoni, 905

Neophylax ornatus, 664 Neoplasta, 224, 946, 947, 998 Neoplea, 48, 206, 540, 541, 557 Neoplea striola, 541 Neoporus, 817, 825, 826, 827 Neoptera, 178, 180,232 Neoscapteriscus, 417 Neoscutopterus, 820, 830, 832, 880

Neobidessus, 816, 824, 825, 880

Neosminthurus, 256, 257

Neocataclysta, 768, 769, 771 Neocataclysta magnificalis, 766, 781

Neostempellina, 1145, 1146, 1272

Noctuinae, 785-86

Neotettix, 418,426

Noctuoidea, 769, 770 Nostima, 954, 1015 Nostima approximata, 958,959 Nostococladius, 1198, 1201 Notaris, 906 Noteridae, 89, 167,189, 210, 792, 794, 796, 803, 805, 833-35, 881-82 Noterus, 189, 834 Noterus crassicornis, 833 Nothifixis, 403 Nothotrichia shasta, 627,696,697, 737 Notiodes, 906

Nemotelus, 939, 1001

Nemotelus kansensis, 942 Nemoura, 442,474,477,478, 510 Nemoura arctica, 67,445

Nemoura trispinosa, 85, 100 Nemouridae, 53, 85, 100, 134, 137, 169, 188, 202, 204, 429,430, 431,433,437, 440,445, 447, 462,463,465,475,477, 478,509 Nemourinae, 510-11 Neoascia, 1022

Neochetina, 905

Neotettixfemoratus, 416

Neochoroterpes, 270, 286, 287, 308, 313, 336 Neochthebius, 793, 846, 847 Neochthebius vandykei, 893 Neocloeon, 269, 276, 306, 326 Neocloeon triangulifer, 60,294, 295 Neoclypeodytes. 816, 817, 819, 824, 880 Neoconocephalus, 417,420,427 Neoconocephalus lyristes, 420 Neocorixa, 529, 530

Neothremma, 589, 590, 655,660,663,671, 723, 764 Neothremma alicia, 88,655

Neocorixa snowi, 560 Neocurtilla, 417,428

Neocurtilla hexadactyla, 422 Neocylloepus, 858, 859, 864,866 Neocylloepus boeseli, 898 Neoelmis, 858, 865, 866 Neoelmis caesa, 898

Neoephemera, 197, 272,275, 297, 299, 334 Neoephemeridae, 167, 197, 272, 275, 297,334

Neoerythromma, 359, 361, 363 Neoerythromma cultellatum, 409 Neogalerucella, 868, 902 Neogerris, 537 Neogerris hesione, 554 Neohaemonia, 189, 868,902 Neohermes, 571, 572, 574, 575, 577, 578, 579, 583 Neohermes concolor, 576

Neohermes filicornis, 575, 576

Neohydronomus affinis, 869,906

Neothremma didactyla, 722 Neotrichia, 593,628,629, 630,694, 737 Neotridactylus, 205,421, 427 Neozavrelia, 1143, 1216, 1217, 1272 Nepa, 206, 522, 538, 542 Nepa apiculata, 557 Nepalomyia, 996 Nephrotoma, 1031 Nepidae, 49, 86, 136, 170, 187, 206, 237, 240, 522, 523, 525, 538, 542, 556-57 Nepinae, 538 Nepini, 538 Neporaorpha, 181, 206, 521, 522, 523, 524 Nepticula, 789 Nepticulidae, 170, 767,789 Nerophilus, 219 Nerophilus californicus, 645, 646,647,662, 711,713,744 Nerthra, 532, 534, 567 Nerthra martini, 534 Nerthrinae, 532

Neted sampler,26 Neureclip.sis, 216, 586, 591, 602,651,652, 718, 731 Neureclipsis bimaculata, 87, 719 Neureclipsis crepuscularis, 87 Neurobezzia granulose, 986 Neurocordulia, 341, 343, 379, 380,400

1035, 1041 Nitella, 812

Nixe, 83, 269, 270 Nobilotipula (Tipula subgenera), 1023, 1038, 1040 Nocticanace, 1004

Noctuidae, 88, 136, 170, 767, 769, 774, 785

Notiphila, 73, 108, 950, 953, 954, 979, 1013 Notiphila aenigma, 970 Notiphila caudata, 91 Notomicrus, 834, 835,882 Notonecta, 206, 522, 538, 563

Notonecta hoffmanni, 86 Notonecta insulata, 74 Notonecta undulata, 86, 100, 522

Notonecta unifasciata, 543 Notonectidae,49, 53, 54, 55, 73, 86, 100, 170, 206, 522, 523, 525, 538, 543, 562-63

Notonemouridae, 180, 181

Nuphar, 73, 868 Nyctiophylax, 586,602,651,652,676, 718,732

Nyctiophylax moestus, 719 Nyholmia, 849 Nymphaea, 73,118, 868, 869 Nymphomyia, 90, 222, 228, 932, 972,990 Nymphomyia walkeri, 935, 964

/

Index

Nymphomyiidae, 90, 168,222, 228, 926, 927, 928, 929, 930, 932, 935, 957, 964, 969, 972, 990

Nymphula, 106, 780 Nymphuta ekthlipsis, 780 Nymphuliella, 766, 770, 771 Nymphuliella daeckealis, 766, 781 Nymphulinae, 765 Nymphulini, 765

Oligophlebodes, 589, 655,660, 723, 763 Oligophlebodes minutus, 721 Oligophlebodes zelti, 88 Oligostigmoides, 766, 769, 771 Oligostigmoides cryptalis, 781 Oligostomis, 605,649, 715, 761 Oligostomis ocelligera, 648 Oligostomis pardalis, 659, 716 Oligotricha lapponica, 649,650, 715, 762 Oliveiriella, 1260 Oliveridea tricornis, 1167

Occidentalia comptulatalis, 784 Ochlerotatus, 1073, 1075 Ochlerotatus fitchii, 937 Ochrotrichia, 593,627,628, 697, 737 Ochteridae, 170, 523, 524, 525, 526, 534, 536, 567 Ochterus, 525, 534, 567 Ochterus barheri, 536 Ochthebius, 797, 805, 845, 846, 847, 893 Ochthera, 954, 1012 Ochthera mantis, 959 Oconoperla, 457, 497, 502 Oconoperla innubila, 457, 497, 500, 502, 517

Octogomphus specularis, 372, 374, 376, 395

Odonata, 2, 13, 37,49, 53, 56, 57,67, 72, 76, 80,81,82, 84,96, 97,98-100, 113, 115, 119, 120, 166, 169, 174, 176, 178, 179,180, 186, 189, 190, 199-201,232, 233, 236, 237,238, 239, 240, 341-409, 910 Odontella, 251

Odontoceridae, 133,134,167, 219, 586, 587, 589,610,611,642-46,662,663,673, 675,680,684, 711-12,713,74445 Odontomesa, 1178, 1180, 1196, 1199, 1232, 1253

Odontomyia, 224, 941, 961, 977, 1001 Odontomyia cincta, 942 Odontomyiina, 941 Oecetis, 220, 587, 630,631,673, 680,698,

Oliveridia, 1165, 1192, 1193, 1259 Omism, 1148, 1159, 1222, 1269 Omophron, 209, 802 Omophroninae, 802 Onconeura, 1260

Oncopodura, 250, 251 Oncopodurinae, 250 Onocosmoecus, 642,703, 758 Onocosmoecus unicolor, 88,643, 659, 702 Onychiuridae, 169, 247, 248, 249, 251,258 Onychiurinae, 247 Onychiurus, 248, 258 Onychiurus debilis, 246 Onychiurus dentatus, 245 Onychiurus litoreus, 246 Onychophora, 177 Onychylis, 906 Ophiogomphus, 199, 372, 375, 376, 395 Ophiogomphus carolm, 347, 354, 378 Ophiogomphus coluhrinus, 378 Opiinae, 909 Opius, 917 Optus caesus, 923 Oplonaeschna armata, 367, 368, 369, 371, 398

Optioservus, 802, 856, 860, 861, 866, 867, 868, 898

Optioservusfastiditus, 89 Optioservus phaeus, 856 Ora, 806, 849, 851,895 Oravelia, 523, 535

1471

Ornithodes, 1024 Orohermes, 571, 574

Orohermes crepusculus, 102, 572, 573, 575, 576, 584 Oroperla, 455,492 Oroperla barbara, 455,458, 517 Oropeza, 1031, 1036 Oropsyche, 622 Oropsyche howelae, 693 Oropsyche howellae, 622, 691, 727 Orphulella, 414, 424,425 Orthacheta, 1018 Orthacheta hirtipes, 91 Orthemis, 382, 389,403 Orthocladiinae, 134, 190,1123,1125, 1127, 1159-76, 1180, 1182, 1183, 1191-1213, 1230, 1236, 1240, 1241, 1253-64

Orthodadiini, 1236, 1240, 1241, 1254 Orthocladius, 966, 1165, 1167, 1168, 1169, 1191, 1192, 1193, 1198, 1199, 1200, 1201, 1203, 1206, 1211, 1212, 1214, 1232, 1260 Orthocladius annectens, 1174, 1179 Orthocladius calvus, 112 Orthocladius obumbratus, 91 Orthonevra, 1022

Orthopodomyia, 1073, 1079, 1082, 1083, 1084, 1087,1091,1096 Orthoptera,2,13, 113,166, 169, 174, 176, 180, 205, 232, 233,236, 237, 238, 239,240,411-28

Orthorrhapha,928 Orthorrhaphous,939 Orthotrichia. 629, 630,695,697 Osmylidae, 181, 234 Osobenus, 455,492

Osobenus yakimae, 455, 458, 460,492, 496,517 Ostrinia, 767 Ostrinia penitalis, 767, 784 Ostrocerca, 442, 447,474, 510

Oravelia pege, 539, 551

Ostrocerca albidipennis, 85,477

Orchelimum, 413,415,427 Orchelimum concinnum, 413,419

Ostrocerca truncata, 478 Oudemansia, 250

Orchelimum jidicinium, 413

Oudemansia georgia, 246,259

Oecetis inconspicua, 88,699

Orchesella, 252, 254 Orcus, 394

Oecotelma cushmani, 923

Orders, classification and key to, 231-43

Oulimnius, 860, 861, 865, 866, 898 Oxycera, 941, 1001 Oxyelophila, 766, 770, 771 Oxyelophila callista, 767, 111,782 Oxyethira. 626, 627,628, 629,665, 695, 697, 737 Oxyethira maya, 81 Oxyinae, 411 Oxyrhiza, 1048 Oxyrhizafuscula, 1029, 1033 Oxytelus, 797

742

Oecetis cinerascens, 664 Oecetis immobilis, 88

Oedenops nudus, 1014 Oedipodinae,412 Oedoparena, 926, 949, 979, 1005 Oemopteryx, 442,464, 509 Oemopteryx contorta, 442,468

Oemopteryx glacialis, 444, 445 Oemopteryx vanduzeea, 468 Oidematopsferrugineus, 1020 Olethreutidae, 767 Olethreutinae, 788 Oligia, 786 Oligochaeta, 133, 134 Oligoneuria, 327 Oligoneuriidae, 167, 197, 268, 273, 282, 291,299, 327

Ordobrevia, 858, 859, 862, 863

Ordobrevia nubifera, 898 Oreadomyia, 1260 Oreodytes, 814, 817, 825,827,880 Oreogeton, 944, 947, 999 Oreoleptidae, 168, 224, 925, 926, 929, 943, 945, 963, 967, 973,999

Oreoleptis, 224, 943, 999 Oreoleptis torrenticola, 945,967 Oreothalia, 999 Orimarga, 1024, 1045, 1067 Orimarga attenuata, 1047 Ormosia, 1023, 1024, 1051, 1063 Ormosia divergens, 1056 Ormosia lineata, 1054

Pachydiplax longipennis, 384, 385, 386, 387, 390, 403

Pachydrini, 824 Pachydrus, 814, 816, 824

1472

Index

Pachydrus princeps, 880 Paduniajeanae, 618, 619, 687, 688, 735 Paduniella nearctica, 656,681, 718, 720, 732 Pagastia, 91, 1177, 1178, 1206, 1207, 1252 Pagastiella, 1147, 1148, 1222, 1223, 1269 Palaeagapetus, 217, 592, 625, 691, 694, 738 Palaeagapetus celsus, 626, 629, 665 Palaemnema, 200, 348, 350, 387

Paradicranota, 1042, 1044 Paraglenanthe bahamensis, 1012 Paragnetina, 203,451, 487,490,514 Paragnetinafumosa, 452, 493 Paragnetina immarginata, 488 Paragnetina media, 85, 100 Paragyractis, 190 Parakiejferiella, 1165, 1167, 1202,

Palaemnema domina, 406

1204, 1260 Paralauterborniella, 1148, 1154, 1223, 1224, 1269

Palaeoptera, 232 Palearctic, 1029, 1030, 1032, 1033, 1034, 1044, 1047, 1048, 1052, 1054 Paleodictyopterida, 178 Paleolimnology, 150 Palingeniidae, 167, 268, 273, 290, 291, 300, 317,338 Palmacorixa, 529, 530, 533, 560 Palpada, 1022 Palpomyia, 221, 939, 987 Palpomyia pruinescens, 940 Palpotnyiini, 90

Paraleptophlebia, 53, 84, 128, 130, 270, 273, 286,313,336

Paraleptophlebia debilis, 285, 301 Paraleptophlebia guttata, 285 Paraleptophlebia strigula, 319 Paraleuctra, 204, 439,448, 479, 512

Paraleuctraforcipata, 481 Paraleuctra occidentalis, 450

Paratanytarsus grimmii, 234 Paratendipes, 1145,1148, 1222,1223, 1269 Paratendipes albimanus, 91 Paratendipes thermophilus, 1119 Paratettix. 418,426 Paratettix cucullatus, 416 Paratissa semilutea, 1006

Paratrichocladius, 1169, 1211 Paratrissocladius, 1194, 1261 Parochlus, 1177, 1182, 1183, 1184, 1238, 1250 Parochlus steinenii, 1119

Paronellinae, 252, 254 Parormosia, 1056

Parorthocladius, 1170, 1171, 1209, 1210, 1212, 1261

Paroxya, 423, 426 Paroxya atlantica, 412 Paroxya clavuliger, 412 Paroxya clavuligera, 412

Pultothemis, 354

Paraleuctra projecta, 481 Paraleuctra sara, 481,483 Paraleuctra vershina, 480

Paltothemis lineatipes, 383, 390, 403

Paralichus, 853

Parthina linea, 713

Paralichus ninyops, 906

Parthina vierra, 645, 647, 663

Paralimna, 1014

Parydra, 926, 954, 1009 Parydra quadrituberculata, 959

Panchaetoma, 253 Pancrustacea, 177 Panhexapoda, 177 Pantala, 384, 389, 391, 403

Pantalaflavescens, 385 Papaipema, 786 Paraboreochlus, 1124, 1177, 1181, 1183, 1184, 1238, 1250 Paracanace aicen, 1004

Paracapnia, 446,466,470, 513 Paracapnia angulata, 85, 447 Paracapnia horisi, 473 Paracapnia disala, 471 Paracapnia ensicala, 471 Parachaetocladius, 1160, 1166, 1172, 1202, 1205, 1260 Parachaetocladius abnohaeus, 91 Parachironomus, 1149, 1152, 1222, 1224, 1225, 1226, 1237, 1269 Parachironomus abortims, 1153

Parachironomusfrequens, 1153 Paracinygmula, 269,270 Paracladius, 1160, 1171, 1211, 1212, 1260

Paracladopelma, 1147, 1149, 1150, 1224, 1225, 1226, 1227, 1269 Paracladopelma doris, 1151 Paracladopelma galaptcra, 1151 Paracladopelma longanae, 1151 Paracladopelma rolli, 1151 Paracladopelma undine, 1151 Paraclius, 996

Paracloeodes, 278, 281, 310, 326 Paracloeodes minutus, 311

Paracoenia. 954, 1009 Paracoenia bisetosa, 960, 981

Paracricotopus, 1162, 1168, 1170, 1193, 1195, 1200, 1260

Paracymus, 838, 839, 843, 844, 846, 887 Paradelphomyia, 1023, 1027, 1029, 1033, 1046, 1065

Paradelphomyia fuscula, 1048

Paralimnophyes, 1260 Paramblopusa borealis, 889 Parameletus, 264, 275, 276, 303, 322 Parameletus croesus, 268

Paramerina, 1130, 1133, 1137, 1185, 1186 Paramerina smithae, 1132 Parametriocnemus, 1162, 1164, 1168, 1169, 1191, 1192, 1194, 1211, 1261 Paramormia, 990 Paranemoura, 446, 474, 510

Paranemoura perfecta, 447, 477 Paraneoptera, 180,181, 232, 233 Paranura, 250

Paranurophorus, 252 Paraperla, 77, 203,429,453,482, 519 Paraperlafrontalis, 456, 484,486 Paraperlinae, 519 Paraphaenocladius, 1164, 1168, 1169, 1170, 1205, 1206, 1261 Paraphrosylus, 996 Paraplea, 540, 541, 557 Parapoynx, 215, 766, 767, 768, 770, 771, 773, 782 Parapoynx allionealis, 768, 776 Parapoynx diminutalis, 167, 775 Parapoynx maculalis, 766 Parapoynx seminealis, 766 Parapsyche, 622,691, 727 Parapsyche apicalis, 87 Parapsyche cardis, 87,621 Parapsyche elsis, 693 Parargyractis, 765, 782 Parasimuliinae, 1109 Parasimulium, 1100, 1101, 1106, 1109, 1117 Parasimulium stonei, 1110 Parasmittia, 1261

Parasyntormon, 996 Paratanytarsus, 1143, 1144, 1146, 1213, 1214, 1215, 1216, 1272

r\

Paravelia, 523

Parthina, 646, 711, 745

Paskia, 52 Patasson, 919

Paucicalcaria ozarkensis, 627, 628,691, 694, 738 Paulinia acuminata, 411 Paxilla, 418,426 Pedicia, 223, 228, 933, 1023, 1024, 1029, 1042, 1058, 1068 Pedicia margarita, 1044 Pediciidae, 167, 223, 228, 1023, 1024, 1025, 1026, 1027, 1028, 1029, 1033, 1042, 1044, 1057, 1058, 1059, 1067 Pediciinae, 1067 Pedomoecus sierra, 612, 613,663, 683, 686, 747 Pelastoneurus, 996

Pelecorhynchidae, 168, 224, 941, 945, 963, 967, 973, 999 Pelenomus, 906

Pelina, 954, 1015 Pelina truncatula, 958 Pelocoris, 538, 558 Pelocoris shoshone, 542 Pelonomus, 806, 855, 857 Pelonomus obscurus, 894

Pelopia, 1250 Pelopiinae, 1244 Peloropeodes, 996 Peltodytes, 89, 104,210, 805, 812, 813,874 Peltoperla, 439, 443,479, 508 Peltoperla arcuata, 443,484 Peltoperlidae, 85, 134, 169, 202,429,431, 433, 437,440, 443,444, 462, 465, 483, 484, 507-8 Pemphigus trehernei, 524 Penelomax, 271

Penelomax septentrionalis, 286, 314, 318, 333 Pentacanthella, 252, 255

r

Index

Pentacora, 207, 540, 543, 565 Pentacora signoreti, 546 Pentagenia, 268, 291, 317, 338 Pentagenia vittigera, 273, 290, 300 Pentaneura, 1124, 1130, 1131, 1135, 1184, 1185, 1248 Pentaneurella, 1134, 1138 Pentaneurini, 1122, 1124,1127, 1230, 1235, 1247 Percidae, 343

Phanocerus, 856, 857, 860, 861 Phanocerus clavicornus, 899

Phanogomphus, 356, 373, 374, 375, 379, 395

Phanogomphus hodgesi, 352 Phanogomphus lividus, 375, 377 Phantolabis, 1025 Phantolabis lacustris, 1042, 1063 Phaonia, 1016 Pherbecta lemenitis, 1020

Pericoma, 938

Pherbellia, 91, 1020

Pericomaina, 990-91

Pherbellia quadrata, 970 Philanisus plebius, 66, 96, 104

Pericoraaini, 936

Perithemis, 354, 382, 390, 404 Perithemis tenera, 84

Perlesta, 429, 432,451,452,487, 490,492, 515

Perlesta placida, 85 Perlidae, 53, 85, 95, 100, 135,166, 169, 180, 203, 204, 429, 431,434, 435, 438, 439, 441,452,454,462, 463, 465, 467, 468,488, 489,491, 493, 494,513 Perlinae, 514 Perlinella, 451,487, 515 Perlinella drymo, 452,489,491 Perlinodes, 455,492 Perlinodes aurea, 518 Perlinodes aureus, 441,455,458,492 Perlodes, 80 Perlodidae, 53, 85, 100, 135, 166,169, 203, 204, 240,429,430,431, 432,436, 438,439,441,443,458, 460,461, 464,465,467, 468,494, 496,498, 500, 501,503, 504,516 Perlodinae, 516-18 Perlomyia, 439, 448, 476, 512

Perlomyia utahensis, 450,480 Petaluridae, 169, 200, 201, 342, 343, 347, 350, 353, 354, 368, 374, 393 Petersen grab, 39 Petersen-type grabs, 20 Petite Ponar grab, 22 Petrophila, 215, 765, 767, 768, 769, 770, 771,782 Petrophila bifascialis, 767 Petrophila canadensis, 88 Petrophila confusalis, 88, 189, 766, 767, 111, 773, 774

Petrophila drumalis, 765 Petrophilajaliscalis, 776 Petrophila santafealis, 768 Phaenocarpa antichaetae, 923 Phaenonotum, 836

Phaenopsectra, 1153, 1154, 1156, 1220, 1221,1229, 1269 Phalacrocera, 223, 1024, 1057, 1061 Phalacrocera replicata, 1032 Phalacrocera tipulina, 1058 Phaleria, 868, 891 Phaleria rotundata, 868

Phaneropterinae,427-28 Phanocelia canadensis, 633, 636,644, 704, 706, 758

Philarctus bergrothi, 638,639, 709, 758 Philocasca, 635, 703, 758 Philocasca demita, 88, 657 Philocasca rivularis, 637, 702 Philonthus nudus, 889

Philopotamidae, 77, 86-87, 135, 167, 216, 586, 591, 595, 598,603, 604,646, 647, 666, 669,676,677, 714, 729 Philopotamoidea, 729-30 Philorus, 932, 983 Philorus californicus, 935 Philorus yosemite, 935 Philotelma alaskense, 1009

Philygria, 1015 Phoridae, 168, 946, 948, 974, 979, 1017

Phryganea, 188, 588, 649, 650, 715, 762 Phryganea cinerea, 716 Phryganeidae, 87, 167, 219, 586, 588, 589, 596, 599,600,605,646-51,662, 664, 672,674,678,681, 715-17,760-62 Phycitinae, 767 Phygadeuon, 923 Phygadueontinae, 909 Phyllocycla, hll, 376, 388 Phyllocycla breviphylla, 388, 395 Phyllodromia, 947 Phyllogomphoides, 373, 376, 395 Phyllogomphoides stigmatus, 378 Phylloicus, 590, 616,687, 740 Phylloicus aeneus, 617,661, 687 Phylocentropus, 216, 586, 591, 603, 604, 616,617,670,672,678, 687, 730 Phylogeny, 175-92 Physa, 133 Physemus, 853 Phytobius, 51 Phytobius leucogaster, 907 Phytotelmatocladius, 1261 Pictetiella, 459,495, 497, 505, 518 Pictetiella expansa, 443, 504 Pictetiella lechleitneri, 498 Piestus, 797

Pilaria, 1026, 1027, 1046, 1049, 1065 Pilaria discicollis, 1029

Pillow cage, 40 Pistia, 767 Pistia stratiotes, 869

Placopsidella grandis, 1012 Plagioneurus univittatus, 996 Planiplax sanguiniventris, 382, 404 Plankton tow net, 38

1473

Platambus, 820 Plateumaris, 902 Plathemis, 354, 382, 389,404

Plathemis lydia, 381 Platycentropus, 606, 640, 706, 759 Platycentropus amicus, 707 Platycentropus radiatus, 641,658 Platygastridae, 170, 914,916 Platygastroidea, 909, 910, 912, 924 Platygymnopa helices, 1012 Platyhelminthes, 177 Platynota rostrana, 787 Platysmittia, 1162, 1174, 1194, 1196, 1197, 1261

Platystictidae, 169, 200, 348, 350, 387, 406 Platytipula (Tipula subgenera), 1024, 1038, 1043

Platyvelia, 523, 544, 548, 552 Platyvelia brachialis, 548 Plauditus, 269, 280, 293, 310, 326 Plauditus cestus, 311 Plauditus dubius, 294 Plecoptera, 2, 9, 10, 11, 12, 13, 14,15, 28, 37,49, 52, 53,55,67, 68,79,81,82, 84-86, 92, 95, 96, 97,100, 112,113, 114, 115, 120, 137, 145, 150, 166, 169, 174, 176, 178, 180, 181, 186, 187, 190, 202-4, 232, 233, 236, 237, 239, 240, 429-520 Plecopteracoluthus, 1191, 1193, 1196, 1228 Plecopterans, 180 Plectrocitemia, 651, 652, 656, 718, 732 Plectrocnemia cinerea, 719 Pleidae, 49,65, 170, 206, 522, 523, 525, 540, 541, 557 Plenitentoria, 746-64 Pleurogyrus, 923 Plhudsonia, 1209, 1261

Plumiperla, 453, 485,520 Plumiperla diversa, 456,467 Plutomurus, 250 Pneumia, 991

Pnigodes setosus, 907 Podisminae,426 Podmosta, 446, 447,474,476, 510 Podmosta delicatula, 478

Podonominae, 1124, 1126, 1177-79, 1180, 1183, 1230, 1250 Podonomini, 1177, 1235 Podura, 194,247 Podura aquatica, 245, 248, 258 Poduridae, 169, 194, 247, 248, 258 Poecilocera harisii, 902

Poecilographa decora, 1020 Pogonocladius, 1169,1200, 1203 Pogonognathellus, 250 Polycentropodidae, 87, 135, 167, 216, 586, 591, 594, 599,602,603, 651, 652-56, 669, 672,676,684, 718, 719, 731-32

Polycentropus, 651,652, 656, 669,676, 718, 732

Polycentropus centralis, 87 Polycentropus colei, 719 Polycentropusflavomaculatus, 60

1474

Index

Polycentropus maculatus, 87 Polychaeta, 66 Polygonum, 767 Polymera, 1046, 1065 Polymera georgiae, 1049 Polymitarcyidae, 84, 167, 197, 267, 271, 272, 273, 297, 300, 339 Polynema, 910,913 Polynema needhami, 920 Polynema striaticorne, 913 Polyneoptera, 180, 232, 233 Polypedilum, 110, 111, 1149, 1152, 1153, 1220, 1221, 1222, 1223, 1224, 1225, 1226, 1227, 1270 Polypedilum beckae, 1153 Polypedilum pemhai, 69, 76 Polypedilum vanderplanki, 69, 76 Polyphaga, 89, 182, 211-14, 791, 792, 794, 795,883-908 Polyplectropus, 651, 652, 718, 732 Polyplectropus charlesi, 719 Polytrichophora, 1012 Pomoleuctra, 448,476,479, 512 Pomoleuctra andersoni, 452,481,483 Pompilidae, 170,912,914,924 Ponar grab, 20, 23, 24, 39 Pontamalota, 847, 848 Pontomalota, 889

Pontomyia, 66, 1143, 1146 Postelichus, 793, 806, 855, 857, 894 Postelichus immsi, 80 Potamanthidae, 168, 197, 271, 272,273, 297, 300, 337 Potamanthus, 190, 271

Potamogeton, 118,766 Potamogeton pectinatm, 118 Potamophylax cingulatus, 114 Potamyiaflava, 621,623,624,625, 691, 692,693, 728 Potthastia, 1176, 1177, 1206, 1207, 1209, 1252 Potthastia gaedii, 1178

Prasocuris phellandrii, 902 Pratanurida, 250

Predator-Prey Index, 131, 132 Prestwichia, 912, 921 Primer bias, 160

Prinocyphon, 806 Prionocera, 933, 1024, 1035, 1058, 1069 Prionocera dimidiata, 1037

Prionocyphon, 214, 848, 849, 851,895 Probabilistic surveys, bioassessment, 155 Probezzia, 939,987 Procanace dianneae, 1004

Procladiini, 1128, 1235, 1246 Procladius, 966, 1128, 1129, 1131, 1132, 1135, 1187, 1188, 1189, 1234, 1239, 1246 Procladius hellus, 1239

Proclinopyga, 999 Procloeon, 269, 276, 279, 281, 292, 306, 326 Procloeon ingens, 295 Procloeon rivulare, 295

Procloeon simplex, 295

Procloeon viridoculare, 295 Proctotriipoidea, 912 Prodiamesa, 1180, 1196, 1199, 1253 Prodiamesa olivacea, 1178 Prodiamesinae, 1127, 1180, 1183, 11911213, 1235, 1252-53 Prodiamesinae, 1180 Progomphus, 343, 372, 374, 376, 396

Psephenidae,49, 55, 70, 89, 106, 122, 126, 133, 167, 182, 189, 190, 214, 243, 791, 792, 793, 794, 799, 802, 807, 808, 84952, 893

Progomphus borealis, 80 Progomphus obscurus, 374 Progonomyia, 1055 Proischnura subfurcatum, 56

Psephenivorus, 919 Psephenoides, 189 Psephenus, 70, 126, 189, 190, 807, 849, 850, 851,852, 893 Psephenusfalli, 106 Psephenus herricki, 849 Psephenus montanus, 89 Psephidonus, 890

Proisotoma, 253, 257

Pseudachorutes, 249

Prokelisia marginata, 524

Pseudanteris insignis, 924

Promoresia, 856

Pseudanurida, 247, 250 Pseudanurida sawayana, 246, 259

Promoresia elegans, 89 Pronoterus, 833

Propsilocerus, 1180, 1211, 1261 Prosimuliini, 1109 Prosimulium, 1100, 1104, 1106, 1109, 1117 Prosimulium clandestinum, 1107 Prosimulium fuscum, 90 Prosimulium mixtum, 90, 1102, 1103, 1105 Prosimulium unicum, 1112 Prosimulium ursinum, 97, 1107, 1110, 1111 Prostoia, 446,474,476, 510 Prostoia besametsa, 85,477

Pseudanurophorus, 252 Pseuderipternus, 923 Pseudiron, 75, 197, 268 Pseudiron centralis, 83, 282, 302, 304, 328 Pseudironidae, 83, 168, 197, 268, 282, 302, 304, 328 Pseudohourletiella, 261

Pseudocentroptiloides, 276, 292, 306, 311,326 Pseudochironomini, 1122, 1127,

Prostoia similis, 85

1230, 1271 Pseudochironomus, 1127, 1181,1216, 1218, 1223, 1224, 1225,1238, 1239, 1271

Protachoutes, 250

Pseudochironomus articaudatus, 1181

Protanyderus, 228, 992 Protanypini, 1252 Protanypodini, 1176, 1235 Protanypus, 1122, 1176, 1178, 1200, 1204,

Pseudochironomusfulviventris, 1181 Pseudochorthippus, 423

1234, 1252 Protanypus ramosus, 1178 Protaxymyia, 934 Protaxymyia thuja. 935, 983 Protochauliodes, 571, 574, 578, 584

Protochauliodes spenceri, 575, 576, 579 Protoneura, 359, 363 Protoneura capillaris, 352, 366 Protoneura cara, 409 Protoneura viridis, 351 Protoneuridae, 169, 366 Protoneurinae, 355, 359

Pseudocloeon, 268 Pseudocorixa, 560 Pseudodiamesa, 1176,1178, 1207, 1209, 1252

Pseudogoera singularis, 642,645,663,684, 711,713,745 Pseudohyadina longicornis, 1015 Pseudokiefferiella, 1177, 1206, 1207, 1252 Pseudolampsis guttata, 868 Pseudoleon superbus, 384, 389, 391, 404 Pseudolimnophila, 1024, 1025, 1027, 1029, 1033, 1046, 1066

Protoplasa, 223, 971 Protoplasafitchii, 935, 964,992 Protoptila, 618,619,690, 735 Protoptila erotica, 688 Protoptila maculata, 668,688 Protoptilinae, 687, 735

Pseudolimnophila inornata, 933 Pseudolimnophila luteipennis, 1048 Pseudolimnophila sepium, 1029, 1033 Pseudoneureclipsidae, 586 Pseudonychiurus, 258 Pseudonychiurus dehilis, 246 Pseudorthocladius, 91, 1160, 1172, 1205, 1206, 1262

Protosialis, 571, 572, 574, 582 Psamathobledius, 890 Psammisotoma, 253 Psammocladius, 1160, 1171

Pseudosinella, 250 Pseudosinella ahainaensis, 260 Pseudosinella lahainensis, 246 Pseudosmittia, 940, 1162, 1201, 1213,1262

Pschomyiidae, 720

Pseudosmittia gracilis, 1234

Psectrocladius, 1160, 1162, 1171, 1172, 1194, 1195, 1196, 1199, 1228, 1261 Psectrotanypus, 1 128, 1 135, 1 187, 1189, 1246

Pseudosmittia mathildae, 1176

Psectrotanypus dyari, 1131, 1132 Pselactus spadix, 907 Pselaphidae, 793

Psilidae, 960 Psiloconopa, 1054 Psilocricotopus, 1196

Pseudostenophylax, 219, 590,635, 703, 759 Pseudostenophylax edwardsi, 637 Pseudostenophylax sparsus, 637, 704

r

e

Index

Psilometriocnemus, 1164, 1168, 1174, 1211, 1212, 1229, 1262 Psilopa, 1006

Psilotanypus, 1188 Psilotreta, 587, 589, 610,645,646,647, 673,680, 711,745 Psilotreta amera, 662 Psilotreta labida, 713 Psorophora, 1071, 1072, 1073, 1079,1081, 1084, 1085, 1087, 1096 Psychoda, 975, 991

Psychoda alternata, 964 Psychodidae,49, 167, 222,228, 926, 927, 928, 930, 936, 938, 957, 962, 964, 969, 975, 990 Psychodini, 936, 991 Psychodomorpha, 185 Psychoglypha, 220, 590, 635,636, 706, 759 Psychoglypha mazamae, 657 Psychoglypha subborealis, 705 Psychomyia, 216, 591, 602, 656,669, 676, 718,733 Psychomyiaflavida, 653, 720 Psychomyiidae, 87, 135, 136, 167, 190, 216, 586, 591, 599,602, 603,653, 656, 669,612,616,681,682,718, 732-33

Psychomyioidea, 730-33 Psychopomporus, 816, 826 Psychopotnporusfelipi, 880 Psychoronia, 642, 709, 759 Psychoronia brooksi, 710 Psychoronia costalis, 643, 659 Pteromalidae, 170, 920 Pteromalidae, 910, 911, 913, 914, 916

Pull-up trap, 21 Pycnopsyche, 88, 130, 137, 634,636, 637, 657, 709, 760 Pycnopsyche guttifer, 56, 110, 137 Pycnopsyche lepida, 56, 140, 586 Pycnopsyche scabripennis, 708 Pyractomena, 213 Pyractonema, 868 Pyralidae, 49, 767, 770 Pyraloidea, 765, 769 Pyraloidea, 770 Pyrausta, 784 Pyraustinae, 767, 784 Pyrrhalta, 73, 110, 903

Quart jar rearing method,40

Radotanypus, 1128, 1136, 1187, 1189, 1246

Ramphocorixa, 530, 533, 560 Ramphocorixa acuminata, 100 Ranatra, 206, 538, 557 Ranatra brevicollis, 542 Ranatra montezuma, 86

Ranatrinae, 538 Ranunculus, 118

Raphidioptera, 181 Raphidocladius, 1206, 1229 Rapoportella, 249 Raptoheptagenia, 196 Raptoheptagenia cruentata, 282, 310, 316, 330 Rasvena, 455,485

Pteronarcella, 430,439, 482,507 Pteronarcella badia, 440,484

Rasvena terna, 456, 458,485,486, 488, 520 Recolonization, 18 Reference condition approach (RCA),

84-85, 134, 169, 202, 204, 237,240, 429,430, 431,433,437, 440, 462, 467,484, 507 Pteronarcys, 10, 11, 12, 14, 15,202,204, 430,439, 482, 507 Pteronarcys dorsata, 84, 437, 467 Pteronarcys proteus, 84 Pteronarcys scotti, 85 Pterygota, 177, 180, 232 Ptiliidae, 168, 793, 795, 799, 804, 805, 809, 845, 883 Ptilocolepinae, 592 Ptilodactylidae, 89, 167, 214, 792, 800, 802, 807, 808, 853, 854, 899-900 Ptilomyia, 1014 Ptilomymar, 910 Ptilomymar magnificum, 910,920 Ptilostomis, 588,648,649, 650, 674, 715, 762 Ptilostomis ocellifera, 717 Ptychoptera, 934, 937,992 Ptychoptera lenis, 90, 108, 964 Ptychopteridae, 49, 90, 108, 168,222, 226, 241, 243, 925, 926, 927, 930, 931,932, 937, 962, 964, 969, 971, 991-92, 1028

Rhaphium slossonae, 968 Rheocricotopus, 1160, 1171,1193,1194, 1195, 1196,1262 Rheopelopia, 1133, 1134, 1138, 1189, 1190,1249 Rheosmittia, 91, 1159,1163,1198, 1201, 1228, 1262 Rheotanytarsus, 190, 1142, 1143, 1144, 1216, 1217, 1273 Rheotanytarsus exiquus, 91 Rheumatobates, 527, 535, 536, 555 Rheumatohates rileyi, 536 Rhinonapaea metallica, 1009 Rhinoncus, 907 Rhionaeschna, 367, 368, 387, 398 Rhionaeschna multicolor, 388 Rhithrogena, 71, 190, 283, 285, 307, 312,330

Pteromicra, 1020

Pteronarcyidae, 9, 10,11, 12, 14, 15, 53,

1475

144-45

Reference studies, bioassessment, 152-55 Remartinia, 367, 398 Remartinia luteipennis, 367, 368 Remenus, 455, 499, 502, 518 Remenus bilobatus, 458 Renocera, 949, 951, 1020 Reomyia, 1139, 1184,1185 Respiration, 43-63 Rhabdomastix, 1024, 1042,1051, 1063

Rhabdomastix setigera, 1044 Rhabdomastix trichophora, 1056 Rhagadotarsinae, 555 Rhagadotarsus, 100 Rhagovelia, 207, 527, 544, 547, 552 Rhagovelia distincta, 547 Rhagoveliinae, 544 Rhamphomyia, 229, 947, 999 Rhantus, 815, 819, 820, 833, 880 Rhantus binotatus, 89

Rhaphidolabis(Dicranota subgenera), 1042

Rhaphidophoridae,412 Rhaphium, 947, 996 Rhaphium cainpestre, 947

Rhithrogena semicolorata, 61, 83 Rhizelmis, 858, 859, 863, 864 Rhizelmis nigra, 899 Rhyacophila, 80, 87, 217, 220, 592,601, 656, 667,676, 718,739 Rhyacophilidae, 87, 122, 135, 167, 190, 217, 220, 238, 586, 589, 592, 599, 601,603, 653, 656,666,667,676, 681,718-20, 738-39 Rhypholophus, 1024, 1050, 1053, 1063 Rhysophora, 1007 Rickera, 455,497, 505 Rickera sorpta, 458,497, 505, 519 Riffle sampler, 18 Robackia, 1122, 1149, 1225, 1270 Robackia claviger, 1151 Robackia demeijerei, 91, 1151, 1153 Rock-outcrop sampler, 18 Roederiodes, 946, 999 Romalea, 421,426 Romalea microptera, 412 Romaleidae,412,415,421 Romaleinae, 426 Rossiana, 219, 590,606 Rossiana montana, 654,656, 662, 720, 721,763 Rossianidae, 167, 219, 586, 590, 606, 611, 654,656, 662, 671,675,680, 681, 720, 721,762-63 Rupisalda, 540, 543, 546, 565 Rusekella, 250

Sacodes, 849, 895 Saetheria, 1151, 1159, 1225, 1227, 1270 Saetheria tylus, 1151 Saetheriella, 1262

Salda, 540, 543, 545, 546, 566 Salda buenoi, 545 Saldidae, 166, 170, 207, 521, 522, 525, 526, 540, 543, 545, 546, 565-66 Saldinae, 540 Saldoida, 540, 545, 566 Saldula, 526, 544, 545, 546, 566 Saldula pexa, 545

1476

Index

Salina, 252 Salina banksi, 260

Salmonidae, 343 Salmoperla, 455, 492

Salmoperla sytvanica, 441,492, 518 Salpingidae, 168, 793, 794, 795, 800, 801, 807, 808, 868, 891

Sahelinusfontinalis, 79 Salvinia, 869

Samea muttiplicaiis, 767, 785 Sameodes alhigutlalis, 767 Samples and statistical considerations, 30-34

Sanfilippodytes, 817, 825, 826, 881 Sarabandus, 849

Sarabandus robustus, 895

Sarcophaga, 913, 949 Sarcophaga dux, 970 Sarcophagidae, 168, 225, 949, 953, 974, 1017

Sargus, 1001 Sasquaperla, 453, 485

Sasquaperla hoopa, 453,456, 485, 488,520 Saunderia, 1263

Savchenkia (Tipula subgenera), 1031, 1038 Savtshenkia (Tipula suhgeneta), 1024, 1036 Scapteriscus, 428

Sensiphonrura, 251 Sensiphorura, 247, 249 Sepedomerus, 949 Sepedomerus macropus, 1021 Sepedon, 91, 108, 225, 949, 951, 1021 Sepedonfuscipennis, 970 Sepedon tenuicornis, 951 Sergentia, 66, 1154, 1220, 1229, 1270 Sericomyia, 1022 Sericostomatidae, 88, 167, 219, 586, 587, 589, 596,604,611, 654, 660,662,670, 675,678,684, 720-23, 745-46

Simuliini, 1109 Stmullum, 108, 190, 223, 228, 1097, 1100, 1101, 1104, 1109, 1113, 1118

Sericostriata, 219, 589

Slmullum trtbulatum, 1102

Sericostriata surdickae, 655,659, 660, 722, 723, 764 Serratella, 83, 271, 288, 314, 333 Serratellafrisoni, 314 Serratella ignita, 83, 98

Stmullum venustum, 938, 1102, 1110 Stmullum vlttatum, 964, 1103, 1108 Stmyra henrlct, 787 Stmyra tnsularls, 767 Stnella, 252

Serratella levis, 308 Serratella serratoides, 290, 308 Serromyia, 987 Setacera, 226, 950, 1009 Setacera atrovirens. 956 Setodes, 630,698, 743 Setodes dixiensis, 664 Setodes incertus, 631,632

Single core sampler, 24, 38 Single corer with pole, 19 Single-plate sampler, 20 Slnotlpula (Tipula subgenera), 1023, 1025,

Setodes oligius, 700

Siphlonuridae, 68, 119, 167, 197, 268, 274,

Scarabandus, 849

Setvena, 455,492, 518

Scatdla, 954, 1009 Scatella hawaiiensis, 958

Setvena bradleyi, 467 Shipsa, 446, 474, 476 Shipsa rotunda, 85,446, 447, 474, 476,

Scatellapicea, 953, 958 Scatella thermarum, 67

Scathophagidae, 91, 167,225, 925, 928, 957, 960, 961, 974, 980, 1017-18

Scatophila, 954, 1010 Scatophila iowana, 956,958 Scelionidae, 912 Scellus, 996

Schaefferia, 249

478,511

Shovel sampler, 26 Shredder Index, 131, 132 Sialidae, 73, 86, 102, 135, 170, 208, 237, 240, 241, 569, 570, 571, 572, 573, 574, 582 Stalls, 73, 86, 208, 571, 572, 574, 582 Stalls aequalls, 86

Stmullum anatlnum, 1111 Stmullum arcttcum, 1108

Stmullum argyreatum, 51 Stmullum bracteatum, 1105

Stmullum jenntngst, 90 Stmullum mertdtonale, 1111 Stmullum montlcola, 51 Slmullum noellerl, 112

Slmullum plctlpes, 69

1037,1038 Slphlontsca, 268 Slphlontsca aerodromta, 68, 264, 276,277, 303, 322

275,277, 300, 303,307, 321-22

Slphlonurus, 198, 274,211, 300, 303, 307, 322

Slphlonurus alternus, 67 Slphlonurus occldentalls, 52 Slphloplecton, 197, 275, 284, 304, 313, 323 Stsko, 646, 712, 730 Stsko oregona, 646 Stsko stsko, 646, 714

Slsyra, 97, 578, 580, 584 Slsyra vlcarla, 580 Sisyridae, 49, 86, 97, 170, 181, 208, 234, 237, 240, 241, 569, 570-71, 578, 579,

Schema salinum, 1014 Schistocerca, 414,421,425

Stalls callforntca, 573

Schizoptera bispina, 553 Schizopteridae, 170, 521, 553

Stalls dretsbacht, 86 Slalls hamata, 572

Skutzla, 1273

Schizoramia, 177

Stalls hasta, 573

Schoenobiinae, 767, 770, 771, 111, 783

Stalls rotunda, 102, 572 Stera ntcoya, 246 Sterracapnta, 446,470,472, 513 Sterracapnta washoe, 473 Sterraperla, 439, 482, 508 Sterraperla cora, 444 Sieving, 34 Slgara, 206, 532, 561 Slgara decoratella, 534 Stgarafallenotdea, 533 Slgara Itneata, 534 Stgara mathesonl, 533 Stgara mcklnstryl, 534 Slgara mullettensls, 534 Sllvlus, 943, 944, 1003 Simuliidae, 37,49, 51, 55, 77, 90, 92, 108, 115, 135, 167,223,228,238,241,243, 925, 926, 927, 928, 929, 930, 936, 938, 957, 964, 973, 976, 1097-1118

Skwala, 203, 457,495, 518 Skwala amertcana, 436,460, 496,498 Smlcrtdea, 622,691, 729 Smlcrtdeafasclatella, 621, 623, 692 Smicrideinae, 586,691, 729 Sminthuridae, 169, 194, 256, 257, 260-61 Smlnthurldes, 245, 247, 256, 260 Smlnthurldes aquatlcus, 257 Smlnthurlnus, 194,256, 257 Smlnthurus, 256, 257 Smlnthurus Incognttus, 261 Smlttla, 1162, 1164, 1174, 1200, 1262 Smlttla aquattlls, 1164 Sollperla, 439,482,508 Sollperla campanula, 484 Sollperla saltsh, 484 Somatochlora, 199, 356, 380, 400 Soyedtna, 442, 445, 474, 511 Soyedlna valltcularla, 85 Soyedtna washtngtonl, All Spanglerogyrus, 810, 811

Schoetella, 249

Schummelia {Tipula subgenera), 1035, 1039 Sciomyzidae, 37, 91, 108, 167, 186, 225, 226, 229, 241, 926, 927, 928, 929, 949, 951,952, 961,970,974,979, 1018-21, 1027 Scirpus, 767 Scirtes, 849, 851, 895 Scirtes tibialis, 89

Scirtidae, 89, 167, 214, 243, 792, 795, 799, 802, 806, 808, 848, 849, 851, 894-95

Scleroprocta, 1023, 1030, 1050, 1053, 1064 Scopeumatidae, 1017 Scudderia, 417, 428 Scudderia texensis, 420

Scydmaenidae, 793 Seira, 250 Semicerura, 250, 251 Sensillanura, 250

Stalls cornuta, 572

Simuliinae, 1109

r\

580, 584

Slsyrldlvora cavlgena, 920

r

Index

Spanglerogyrus alhiventris, 872 Sparharus, 289, 296, 317, 335 Sparbarus choctaw. 296 Sparharus maculatus, 295, 296, 305 Sparbaus, 269 Sparganothis sulfureana, l&l Spartina, 413, 524 Spaziphora cincta, 1018 Sperchopsis, 839, 840, 843, 844 Sperchopsis tesselata, 887 Sphaeridiinae, 836, 837, 839 Sphaeriidae, 49, 51 Sphaerius, 799, 801, 803, 805, 836, 883 Sphaeriusidae, 168, 793, 794, 799, 801, 803, 805, 836, 883 Sphaeromias, 987 Sphaeromiini, 90 Sphagniana, 415 Sphagniana sphagnorum, 413,419 Sphagnophylax meiops, 88,640,641, 644, 701,703, 760

Sphingidae, 767 Sphyrotheca, 256,257 Spilogona, 1016 Spilomelinae, 767, 784-85 Spilomyia, 1022 Spinactaletes. 250, 260 Spinactaletes honeti, 246 Spinactaletes inyoptesimus, 254 Spinactaletes nemyops, 246 Spinadis, 269 Spinadis simplex, 283, 310, 316, 330 Spinisotoma, 252 Spodoptera pecticornis, 769 Spodoptera pectinicornis, 767 Stachanorema, 252

Stackelbergina, 1124, 1170, 1173, 1211, 1262

Stactobia, 696 Stactobiella, 627,697, 738 Stactobiella delira, 626, 629

Staphylinidae, 166, 167, 214, 793, 794, 795, 797, 798, 799, 802, 803, 847^9, 887-90

Stationary screen trap, 19 Stegopterna, 1100, 1101, 1104, 1106, 1113, 1118 Stegopterna acra, 1111 Stegopterna mutata, 1102, 1103, 1105, 1108, nil

Steinovelia, 523, 544, 548 Steinovelia stagnalis, 552 Stempellina, 1142, 1145, 1215, 1216, 1273 Stempellinella, 1139, 1142, 1216, 1218, 1273 Stenacris, 421, 425 Stenacris vitreipennis, 412 Stenacron, 198, 283, 285, 307, 312, 330 Stenacron interpunctatum, 83, 264 Stenelmis, 51, 211, 806, 856, 858, 859, 861, 862, 863, 899

Stenochironomus, 1151, 1153, 1154, 1225, 1227, 1270 Stenocolus, 212, 800, 808, 853, 854 Stenocolus scutellaris, 853,900 Stenogastrura, 249

Stenogastura, 251 Stenogomphurus, 373, 377, 396 Stenonema, 83, 269, 283, 285, 307,312, 331 Stenonemafemoratum, 269, 283, 285, 307,312

Stenopelmatidae,412 Stenopelmus rufinasus, 907 Stenophylax, 190 Stenopsychidae, 586

1477

Suragina, 941 Suragina concinna, 967,993 Surber sampler, 18, 19, 26, 27, 28, 38

Surface film sampler, 19 Susperatus, 269, 289, 315, 335 Susperatus prudens, 296, 305 Susperatus tuberculatus, 296 Susulus, 457,495 Susulus venustus, 457,460, 495, 498, 518 Suwallia, 453, 485, 520 Suwallia pallidula, 456 Suwallia starki, 488 Sweep net, 21 Sweltsa, 86, 203,204,453, 485, 520

Stenotabanus, 944, 1003 Stenus, 847, 890 Steremnius, 907

Sweltsa horealis, 486

Stethophyma, 423,425

Sweltsa oregonensis, 438, 456,486 Sweltsa palearata, 440,443 Symbiocladius, 1161, 1166, 1172, 1174, 1198, 1201, 1263 Sympetrum, 341, 342, 383, 384, 390, 404 Symplecta, 1024, 1051, 1064 Symplecta hybrida, 1056 Symplecta pilipes, 1050, 1053 Symplecta rainieria, 1054 Symposiocladius, 925, 1165, 1167, 1211 Sympotthastia, 1177, 1191, 1192, 1252 Sympycnidelphus, 996 Sympycnus, 996 Synaptonecta, 538 Synaptoirecta issa, 549, 561 Synclita, 765, 780 Syndiamesa, 1177,1179, 1206, 1229, 1252 Synendotendipes, 1154, 1157, 1225, 1271 Synorthocladius, 1170, 1171, 1202,

Stictochironomus, 128, 1148, 1159, 1222, 1270 Stictocladius, 1262 Stictotarsus, 816, 817, 825, 827, 881 Stictotarus, 817

Stigmellidae, 789 Stilobezzia, 939, 987 Stilobezzia bulla, 966 Stilocladius, 1170, 1173, 1211,1212,1263 Stovepipe sampler, 18, 19,23, 38 Stratiomyia, 1001 Stratiomyidae, 49, 167, 910, 926, 927, 928, 939, 942, 961, 963, 967, 973, 977, 1000-1001

Stratiomyiidae, 224, 229 Stratiomys, 939, 942, 1001 Stream bottom T-sampler, 38 Strophopteryx, 442, 464, 509 Strophopteryx appalachia, 442 Strophopteryx cucullata, 469 Strophopteryxfasciata, 442, 444, 445,468 Strophopteryx limata, 85 Stupkaiella, 936, 991 Stygoparnus, 855 Stygoparnus comalensis, 77, 894 Stygoporus, 792, 816, 826 Stygoporus oregonensis, 881 Stylogomphus, 372, 377, 396 Stylogomphus albistylus, 375 Stylurus, 346, 373, 377, 396 Stylurus ivae, 381 Stylurus laurae, 375 Subaquatic light trap, 40 Subaquatic light traps, 22 Suhlettea, 1142, 1143, 1144, 1213, 1215, 1273

Sublettiella, 1263 Subsampling, 34

Suction samplers, 18, 19, 20, 21, 22, 23

Stenelmis crenata, 82, 89 Stenelmis nr. bicarinata, 89

Sulcarius, 910 Sulcarius medius, 923 Superodontella, 249 Suphis, 805, 833, 834,835 Suphis inflatus, 882 Suphisellus, 833, 835, 882

Stenelmis sexlineata, 89, 106

Suphisellus puncticollis, 89

Sweltsafidelis, 488 Sweltsa onkos, 86

1205, 1263

Syntormon, 996 Syrphidae,49, 50, 73, 167, 192, 225,229, 243, 926, 927, 928, 929, 930, 946, 948, 961,970,974,978, 1021-22 Systenus, 997

T-sampler, 18 Tabanidae, 37, 49, 73, 91, 108, 167,186, 224, 229, 238, 925, 926, 927, 928, 929,930, 941,945,961,963, 967, 973, 978, 1002-3 Tahanus, 944, 945, 961, 1003 Tahanus atratus, 91, 108 Tahanusfumeus, 91 Tahanusfumipennis, 91 Tahanus reinwardtii, 945 Tachopteryx thoreyi, 368, 374, 393 Tachytrechus, 997 Taenionema, 442,464, 509 Taenionema atlanticum, 469

Taenionema pacificum, 444, 445 Taenionema raynorium, 468 Taeniopterygidae, 53, 85, 169, 202, 204, 240,429, 430,431,433, 437, 440, 444, 445, 462,463,468, 469, 508

1478

Index

Taeniopteryginae, 508-9 Taeniopteryx, 53, 190, 202, 204, 439,444, 464, 509 Taeniopteryx maura, 437,440, 463 Taeniopteryx metequi, 468 Taeniopteryx nivalis, 85 Taeniorhynchus, 48, 50 Tafallia, 249, 251 Tallaperla, 110, 202,439,443,479, 508 Tallaperla anna, 465,483 Tallaperla maria, 85, 111,437, 484

Tendipedidae, 1243 Tendipes, 1265 Tenebrionidae, 167, 793, 795, 800, 801,

Thoronella, 910,924

808, 868, 891 Tetanocera, 91, 949, 1021 Tetanocera loewi, 91 Tetanocera montana, 952 Tetanocera vicina, 952 Tetanoceratidae, 1018 Tetanoceridae, 1018

Thraulodes brunneus, 290 Thremmatidae, 167, 586, 589, 590,605, 607,655,660,664,677,681, 721,

Tanarthrus, 868

Tetracanthella, 252, 255 Tetragoneuria, 84, 100, 380, 400 Tetragoneuria cynosura, 84

Tanychela, 912 Tanychela pilosa, 909,923 Tanyderidae,49, 168, 224, 228, 926, 927, 928, 930, 935,936, 962, 964, 969, 971, 992, 1028 Tanypodinae, 136, 1122, 1123, 1124, 1126, 1127-39, 1129, 1180, 1183-90, 1228, 1230, 1235, 1244 Tanypodini, 1128, 1235, 1249 Tanypteryx, 200, 201 Tanypteryx hageni, 354, 368, 393 Tanypus, 1128, 1129,1135,1189, 1190, 1233, 1239, 1250 Tanypus concavus, 1135 Tanypus neopunctipennis, 1135 Tanysphyrus, 802, 907 Tanytarsini, 77, 135, 1122,1123,1127, 1139-^5, 1230, 1235, 1271-74 Tanytarsus, 73, 1143, 1144, 1145, 1146, 1213, 1215, 1216, 1217, 1218, 1229, 1233, 1237, 1273 Tanytarsus halteatus, 191 Tanytarsus dissimilis, 58 Tardigrada, 177 Targeted studies, bioassessment, 155

Tethymyia, 1236, 1240, 1254

Tetrastichus, 913

Tetrastichus polynemae, 919 Tetrigidae, 169, 205, 411, 412, 415, 416, 418,426

Tetriginae, 426-27 Tetrix, 205,418,427 Tetrix subulata, 416

Tettigidea, 205, 418, 426 Tettigoniidae, 169, 205, 237,411,413, 415, 417,419,420,427

Tettigoniinae, 428 Teuchogonomyia, 1055 Teuchophorus, 997 Thalassaphorura, 258 Thalassaphorura debilis, 246 Thalassaphorura halophila, 246 Thalassaphorura litoreus, 246 Thalasselephas testaceus, 908 Thalassomya, 1240, 1241, 1243 Thalassosmittia, 1236, 1240, 1241, 1263 Thalassotrechus barbarae, 873

Thambemyia borealis, 997 Thaumalea, 940,975

Tarnetrum, 404

Thaumalea verralli, 992

Tarphiota, 890 Tauriphila, 383, 390,404 Tauriphila argo, 341

Thaumaleidae, 168, 223, 228,926, 927, 928, 929, 930, 936, 940, 962, 965, 969, 975, 992 Thaumastoptera hynesi, 1045, 1067

Tavastia, 1263 Taxodium, 767

Telebasis, 358, 363, 409 Telebasis salva, 84, 364 Telenomus, 924

Telmatogeton, 66, 1231, 1241, 1243 Telmatogetoninae, 1230, 1236, 1240, 1241, 1243

Telmatogon Japonica, 80 Telmatopelopia, 1134, 1138, 1188 Telmaturgus parvus, 997 Teloganopsis, 271 Teloganopsis deficiens, 286, 290, 314, 318,333 Teloleuca, 540, 545, 566 Teloleuca bifasciata, 546 Telopelopia, 1133, 1134, 1185,1249 Telopelopia okoboji, 1141

Theliopsyche, 588,630, 631,664, 697,750

Theliopsyche corona, 698 Theliopsychinae, 750 Thermonectus, 815, 818, 819, 829, 830, 881

Thoronidea, 924 Thraulodes, 285, 286, 308, 313, 337

723, 763 Threticus, 991

Throscinus, 853,900

Timpanoga, 271, 289, 315 Timpanoga hecuba, 333 Tinodes, 653,656, 718, 733 Tinodes cascadius, 720 Tiphodytes gerriphagus, 910, 916, 924 Tipula, 13, 15, 123, 134, 137, 223, 1023, 1024, 1025, 1026, 1031, 1032, 1035, 1070 Tipula abdominalis, 90, 1041

Tipula caloptera, 1040, 1058 Tipula commiscihilis, 1039 Tipula eluta, 965 Tipula hovsgolensis, 1041 Tipula ignobilis, 1036 Tipula luteipennis, 1043 Tipula nobilis, 1040 Tipula oropezoides, 1039 Tipula paludosa, 1041 Tipula pruinosa, 1040, 1056 Tipula pura, 1043 Tipula rohweri, 1036 Tipula sacra, 90, 106, 1041 Tipula salicetorum, 1041 Tipula spenceriana, 1043 Tipula strepens, 933, 1040 Tipula synchroa, 1039 Tipulidae, 13, 37,49,81, 82, 90, 106, 115, 136, 137, 167, 223,238, 241, 243,928, 1023-70

Tipuloidea, 185, 926, 927, 929, 930, 931, 933, 962, 965, 969, 971

Tlalocomyia, 1106, 1109,1118 Tlalocomyia ramifera, 1103, 1110 Tokunagaia, 1200, 1203, 1208, 1209, 1210, 1263 Tomoceridae, 194 Tomocerina, 250

Tomocerinae, 250 Tomocerus, 194, 250, 251

Thermonectus ornaticollis, 89

Tomolonus, 250

Thienemannia, 1211,1263 Thienemanniella, 1159, 1163, 1191, 1192,

Torridincolidae, 49, 51

1254

Thienemannimyia, 1133, 1134, 1139, 1140, 1141, 1189, 1190, 1249 Thienemanniola, 1139, 1142 Thinobius, 890

Thinophilus, 997 Thinopinus, 797, 802, 847,848 Thinopinus pictus, 890

Tortopsis, 264, 265, 271, 291, 317, 339 Tortopsis primus, 290 Tortopus, 271,291,317,339 Tortopus circumfluus, 271 Tortopus igaranus, 271 Tortriddae, 170, 767, 787 Tortricinae, 787-88 Tournotaris, 908

Toxidty testing, 150-51 Toxorhynchites, 1072, 1073, 1075, 1077,

Temelucha chilonis, 923 Temnostoma, 225, 946

Thinoscatella, 1010 Thinusa, 847, 890

Tempisquitoneura, 1162, 1174, 1263 Tenagobia, 529, 538 Tenagobia mexicana, 561

Tholymis citrina, 382, 389, 391, 404 Thornburghiella, 936, 991

Trachypachidae, 182

Thoron, 924

Trapezostigma, 404

1079, 1082, 1087, 1088, 1096 Tramea, 199, 384, 385, 391,404

Index

Trathala, 924 Traverella, 196, 275, 277,284, 308, 313,337 Traverella primanus, 270 Trechus, 873

Treopalpus llthotarinus, 890 Trepohates, 535, 536, 537, 555 Trepobates hecki, 536 Trepobatinae, 535, 555 Triacanthagyna, 363, 368, 398 Triacanthagyna trifida, 369, 371 Triacanthella, 249

Triaenodes. 131,188, 587, 608,631,633, 697, 743

Triaenodesfrontalis, 88 Triaenodes injustus, 88 Triaenodes marginatus, 665 Triaenodes tarda, 631, 632 Triaenodes tardus, 699 Tribelos, 76, 1152,1156, 1222,1271 Trichoceridae, 1023, 1028 Trichochilus, 1162, 1174, 1264 Trichoclinocera, 946,999 Trichocorixa, 530, 531, 561 Trichocorixa reticulata, 86, 531 Trichogramma, 910, 912, 913, 921 Trichogrammatidae, 170, 910, 914, 915,

Triogma, 1057 Triogma exsculpta, 1061 Triogma trisulcata, 1032 Trissopelopia, 1133, 1137, 1185, 1228, 1249 Triznaka, 453,482,520 Triznaka pintada, 456 Triznaka signata, 86,486 Trochopus, 527, 547 Tronamyia lindsleyi, 1012 Tropidopolinae, 411 Tropisternus, 213, 793, 836,837, 839, 840,

Veliinae, 544 Vesicephalus, 256, 257 Vespoidea, 909, 912, 924 Vial rearings, 40 Viehoperla, 439,479, 508 Viehoperla ada, 443,484 Virgatanytarsus, 1145, 1146, 1274 Visoka, 446, 472 Visoka cataractae, 445,446, 472, 511

842, 887 Tropisternus ellipticus, 89 Trypetoptera canadensis, 1021

w

Tsalia, 271

Tsalia berneri, 288, 290, 308, 314, 333 Tullbergia, 248 Tullberginae, 247 Tvetenia, 1170, 1209, 1210, 1264 Tvetenia bavarica, 1167 Tvetenia discoloripes, 1167 Twinnia, 1099, 1101, 1106, 1109, 1118 Twinnia tibblesi, 1102, 1111 Typha, 118, 767

1250

Trichotendipes, 1221 Trichothaumalea, 992 Trichothaumalea elakalensis, 965

Trichotipula (Tiptda subgenera), 1024, 1038, 1039

V

Tricoryhyphes, 270, 271 Tricorythidae, 75, 81, 270, 333 Tricorythodes, 53, 75, 83, 196, 265, 270, 271,273,289, 304, 305,315, 334 Tricorythodes allectus, 83 Tricorythodes atratus, 83 Tricorythodes minutus, 83 Tricyphona, 1023, 1024, 1042, 1068 Tricyphona immaculata, 1044 Tricyphona inconstans, 1044, 1058 Tridactylidae, 169,205,411, 413, 415, 416,421,427 Tridactylinae, 427 Trigonidiinae, 417,428 Trimerina madizans, 1007

Trimerinoides adfinis, 1007

Vitis, 767

Vivacricotopus, 1193, 1228, 1264

Water column sampler, 21, 22, 23 Waynokiops. 268, 302 Waynokiops dentatogriphus, 268, 276, 326 Weberacantha, 253 Whitneyomyia, 944 Wiedemannia, 946,999

Wilding sampler, 18, 19, 21, 23, 38 Willemia, 249, 251 Williamsonia, 379, 380,400 Willowsia, 250, 254 Wirthiella, 1220, 1223 Wormaldia, 216,646,647, 712, 730

Typhlogastrura, 249 Typopsilopa, 1014

Wormaldia moesta, 87

u

Wyeomyia, 1073, 1079,1082,1083, 1084, 1087,1089,1096

Wormaldia occidea, 714

920-21

Trichogrammatidae, 913 Trichomalopsis, 916, 920 Trichomycetes, 1120 Trichopria, 910, 912, 915,922 Trichopria columbiana, 910 Trichoptera, 2, 13,28, 32, 37, 49, 52, 54, 55,67,68,72, 76,77, 80,81,82, 86-88,92, 96, 102^, 112, 113, 115, 119, 120, 125, 126, 137, 145, 150, 166, 167-68, 174, 176, 183, 184, 186, 187, 189, 190, 192, 216-20, 232,233, 234, 236, 238, 239,240, 241, 242, 585-764, 791 Trichotanypus, 1177, 1181, 1183, 1184,

1479

Uenoidae,68, 88, 94, 126, 130, 133, 167, 219, 586, 589, 590, 595,607, 655, 660, 663,671,675,680, 681,722, 723, 764 Ula, 1027

Ulomorpha, 1026, 1027, 1046, 1066 Ulomorpha pilosella, 965 Ulomorpha vanduzeei, 1034, 1049 Vndulambia, 783 Unniella, 1170, 1173, 1194, 1264 Uranotaenia, 1071, 1073, 1078, 1079, 1083, 1084, 1087, 1088, 1096

Urolepis, 912,920 Urolepis rufipes, 912 Usingeriessa, 766, 769, 770, 771, 783 Usingeriessa onyxalis, 767 Usingerina, 558 Utacapnia, 448,466,470, 472, 513 Utacapnia lemoniana, 440,449,471, 473 WtipeWfl, 453,482, 519 Utaperla gaspesiana, 453 Utaperla sopladora, 454,486 Uvarus, 817, 823, 881 Uzelia, 252

Xenelmis, 860, 866 Xenelmis sandersoni, 899

Xenochironomus, 1152, 1225, 1227, 1271 Xenochironomus xenolahis, 1155

Xenopelopia, 1134,1184, 1185, 1249 Xenylla, 249 Xenyllodes, 249 Xestochironomus, 1154, 1156,1225, 1271 Xiphocentron, 217,604, 655, 669,678, 722 Xiphocentron messapus, 666, 723, 733 Xiphocentronidae, 168, 217, 586, 603, 604,655, 666,669,672,678, 682, 722, 723, 733

Xylena nupera, 786 Xylotopus, 925, 1124, 1166, 1170, 1172, 1196, 1199, 1264

Xylotopus par, 91

Yamatotipula (Tipula subgenera), 965, 1023, 1024, 1038, 1040, 1056, 1058 Ylodes, 88

Vacupernius, 289, 305 Vacupernius packeri, 290, 315, 334 Varipes, 282, 293, 302 Varipes lasiobrachius, 326 Vatellini, 823 Vatellus, 816, 823 Veliidae, 170, 207, 522, 523, 527, 544, 547, 548,552

Yoraperla, 439,482 Yoroperla brevis, 440, 443,444 Yoroperla nigrisoma, 484 Yphria, 605 Yphria californica, 646,648,650,662, 715, 716, 762 Yugus, 459,497,499, 502, 518 Yugus bulbosus, 461, 500 Yugus kondratieffi, 465

1480

Index

Zaitzevia, 860, 861, 862, 863, 899 Zalutschia, 1161, 1171, 1192, 1193, 1195, 1196, 1264 Zalutschia zalutschicoia, 1171

Zapada, 137, 202, 204,442, 472, 511 Zapada cinctipes, 53, 85 Zapada columhiana, 85 Zapada haysi, 440,445 Zapada oregonensis, 465, 475

ry

Zealeuctra claasseni, 441

Zetterstedt, 661, 711 Zeuximyia, 943 Zoniagrion exclamationis, 358, 362, 364, 409 Zoraena, 379, 399 Zoraena diastatops, 388 Zumatrichia notosa, 625,626,696, 697, 738 Zygentoma, 177

Zealeuctra warreni, 480,483

Zygoptera, 53, 55, 81, 84, 342, 343, 344,

Zavrelia, 1139, 1142, 1216, 1218, 1229, 1274 Zaweliella, 1154, 1156, 1157, 1222, 1223, 1271 Zavrelimyia, 1130, 1132, 1133, 1135, 1137, 1139, 1184, 1185, 1186,1249 Zealeuctra. 85,430,439, 448, 476, 479, 512

Zealeutra claassenia, 450

345, 346, 347, 348, 349, 351, 355-63,

Zeros, 1015

405-9

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.Fifth Edition

An Introduction to the

AQUATIC INSECTS of NORTH AMERICA

vS>

S^SaSi

^

Edited by

R.W. Merrltt ■ K.W. Cummins ■ M.B. Berg

Dr. Richard W. Merritt, Ph.D., is an aquatic entomologist and ecologist specializing in the feeding ecoiogy

and biology of aquatic insects, especially the Diptera. He is currently Michigan State University Distinguished Professor Emeritus and previous Chair of the Department of Entomoiogy. Dr. Merritt has received the Award of Exceilence from the North American Benthological Society (NABS, now the Society for Freshwater

Science) and was recently made an inaugural Fellow of this Society. While at MSU, Dr. Merritt conducted lab and field research, directed graduate students, published research papers and book chapters, and taught courses in Aquatic Insects, Stream Biomonitoring, and Forensic Entomology. He is a Diplomate of the American Board of Forensic Entomology, a Fellow in the American Academy of Forensic Sciences and received the Life Time Achievement Award from this Academy.

Dr. Kenneth W.Cummins,Ph.D., is an Adjunct Professor of Humboldt State and Michigan State Universities.

He held faculty positions at Northwestern, Pittsburgh, Michigan State, Oregon State, and Maryland Universities and was the Distinguished Scientist at the South Fiorida Water Management District. For over 50

years. Cummins taught ecology and taxonomic classes, trained graduate students, and conducted funded research. He has an extensive publication record that includes three citation ciassics and the wideiy cited

River Continuum paper in 1980 Can, J. Fish. Aquat. Sci.). He is the recipient of the ASLO Martin Award and the NABS Award of Excellence in Benthic Science.

Dr. Martin B. Berg, Ph.D., is an aquatic entomoiogist and ecologist speciaiizing in the ecology of Chironomidae and assessing energy fiow in aquatic systems. He is a Professor of Biology at Loyola University

Chicago, where he teaches courses in aquatic insects, stream ecoiogy, general ecology, and biostatistics and experimental design. Dr. Berg received his B.S. and M.S. from Indiana University of Pennsylvania and his Ph.D.from the University of Notre Dame. He has published numerous book chapters and research papers in scientific journais and has been an active member of the Society for Freshwater Science (formeriy the North American Benthological Society) since 1983.

Kendall Hunt pub l i sh i ng

company