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An Introduction to the Aquatic Insects of North America
5th Editions Entomology Richard W. Merritt Digital
Instant Download
Author(s): Richard W. Merritt, K. W. Cummins, M. B. Berg
ISBN(s): 9781524968540, 1524968544
Edition: 5
File Details: PDF, 208.00 MB
Year: 2008
Language: english
 yt«U)1

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ji'
 An Introduction to the
AQUATIC INSECTS of
NORTH AMERICA
r\

ry
 • • Jf

■ V, ?€>--•

-5i>-

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.
r\
 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

vii
viii 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 ix

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
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 photographs of the majority of aquatic insect families
revised 5th edition of An Introduction to the Aquatic as a supplement to identification by two of the best
Insects of North America. Ken, Marty, and I are insect photographers in North America (Courtney
excited about the new and revised additions to this and Marshall). Important changes have been made,
new edition. When first published in 1978, the book including revision and expansion of keys, along with
had 22 authors, keys only to the family level, and new figures added to the taxonomic chapters. Figures
1,712 references. This 5th edition has 45 authors, have also been added to the General Classification and
expanded generic level keys,and over 7,000 references. Key to Orders chapter so that students do not need to
Sadly, nine authors have passed since the last edition refer to figures in other parts of the book when keying
and are listed in the Dedication. We have added 18 out aquatic insects to Order level. There have been
new authors in this 5th edition. As with previous particularly significant revisions to the chapters on
editions, this new one is intended to serve as a standard Ephemeroptera, Plecoptera, Trichoptera, Coleoptera,
guide to the aquatic and semiaquatic insects of North Diptera,and Tipuloidea.The Trichoptera and Diptera
America,including keys to the immatures and in most chapters have each been combined into one chapter
cases adults, with pupal keys to the Trichoptera, for each order. Larvae, pupae,and adults together are
Diptera, Culicidae, Simuliidae, and Chironomidae. treated as one chapter for each order. As with any
There have been substantial additions and expanded comprehensive treatment on aquatic insects, coverage
coverage to some of the introductory chapters, ofthe literature can only be partial because it continues
especially the Bioassessment, Respiration, Habitat, to grow exponentially. This is a true measure of the
and Life History chapters. A separate chapter on popularity of the subject matter. We added to the
Ecological Tables of Aquatic Insects, the hallmark of references submitted by the authors by surveying
the first four editions, has been added to summarize other literature sources. As before, we strongly
and elaborate on the ecological information for each encourage users of the book to continually update
taxon,as well to update,revise and expand the content. material in their own areas of interest. Also, we hope
In addition,a new chapter on A Photographic Overview this new edition will be of even greater use to both
ofAquatic Insects of North America has been added to professional and lay groups interested in aquatic
the book. This chapter includes outstanding color insects.

XI
ry
 ACKNOWLEDGMENTS
The editors would like to thank all the contribu Acknowledgments, thanks, and credits by con
tors, both old and new, for their cooperation during tributors and the editors for specific chapters are as
the production of the 5th edition of this book. This follows:
new edition would have not been possible without
their help and expertise. We also would like to thank Chapter 7: Biomonitoring
our publisher, Kendall/Hunt Publishing Company, We thank Marcus Beck, Dave Buchwalter, Joyce
Dubuque, Iowa for their cooperation and patience Chou, Matthew Cover, Cody Fees, David Gillett,
with this endeavor. They have been our publisher Charles Hawkins, Ryan King, Katrina Krievins,
since the first edition in 1978 and we have developed Jason May, Alvina Mehinto, Patina Mendez, Peter
a good working relationship over the years to help Ode, Alison O'Dowd, Ashley Park, Andrew Rehn,
make this book a success. At MSU, we would like to Eric Stein, Stephanie Strachan, Susanna Theroux,
thank Dr. Bill Ravlin, Chairperson of Entomology, and Rebecca Willison.
MSU,for his continued support of this project over
the past 2.5 years. We would especially like to thank Chapter 10: Aquatic Insects of N. A.: A Photographic
Mr. Scooter Nowak for assistance with the computer Overview

program to merge old and new references and all We are grateful to numerous colleagues for shar
other issues dealing with computer programming in ing their knowledge and advice on certain families
this edition. of Ephemeroptera (B.C. Kondratieff, D. Lemkuhl,
I(RWM)would like to thank my wife Pam for and J. Webb) and Trichoptera (N.H. Anderson,
her continued patience during another edition of this D.E.Bowles,J. Giersch,L.Myers,and R.W.Wisseman).
book and my close friends and colleagues, Gary Without their generous assistance, we would have
Lamberti, Eric Benbow, Marty Berg, and Johnny been unable to locate and photograph many fami
Wallace who continually harassed me along the way lies included in this chapter. Several individuals
with their texting! graciously helped with identification of our images:
I (KWC) Over 60 years as an aquatic ecologist Ephemeroptera (S. Burian), Odonata (Ken Tennes-
there have been so many who have helped, collabo sen), Plecoptera (B.C. Kondratieff and R.E. DeWalt),
rated and inspired me along the way. The list is too Coleoptera(D.R. Maddison and A. Short), Trichop
long to include here, but I would single out Noel tera (J.C. Morse, A.K. Rasmussen, D. Ruiter, and
Hynes, the two other editors of this book. Rich and R.W. Wisseman), and selected Diptera (A. Fasbender
Marty, and Margaret Wilzbach, Clyde Eriksen, Bill and B.J. Sinclair). Jon K. Gelhaus kindly provided
Coffman, and Bob Peterson. specimens of larval Phalacrocera (Cylindrotomidae)
I (MBB), first and foremost, thank my wife Pat for us to photograph. We thank the following indi
for all of her support and for enduring yet another viduals for allowing use of their images: C.R. Nelson
edition of the book. I also thank my children, Juliana (Ephemeroptera: Euthyplociidae), M. Garrison
and Ethan, for their understanding, patience, and for (Odonata: Corduliidae), Ken Tennessen (Odonata:
agreeing not to ask "Is it done yet?" Finally, I thank Lestidae), and U.G. Neiss(Odonata: Platystictidae),
Rich and Ken for the opportunity to join them on this and J.C. (Skip) Hodges, Jr. (Trichoptera: Dipseu-
and the previous edition, and Gary Lamberti, my dopsidae, Ecnomidae, and Xiphocentronidae). Field
good friend and valued colleague for his sage advice work associated with this chapter was supported
over the years. in part by a National Science Foundation grant

Xlll
xiv Acknowledgments

(DEB-0933218)to G.W.Courtney and the National Figs. 13.244-13.245 (Allen and Edmunds 1963a);
Institute of Food and Agriculture, Project No's. Fig. 13.246 (Allen and Edmunds 1963b); Fig. 13.72
6693 and 5473. (Bednarik and Edmunds 1980); and Fig. 13.76(Traver
and Edmunds 1967). Figures 13.93-13.105 and
Chapter 13: Ephemeroptera Figs. 13.197-13.198, 13.202-13.207 are taken from
The keys to nymphs and adults in this chapter Wiersema and McCafferty (2000); Figs. 13.265-13.266,
were originally developed based on those presented Figs. 13.268, 13.270 are taken from McCafferty
by Edmunds et al. (1976) and first appeared in the and Provonsha (1985), courtesy of the authors; Fig
second edition of this book. Since then the keys have 13.259-13.260 are taken from McCafferty and
changed considerably with each edition ofthe book as Lugo-Ortiz (1998); Fig 13.123 is taken from Lugo-
did our knowledge of the taxonomy of North Ameri Ortiz and McCafferty (1998c); Figs. 13.179-13.180 are
can mayflies. 1 am indebted to George Edmunds, taken from Lehmkuhl (1976); Figs. 13.177-13.178,
Jr. and Robert Waltz for all of their great work on pre 13.152 are taken from Kluge (2004); Figs. 13.77 and
vious editions of this chapter. The keys in this edition 13.205 are taken from Flowers and Dominguez(1992).
also have benefited from the generous contributions by
bench taxonomists and aquatic biologists that use keys Chapter 14: Odonata
as part of their jobs and discovered problems or John C. Abbott (University of Alabama, Tusca-
observed variations in characters that created stum loosa, Alabama)for providing a wing scan of Lepto-
bling blocks in parts ofthe previous keys. I also want to basis and proofreading the draft for the 4th edition;
thank all my mayfly colleagues that were willing to Maria C. Garrison (McHenry County College,
share their ideas and time discussing some of the per Illinois) for proofreading and critiquing the entire
sistent problems still plaguing mayfly taxonomy rele chapter for the 5th edition, as her comments and
vant to the keys presented here. questions resulted in many corrections and improve
In this edition there are several new figures, but ments in the draft manuscript.
most of the figures from the previous edition are
retained. I grant permission to use figures from my Chapter 16: Plecoptera
publications; Fig. 13.229 (Burian 2001) and Fig. 13.234 We graciously acknowledge that the following
(Burian 1995). I am grateful to the University of Min keys to adult and nymphal stoneflies are based upon
nesota Press for the continued use of figures from the groundbreaking work of Harper and Stewart
Edmunds et al. (1976). I greatly appreciate the permis (1984), Stewart and Harper(1996), Stewart and Stark
sion of Dr. R.D. Waltz to continue to use illustrations (2002), and Stewart and Stark (2008). Their contribu
from his publications: Fig. 13.255 (Waltz and tions were influenced by regional keys to stoneflies by
McCafferty 1989); Fig. 13.263(Waltz and McCafferty Jewett (1956) and Hitchcock (1974). We have incor
1987a), Fig. 13.108(Waltz eta/. 1985); Fig. 13.49(Waltz porated the new stonefly genera proposed since the
2002); Fig. 13.124-13.125(Waltz and McCafferty 1985); last edition and made other modifications suggested
Figs. 13.116 and 13.261 (Waltz and McCafferty 1999). by colleagues and students.
The late Dr. R.K. Allen allowed us to republish The update of this chapter has built a solid foun
Figures 13.79 (Allen 1974), 13.90, 13.92 (Allen 1973), dation for its future improvement. Repeated photo
13.91 (Allen 1976), and 13.246 (Allen 1965). Figures copy reproduction of previous nymphal illustrations
13.80, 13.223-13.227, 13.230, and 13.267 are from degraded image quality. Therefore, we replaced most
Burks(1953)and are published with permission of the of the nymphal illustrations of Stewart and Stark
Illinois Natural History Survey. Figure 13.44 is from (2008) with those scanned from Stewart and Stark
Needham et al. (1935) and is published courtesy of (2002). Adult images from the 4th edition were near
Cornell University Press. Figure 13.264 is from Prov- pristine, requiring only scanning with minor editing.
onsha and McCafferty(1982),courtesy ofthe authors. All images are now saved as tiffs and deposited for
Figures 13.119, 13.269, and 13.271 are from Davis safe keeping. Many new figures were added from var
(1987). Figures 13.120, 13.122 are from Peters (1971) ious sources and we extensively edited the keys to
with permission ofthe author. Figures 13.66-13.68 are improve their usefulness. New references have been
from Bednarik and McCafferty (1979), courtesy of added and some older ones retired.
the Canadian Bulletin of Fisheries and Aquatic
Sciences. The late Dr. G.F. Edmunds, Jr. provided Chapter 19: Trichoptera
Figs. 13.83-13.84, 13.240 (Allen and Edmunds Larval and pupal keys are based on the work of
(1962); Fig. 13.243 (Allen and Edmunds 1965); Wiggins and Currie(2008). Assistancefrom D.E.Ruiter
 XV Acknowledgments

is gratefully acknowledged and we would like to thank J.K. Moulton (Dixidae), and B.J. Sinclair (Empi-
James C.(Skip) Hodges, Jr. for allowing us to use his doidea). 1 am especially grateful to Brad Sinclair for
ry excellent caddisfly case photographs. his revision of all couplets pertaining to empidoid
flies. I would also like to thank the scientists at
Chapter 21: Coleoptera Rhithron Associates, Inc. for beta-testing the keys.
Rob Roughley contributed extensively as the This work was supported in part by the National
senior author of the 4th edition. We thank Stephen Science Foundation (grants DEB-0933218 and
Baca (Noteridae), Grey Gustafson (Gyrinidae), Crys EF-1115156) and the National Institute of Food and
tal Maier (Lutrochidae), and Phil Perkins (Hydraeni- Agriculture, Project No's. 6693 and 5473.
dae)for reviewing various portions ofthe text and key
for the 5th edition. Chapter 24: Tipuloidea
We acknowledge a great debt to the late Prof. George
Chapter 23: Diptera Byers, University of Kansas. His extensive studies ofthe
I wish to acknowledge numerous colleagues for North American Tipuloidea, spanning more than 50
sharing their knowledge and advice on selected Dip years, and his development of the first comprehensive
tera, especially A. Borkent (Ceratopogonidae), keys to aquatic crane fly larvae, provide much of the
G.R. Curler (Psychodidae), J.K. Gelhaus (Tipuloidea), basis for the present keys(Gelhaus et al. 2018).
 LIST OF CONTRIBUTORS
P. H.ADLER Department of Plant and Environmental R. W. HOLZENTHAL Department of Entomology,
Sciences, Clemson University, Clemson, SC 29634 Hodson Hall, 1980 Folwell Ave., University of
D. P. BATZER Department of Entomology, Univer Minnesota, St. Paul, MN 55108
sity of Georgia, Athens, GA 30602 A. D. HURYN Department of Biology, University of
M. E. BENBOW Department of Entomology and Alabama, 2107 Bevill Building, Box 870206,
Department of Osteopathic Medical Specialties, Tuscaloosa, AL 35487
Michigan State University, 243 Nat. Sci. Bldg. J. K. JACKSON Stroud Water Research Center, 970
288 Farm Lane, East Lansing, MI 48824 Spencer Road, Avondale, PA 19311
A. M. R. BENNETT Canadian National Collection B. C. KONDRATIEFF Colorado State University,
of Insects, Arachnids and Nematodes, Agricul Department of Bioagricultural Sciences and Pest
ture and Agri-Food Canada,960 Carling Avenue Management, 1177 Campus Delivery, Fort Collins,
Ottawa, Ontario Canada KIA 0C6 CO 80523
M.B.BERG Department of Biology,Loyola University G. A. LAMBERTI Department of Biological Sci
Chicago, 1032 W. Sheridan Rd., Chicago, II60660 ences, University of Notre Dame, Notre Dame,
D. E. BOWLES Department of Biology, Missouri IN 46556-0369
State University, Springfield, MO 65897 S. A. MARSHALL University of Guelph Insect Col
D. B. BUCHWALTER Department of Environmen lection and Insect Systematics Laboratory,
tal & Molecular Toxicology, Campus Box 7633, School of Environmental Sciences (Bovey), 1216
NC State University, Raleigh, NC 27695-7633 Edmund C. Bovey Building, University of
S. K. BURIAN Department of Biology, Southern Guelph, Guelph, ON,Canada NIG 2W1
Connecticut State University, 501 Crescent R. D. MAZOR Southern California Coastal Water
St., New Haven, CT 06515 Research Project, 3535 Harbor Blvd, Suite 110,
A. CONTRERAS-RAMOS Instituto de Biologia, Costa Mesa, CA 92626
UNAM,Depto. de Zoologia,Apdo Postal 70-153, R. W. MERRITT 1005 Cormorant Terrace, The
04510 Ciudad de Mexico, Mexico Villages, EE 32162
G. W.COURTNEY Department of Entomology,Iowa B. W. MERRITT Via dei Gilardi 17,6926 Montagnola,
State University, 401 Science II, Ames,IA 50011 Switzerland
K W. CUMMINS California Cooperative Fisheries J. C. MORSE Department of Plant and Environ
Research Unit,Humboldt State University, Arcata, mental Sciences, Clemson University, Clemson,
CA 95521 SC 29634
D C. CURRIE Department of Natural History, S. NOWAK School of Informatics, Computing and
Royal Ontario Museum, 100 Queen's Park Cyber Systems, Northern Arizona University,
Toronto, ON, Canada M5S 2C6 Building 90, 1295 S. Knoles Dr., Flagstaff, AZ
R.E. DEWALT University of Illinois Prairie Research 86011
Institute, Illinois Natural History Survey, 1816 S. G. E. PARSONS Department of Entomology,
Oak St., Champaign, IE 61820 Michigan State University, 243 Nat. Sci. Bldg., 288
L. C. FERRINGTON,JR. Department of Entomol Farm Lane, East Lansing, MI 48824
ogy, Hodson Hall, 1980 Folwell Avenue, Univer V. PODENIENE Institute of Biology, Life Sciences
sity of Minnesota, St. Paul, MN 55108 Center, Vilnius University, Sauletekio str. 7,
J. K. GELHAUS Department of Entomology, The LT-10257 Vilnius, Lithuania
Academy of Natural Sciences of Drexel University, D. A.POLHEMUS Department of Natural Sciences,
1900 Benjamin Franklin Parkway, Philadelphia, Bishop Museum, 1525 Bernice St., Honolulu, HI,
PA 19103 96817

xvii
xviii List of Contributors
o

A.K.RASMUSSEN Center for Water Resources, 113 M.A.SOLIS SEL,USDA,Smithsonian Institution, n

S. Perry-Paige Building, 1740 S. Martin Luther P.O. Box 37012, National Museum Natural
King Jr. Blvd., Florida A&M University, History, E-517, MRC 168, Washington, DC
Tallahassee, FL 32307 20013-7012
J. P. RFCFVFUR Department of Entomology, H. SONG Department of Entomology, Texas A&M
Michigan State University, 243 Nat. Sci. Bldg., University, College Station, TX 77843-2475
288 Farm Lane, Fast Lansing, MI 48824 K. J. TENNESSEN P.O. Box 585, Wautoma, WI 54982
V. H. RFSH University of California, FSPM J. B. WALLACE Department of Entomology and
Department, Organisms & Environment Divi Odum School of Ecology, University of Georgia,
sion, Berkeley, CA 94720 Athens, GA 30602
D. R. ROBERTSON Integrative Research Center, J. R. WALLACE Department of Biology, Millersville
Field Museum of Natural History, 1400 S. Lake University, Millersville, PA 17551
Shore Drive, Chicago,IL 60605 D.S. WHITE Hancock Biological Station, Murray State
D. M. ROSENBERG 280 Waverley St., Winnipeg, University, 561 Emma Drive, Murray, KY 42071
MB,Canada R3M 3L3 W. C. E. P. VERBERK Department of Animal
A. E. Z. SHORT Department of Ecology and Evolu Ecology and Physiology, Radboud University,
tionary Biology, University of Kansas, 6002 Nijmegen, The Netherlands
Haworth Hall, Lawrence, KS 66045 K. W. WILL University of California, ESPM
R. J. SNIDER Department of Integrative Biology, Department, Organisms & Environment Division,
Michigan State University, 203 Nat. Sci. Bldg., Berkeley, CA 94720
288 Farm Lane, East Lansing, ML 48824
 ffrls. -: / V-

INTRODUCTION
Richard W. Merritt Martin B. Berg
Michigan State University, East Lansing Loyola University Chicago, Illinois
Kenneth W. Cummins
Humboldt State University, Arcata

The emphasis on aquatic insect studies, which has In addition, fly anglers have enthusiastically sought
expanded exponentially in the last five decades, has more knowledge about aquatic insects, both as fish
been largely ecological. This interest in aquatic insects foods to be imitated and as interesting cohabitants with
has grown from early limnological roots (e.g., Forbes their quarry (e.g., Swisher and Richards, 1971, 1991,
1887) and sport fishery-related investigations of the 2018; Schweibert 1973; Caucci and Nastasi 1975,2004;
'30s and '40s (e.g., Needham 1934), to the use of Borger 1980, 1995; LaFontaine 1981; Whitlock 1982,
aquatic insects as indicators of water quality during 2014; Hafele and Roederer 1995; Knopp and Cormier
the '50s and '60s (e.g., Kuehne 1962; Bartsch and 1997; Ames 2005; Fauceglia 2005; Weamer 2017).
Ingram 1966; Wilhm and Dorris 1968; Warren 1971; More recently, aquatic entomology and its applications
Cairns and Pratt 1993). In the '70s and '80s, aquatic have experienced many improved and/or new methods,
insects became the dominant forms used in freshwater methodologies, and coupled technologies (DeWalt
investigations of basic ecological questions (e.g., 2010; Hauer and Lamberti 2017; Lamberti and
Barnes and Minshall 1983). During the past 10 years Hauer 2017).
(2008-2018), emerging studies have dealt with inva Initially, the primaryjustification for this book was
sions of alien invertebrate species, and forecasting that the systematics ofaquatic insects had lagged behind
responses of benthic insect community structure and the needs ofaquatic ecologists and water managers,and
function to anthropogenic climate change (Poff et al. the inquisitiveness of anglers. This is still true, but to a
2010; Strayer 2010; Ricciardi 2015; Fenoglio et al. lesser extent, although the sophistication of scientists,
2016). Ecological applications of observed habitat managers, and anglers perpetuates the need for ever
affinities and traits associated with the physical tem better taxonomic and ecological treatments.
plate ofstreams will become increasingly important in Aquatic insects also are of concern to those
predicting how aquatic insects respond to changing involved in teaching (e.g., Resh and Rosenberg 1979;
hydroclimate and flow regime (Pyne and Poff 2017; Li and Barbour 2011; Hauer and Lamberti 2017;
Flerbst et al. 2018). In addition, DNA barcoding also Lamberti and Hauer 2017; Merritteta/. 2017; Gushing
will likely become more widely used in the identifica 2016), and in outdoor recreation activities because
tion of aquatic insects (e.g., DeWalt 2010). Our certain groups (e.g., mosquitoes, black flies, horse
expanding knowledge of biodiversity and the role that flies) are frequently pests of humans and other ani
different aquatic insects play in water quality assess mals in water-based environments (Kim and Merritt
ment is the only way to sustainably manage ecosys 1987; Malmqvist et al. 2004; Lemelin 2013). Identifi
tems in an ever changing global environment(Foottit cation is the first step toward a basic understanding of
and Adler 2009; Dijkstra et al. 2014). the biology and ecology of aquatic insects that even
The work on aquatic insects has embraced most tually allows for the development of proper manage
major areas of ecological inquiry including population ment strategies. The amateur naturalist and primary,
dynamics, predator-prey interactions, physiological secondary, and post-secondary school educators also
and trophic ecology,competition(Resh and Rosenberg require basic identification as an important initial
1984; Allan 1995), and management applications of step in familiarization. Thus, for all concerned, iden
this basic research (Wright eta/. 1991; Rosenberg and tification and basic ecological and life history infor
Resh 1993a; Dodds 2002; Benke and Gushing 2005). mation is important for categorizing the aquatic

1
 Chapter 1 Introduction

insects collected. This 5th edition continues to offer terrestrial species are encountered with varying fre
information on functional adaptations of aquatic quency. A specimen not fitting the keys in this edition
insects that allows an additional tool for categorizing probably belongs to a terrestrial taxon and generally
aquatic insects (e.g., Chapter 6). can be identified using Triplehom and Johnson (2005).
A number of well-known general works(Usinger An annotated list of general references to works
1956a; Edmondson 1959; Klots 1966; Pennak 2001) dealing with aquatic insect taxonomy and ecology
and specific studies(Ross 1944; Burks 1953)are taxo- is given in Table 1. As indicated above, many of the
nomically and ecologically, at least for the most part, taxonomic works are outdated; however,they include
out-of-date. More recent comprehensive treatments a great deal of useful biological information on
of the Odonata (Westfall and May 1996; Needham et the groups. More specific references are given at the
al. 2000), Plecoptera (Stewart and Stark 2002), and end of the appropriate order (or family) chapter.
Trichoptera(Wiggins 1996)currently are available, as Various combinations of taxa can be categorized so
are some general works (e.g., Stehr 1987,1991; Thorp as to permit ecological questions to be addressed at
and Covich 2010; Thorp and Rogers 2016). However, the functional level. For example, some groups are
at the present time only this 5th edition gathers based on morpho-behavioral adaptations for food
together comprehensive, updated, generic treatments gathering, habitat selection, or habits of attachment,
of immature and adult stages of aquatic and semi- concealment, and movement(Chapter 6, and ecolog
aquatic insects of North America. This edition is ical tables at the end of each taxonomic chapter).
intended, as were previous editions, to serve as a stan Different levels of taxonomic identification are
dard reference on the biology and ecology of aquatic required to functionally classify aquatic insects. For
insects with updated keys to separate life stages of all example, the ordinal level may be sufficient to define
major taxonomic groupings. To this aim, we have functional trophic relations for the Odonata, but
provided additional color photographs of most all even the generic level may be insufficient in some of
families of immature aquatic insects to assist the stu the Chironomidae (Diptera). Fcologists also have
dent and professional with correct identifications. resorted to "habitat taxonomy" of a single aquatic
The taxonomic coverage is coupled with summa system (e.g., Coffman et al. 1971) or regionalized
ries of related information on aquatic insect phylog- keys (e.g., Brigham et al. 1982; Peckarsky et al. 1990;
eny,ecology,water quality,bioassessment,respiration, Ward and Kondratieff 1992; Bouchard 2004; Hudson
sampling, rearing, life history, and behavior. Generic et al. 2012; Morse et al. 2017), where the fauna of a
keys to immatures and adults are provided for all but given system or region is studied for an extended
a few groups of Diptera. Further, pupal keys are now period in sufficient detail to permit such system-spe
provided for most holometabolous orders. All the cific keys to be written. Although this allows for sig
keys have been revised, and very significant revisions nificant simplification in such keys by excluding taxa
have been made to many groups, such as the Ephem- from other systems or regions, changes in species
eroptera, Odonata, Plecoptera, Trichoptera, aquatic composition, the key element in disturbance or intro
Coleoptera, and Diptera. Because of the size of the duction of exotics can be masked by the restricted
order Diptera,separate chapters have been devoted to nature of this approach. This means that such keys
individual families or superfamilies (i.e., Chironomi- must be used cautiously and verified continually
dae, Simuliidae, Culicidae, Tipuloidea). against the full range of taxonomic information
The distinction between aquatic or semi-aquatic available.
and terrestrial insects is arbitrary. In this 5th edition, The particular emphasis on ecology and field
those orders and families with one or more life stages techniques in all previous four editions of this book
associated with aquatic habitats and frequently encoun has reflected our conviction that the most critical task
tered in collections made from aquatic environments at hand is the integration oftaxonomic and ecological
are covered. This includes the Collembola,Orthoptera, approaches. This integrated approach will permit
and Hymenoptera, even though they are only margin important questions concerning environmental qual
ally associated with aquatic habitats. Because terrestrial ity and management to be addressed. It is our hope
insects frequently become trapped in the surface film that this expanded 5th edition will provide another
of aquatic systems (e.g., Collembola), a wide range of significant step toward this goal.
 Chapter 1 Introduction

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

Taxonomic
Treatments GENERAL COVERAGE

Source Immatures Adults Biology General Comments

Ward and Whipple x Generic Contains much information on biology.

(1918)
Chu(1949) X Family Generalized treatment of immatures.
Peterson (1951) x Family Limited to hoiometabolous groups;
descriptive in nature.
Usinger (1956a) x x^ Primarily generic level, x Considerable information on West Coast
with keys to Calif, species species.
Edmondson (1959) x X* Generic A standard reference on freshwater
invertebrates; some keys outdated.
Eddy and Hodson (1961) x Order Keys to common animals, including water
mites, of the North Central states.
Needham and Needham x X* Generic Keys to many genera; field manual.
(1962)
Klots(1966) X X* Primarily family level, with x^ Field manual; some keys based on ecology
keys to some genera and behavior of group.
Borror and White (1970) x^ X Orders, some keys to families x Comprehensive field manual on insects;
primarily based on examination of insects
in the hand; coior plates.
Swisher and Richards X Generic (mayflies only) x Anglers' guide, primarily to mayflies; color
(1971, 2018) photographs; seasonal data.
Schweibert(1973) None X Extensive treatment of immature aquatic
insects for anglers; color plates; seasonal
and distributional data.

Caucci and Nastasi X None x Anglers' guide to mayflies and stonefiies;

(1975) color photographs; seasonal and
distributional data.
Parrish (1975) Generic Keys only to Southeastern United States;
limited to water quality indicator organisms.
Smith and Carlton x Primarily family level, with x^ Keys only to intertidal insects of the central
(1975) keys to some species California coast.
Tarter (1976) x* Generic x^ Keys only to West Virginia genera and
occasionally species.
Merritt and Cummins x Orders, families, x Chapters on morphology, ecology,
(1978, 1984, 1996), and genera of North American phylogeny, life history, behavior, bio-
Merritt ef a/. (2008) aquatic insects monitoring, and sampling; summary tables
on ecology and N.A. distribution with
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* Families x^ Field guide to aquatic insects.
Borger(1980, 1995) Orders and families x'^ An angler's guide to the major food
organisms of trout and their presentations.
Hilsenhoff (1981) X* 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 Introduction

Table 1 Continued

Taxonomic
Treatments GENERAL COVERAGE

Source Immatures Adults Biology General Comments

LaFontaine (1981) None An angler's guide to the caddisflies; good

biological section on caddis.
McCafferty (1981) Pictorial keys to aquatic A thorough scientific introduction to
insect families aquatic insects for the fly anglers; excellent
illustrations.

Brigham etal.(1982) Families and genera for A thorough treatment of the aquatic
eastern North America; insects and oligochaetes of the Carolines.
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.
Stehr(1987, 1991) Orders and families of A thorough treatment of terrestrial and
terrestrial and aquatic aquatic immatures with notes on
insect immatures relationships. Literature sources provided.
Intended as text for classes on immature
insects. Excellent illustrations.
Arbona(1989) X None Good section on mayfly biology for anglers.
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.
Peckarsky etal.(1990) Families and genera of Good regional treatment of aquatic
immature freshwater macroinvertebrates of northeastern North
macroinvertebrates America.

Pobst(1990) None A streamside guide for anglers to the major

trout-stream insects with excellent color
photos.
Clifford (1991) 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.
Swisher and Richards None A good presentation on the emergence
(1991) of mayflies, caddisflies, and stoneflies for
anglers.
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 Introduction

Table 1 Continued

Taxonomic
Treatments GENERAL COVERAGE

Source Immatures Adults Biology 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
identification.
Gooderham and X* Common families Covers temperate Australia but applicable
Tsyrlin (2002) to temperate N.A.; color photographs of
live specimens.
Voshell (2002) X* Order and families 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 Mostly adult coverage; newly described
Johnson (2005) and aquatics orders and families incorporated.
Widely used book for terrestrial insect
identification.
Izaak Walton League X Orders and some families A handy resource for anglers, students,
of America (2006) and biologists spending time near rivers
and streams. Gives tips for distinguishing
similar species on behavior and their role in
stream ecosystems.
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
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 Introduction

Table 1 Continued

Taxonomic
Treatments GENERAL COVERAGE

Source Immatures Adults Biology General Comments

Tzilkowski and None Identification, habitat and life history

Stauffer (2011) information for fly fishers and fly tiers on
Eastern North American nymphs of several
orders of aquatic insects, along with
imitations.

Thorp and Rogers (2016) Orders and families of Deals with inland water invertebrates
aquatic insects and of the Nearctic, primarily include
differential taxonomic taxonomic keys supplemented by
treatment of other invertebrates an introduction to the group and
sections on limitations to taxonomy of the
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) 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.
Ward and Stanford (1979) Comprehensive treatment on the ecology of stream regulation, with an emphasis on downstream effects
on biotic (especially aquatic insects) and abiotic components.
Lock and Williams(1981) 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
Investigation. Good reviews on several current topics in aquatic ecology. Widely used reference source.
Williams(1987) 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.
Ward (1992) 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.
Williams and Feltmate An introductory text to the study of aquatic insects, with background information on the aquatic insect
(1992, 2017) 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 Introduction

Table 1 Continued

Ecological Treatments GENERAL COVERAGE

Wotton (1994) 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.
Allan (1995) 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.
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) 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 multi-
metric Indices and how they can be Incorporated into environmental policy and management. A practical
bock for students, researchers, and managers.
Gushing and Allan (2001) 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.
Mackle (2001) 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.
Dodds (2002) 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.
Resh and Garde This encyclopedia of Insects contains subject area coverage of many aquatic topics (e.g., aquatic habitats,
(2003, 2009) 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.
Benke and Gushing (2005) 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
often Includes a section on aquatic Invertebrates, their diversity, abundance, and ecology.
Brdnmark and A thorough overview of lake and pond ecology. Discusses the structure and function of lentic
Hansson (2005) ecosystems, emphasizing the importance of abiotic factors and blotic Interactions.
Williams (2005) 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 A detailed description of field and laboratory methods commonly employed in the study of physical,
(2006, 2017), Lamberti chemical, and biological components of stream ecosystem structure. Reflects latest advances in the
and Hauer (2017) 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.
Marshall (2006) 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.
Lancaster and A biological approach to aquatic entomology structured around four sections; distribution patterns and
Downes(2013) environmental gradients, dispersal and movement, population dynamics and persistence, and trophic
relationships.
Thorp and Rogers (2015) 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.
 ^ 4^..V m
'Vf-
y^ f"

S&s'> - 'Sfes*/^ ■l'.^

GENERAL MORPHOLOGY
OF AQUATIC INSECTS
Martin B. Berg Richard W. Merritt
Loyola University Chicago, Illinois Miehi^an State University, East Lansing
Kenneth W. Cummins
Humboldt State University, Areata

OVERVIEW and developmental data (Truman and Riddiford

A stonefly (order Plecoptera, family Pteronarcy- 2002; Chapter 11 in this volume).
idae) serves to illustrate the general external morpho
HEAD
logical features of aquatic insects used in taxonomic
determinations. This prototypical insect exhibits The generalized insect head represents the evolu
basic morphological features in a relatively unmodi tionary fusion of six or seven anterior segments in the
fied or nonspecialized form. However, modifications ancestral Annelida-Arthropoda line (e.g., Snodgrass
of the general morphological plan are found in each 1935; Rempel 1975). Two or three preoral (procephalic)
insect order having aquatic representatives. These segments, or somites, were fused and now bear import
modifications and associated terminology are pre ant sensory structures used by present-day insects to
sented with the introductory material for each group monitor their environment—the compound eyes,
and should be carefully studied before attempting to light-sensitive ocelli (simple eyes), and the antennae
use the keys in the following chapters. (Figs. 2.1 and 2.2). The labrum, which forms the upper
The insect body represents the fusion and modifi lip, is joined at its base to the clypeus, which in turn is
cation of the basic segmentation plan characteristic of fused to the frons, or face. The margins of the clypeus
the Annelida-Arthropoda evolutionary line (e.g., and frons are bounded by the anterior portion of the
Snodgrass 1935; Manton and Anderson 1979). Each Y-shaped epicranial suture (in Fig. 2.1, the line ofjoining
segment of the body can be compared to a box, with between the clypeus and frons, termed a suture [sulcus],
the dorsal (top) portion, the tergum or notum, joined is not externally visible so the structure is referred to as
to the ventral (bottom) portion, the sternum, and to the frontoclypeus [Nelson and Hanson 1971]).
the sides or lateral portions, the pleura, by mem Three postoral (gnathocephalic) segments are
branes. The legs and wings are hinged (articulated) on fused in modern insects to form the posterior portion
the pleura of the mid-body region, the thorax. The of the head and bear the remaining structures of the
body regions, head, thorax, and abdomen, and asso feeding apparatus (Snodgrass 1935). As described
ciated appendages of a stonefly nymph are shown in above, the labrum forms the upper lip and the paired
Figs. 2.1 and 2.2. The life cycle of stoneflies is repre mandibles and maxillae form the mouth region laterally
sentative of those orders characterized by simple (Figs. 2.2 and 2.3). The bottom of the mouth is set by
{incomplete or hemimetabolous by some authors) the labium or lower lip (Figs. 2.2 and 2.3). The maxillae
metamorphosis, consisting of egg, nymph (imma and the labium bear palps (palpi), which are sensory in
ture), and adult stages; more advanced orders exhibit function (Figs. 2.2 and 2.3). The mandibles are used for
complete (holometabolous) metamorphosis, consist chewing or crushing food or may be modified for pierc
ing of egg, larva (immature), pupa, and adult stages. ing (piercing herbivores or predators) or scraping
Distinction between the terms "nymph" for immature (scraping herbivores that graze on attached algae). The
hemimetabolous insects and "larva" for immature maxillae and labium are variously used for tearing and
holometabolous insects is supported by endocrine 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 epicranlal suture
occiput
cervix 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 11

antenna
labrum

mandible

pedicel
maxillary palp scape
compound eye
coxa of foreleg
maxilla
trochonter
'a'''^^^]^^__^pronotum (pronotal shield; tergum)
femur of prothorox
tarsus

tibia
prosternum (sternum) of prothorox
branched gill

mesosternum (sternum) of
mesothorox
forewing pad

metasternum (sternum) of
metathorax
hind wing pad

8th obdominal sternum

paraproct (subanal lobe)

epiproct (supraonol
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.
12 Chapter 2 General Morphology of Aquatic Insects

1
labrum (upper lip)

right mandible
^terminal (distal) hypopharynx
incisor lobe
of teeth

basal
(proximal)
molar lobe of
points of teeth
articulation

maxillary palp (palpus) showing

palpal segments

lacinia

palpifer

cardo
(base of maxilla)

ventral

right maxilla

glossa
labial palp paraglossa
(palpus)
showing
femur palpal 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 Figure 2.4 Foreleg of Pteronarcys sp.(Plecoptera;
of Pteronarcys sp. (Plecoptera; Pteronarcyidae): A. Pteronarcyidae) showing segments.
ventral view of head; B. lablum; C. right maxliia; D. right
mandible; E. hypopharynx; F. labrum.
 Chapter 2 General Morphology of Aquatic Insects 13

in the Hemiptera, adult Lepidoptera, Hymenoptera, types of wing venation are shown in Figs. 2.5 and 2.6.
and Diptera. The hypopharynx or insect "tongue," The prototypical stonefly wings have many branches
located just anterior to the labium,is a small inconspic of the major veins with many crossveins between
uous lobe in some larval forms,but is subject to extreme them. The highly evolved wing of a dipteran Tipulidae
modification in some orders (e.g., Diptera). (Tipula sp.) is characterized by the fusion of veins and
The sides of the head are referred to as genae the loss of branches and crossveins.
(singular, gena; Fig. 2.2)and the top ofthe head as the The general venation pattern (Figs. 2.5 and 2.6)
vertex. Immediately behind the vertex is a large area consists of: a costal vein (C), the anterior marginal
called the occiput(Fig. 2.1). The head is joined to the vein; a subcostal vein (Sc)just behind the costal vein
thorax by a membranous neck region or cervix and often with two branches near the wing tip; a
(Fig. 2.1). If the head is joined to the thorax so that radial vein (R), often the heaviest vein of the wing,
the mouthparts are directed downward (ventrally), which forks near the middle of the wing, with the
the condition is termed hypognathous(e.g., many cad- main part forming the radial sector vein (Rs) that
disfly larvae). Mouthparts directed forward (anteri typically divides into two branches, each of which
orly) are prognathous (e.g., beetle larvae) and those may divide into two or more branches near the wing
directed backward (posteriorly) are opisthognathous margin; a medial vein (M)(the fourth major vein),
(e.g., some true bugs). which has a maximum offour major branches (typi
In aquatic insects that are dorsoventrally flat cally two or three); a cubital vein(Cu), which has two
tened, such as some stoneflies and mayflies, the sen major branches, the anterior of which usually forks
sory structures (eyes, ocelli, and antennae) are dorsal into two branches; and an anal (vannal) vein (A),
and the food-gathering apparatus is ventral. These which has a maximum of three major branches with
modifications allow certain groups to move through considerable secondary branching, particularly in
interstices of coarse sediments and cling to exposed more ancestral forms. Although crossveins are
surfaces in rapidly flowing streams. 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
THORAX subcosta; a radial crossvein (r) between the radius
The midregion of the body, or thorax, bears the and the first branch of the radial sector; a radial-me
jointed legs (Fig. 2.4) and the wings, and is divided dial crossvein (r-m) between the lower first fork of
into three segments (Figs. 2.1, 2.2, 2.5, and 2.6). The the radial sector and the upper first fork of the
prothorax bears the forelegs, the mesothorax the mid- medial vein; and a medial-cubital crossvein (m-cu)
legs and forewings, and the metathorax the hind legs between the lower first fork of the medial and the
and hind wings (if wings are present). upper first fork of the cubital (see Snodgrass 1935;
The jointed legs are five-segmented: the coxa, Daly eta/. 1978; Bonoretal. 1981; and discussion of
trochanter, femur, tibia, and the three- to five-seg taxonomically significant wing veins given in the
mented tarsus, which terminates in one or two tarsal 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 ABDOMEN

larval and adult Hemiptera,and a few larval Trichop- The prototypical insect abdomen is composed of
tera. The forelegs are modified for burrowing in eleven segments, although in most adults fusion ofthe
Ephemeridae(Ephemeroptera), Gomphidae(Odonata), last two makes them difficult to distinguish. In some
and some semiaquatic Orthoptera. immature forms (notably Ephemeroptera and Mega-
Most adult forms of aquatic insects bear two loptera), gills arise from the pleural regions—being
pairs of wings(mesothoracic and metathoracic);some extensions of the tracheal system borne in variously
mayflies and all Diptera have only one pair. The sec shaped plates or filaments (finger-like gills). In the
ond pair of wings (metathoracic) in Diptera is modi stonefly shown in Fig. 2.2, the branched filamentous
fied into balancing organs (halteres. Fig. 2.6) that gills are attached to the sterna of the thorax and the
function somewhat as gyroscopes. Collembola are first two abdominal segments.
wingless (apterous), as are females of certain species The end of the abdomen of hemimetabolous
of Trichoptera and Diptera. insects (i.e., Hemiptera, Orthoptera, Ephemerop
The structures that extend into the wings are tera, Odonata, and Plecoptera) bears the reproduc
termed veins. The form and location of these veins are tive structures (Figs. 2.1, 2.2, and 2.7-2.9). The
used extensively in insect taxonomy. Two extreme terminal segment bears the anus at its apex and the
 .antenna (terminal portion cut away)
pedicel
scape radial vein
medial forewing
ocelli tiumeral subcostal vein
radial sector
costal crossvems
compound eye
pronotum vein

cubital
pronotum vein
branches of
radial sector
\

mesonotum
branches of
medial vein
branches of
anal veins costal vein cubital vein

subcostal vein
radial vein
metanotum-
- radial sector vein
- medial vein
1st abdominal
tergum anterior cubital vein

hind wing

posterior cubital vein

anal cell

3 2 1

anal veins
told lines

anal (vannal) region

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

) ) ) )))) ) ) ) )) 1 ) ) ) ) ) ) i ))) ) ) ) )
 Chapter 2 General Morphology of Aquatic Insects 15

- antenna

.compound eye

subcostal radial radial stigma

costal sector
humeral
crossvein

axillary region

halter
meson otum

metanotum cubital-anal
crossvein medial vein
anal veins cubital vein
1st abdominal tergum

Figure 2.6

median
supraanal lobe
hemitergal anterior cercus
epiproct (supraanal process) posterior lobe / hemitergal (terminal portion
hemitergal lobe cut away)
cercus (terminal portion cut away)
lobe

remnant ot 11th tergum 9th tergum tergum

epiproct
posterior hemitergal lobe (supraanal process)
10th tergum 8th abdominal
tergum
r^ median hemitergal lobe
9th tergum supraanal lobe
anterior hemitergal lobe
paraproct
(subanalprocess)
pleuron toth sternum
8th tergum
9th sternum
Bth abdominal sternum

Figure 2.7 Figure 2.8

epiproct
cercus

(terminal portion cut away)

paraproct (subanai lobe)
11th abdominal sternum
toth sternum

9th abdominal tergum

9th sternum
vaginal protection

8th sternum genital opening (gonopore)

8th abdominal tergum

imcKsrr

Figure 2.9

Figure 2.6 Dorsal view of adult Tipula sp.(Diptera: Figure 2.8 Lateral view of terminal male
Tipulidae) showing head, thorax, basal portion of abdominal segments of Pteronarcys sp.(Plecoptera:
abdomen, forewing and halter. Pteronarcyidae).
Figure 2.7 Dorsal view of terminal male abdominal Figure 2.9 Ventral view of terminal female
segments of Pteronarcys sp.(Plecoptera: abdominal segments of Pteronarcys sp.(Plecoptera:
Pteronarcyidae); terminology after Snodgrass(1935) Pteronarcyidae).
and Nelson and Hanson (1971).
16 Chapter 2 General Morphology of Aquatic Insects

cerci laterally. The dorsal surface is covered by a Specific morphological modifications in each of
triangular or shield-shaped tergal plate,the epiproct, the orders(or families receiving special treatment)are
and the ventral surface bears two lobes, the parap- detailed in the introductory material covering the
rocts. In males, the ninth sternum often bears two respective groups.The modifications usually represent
lateral styli or claspers (harpagones). These acces fusion or specialization of the basic structures
sory structures bound the phallobase and aedeagus discussed above. However, some of the terms used in
that comprise the main reproductive organ, the naming the various segments of the genitalia have
penis or phallus. restricted meanings, and homology with primitive
The terminal segments of adult females, in addi forms is not always possible (Tuxen 1970; Scudder
tion to the dorsal epiproct and lateral paraprocts 1971a). For a more complete treatment of insect
below the cerci, generally consist of three pairs of morphology, the student should consult Snodgrass
lobes or valvae (valves), which form the visible por (1935), DuForte (1959), Matsuda (1965, 1970, 1976),
tion of the ovipositor and arise from the eighth and Chapman (2013), Resh and Garde(2009), and Gullan
ninth sterna. The bases of the valvae are usually cov and Cranston(2005). Consult Torre-Bueno(1937)for
ered by the projecting eighth sternum (Fig. 2.9). further explanation of terms.
 iMkmm

^"^'4 ^
mr '%

SAMPLING AQUATIC INSECTS

Collection Devices, Statistical Considerations,
AND Rearing Procedures

John K. Jackson Richard W. Merritt

Stroud Water Research Center, Avondale, Michigan State University, East Lansing
Pennsylvania
Kenneth W, Cummins
Vincent H. Resh Humboldt State University, Arcata
University of California, Berkeley
Darold P. Batzer
University of Georgia, Athens

The study of aquatic insects depends on our abil COLLECTING AND SAMPLING DEVICES
ity to collect these organisms, which leads to the need A variety of approaches and devices have been
to make decisions about appropriate sampling used to collect aquatic insects or provide quantitative
devices and/or laboratory processing procedures to information on their richness, abundance, or biomass
separate the animals of interest from the abiotic (Table 3A). The table provided is not a complete list
material they reside in. However, the first step in ing of the references on sampling methods, and stu
planning any scientific study, before any sampling dents and researchers should continually check the
decision can be made, is to answer the WHY current literature for methods suited to their specific
question—why are we collecting samples? At the core objectives. The bibliographies of the Society for
of this question is the need to have a clearly defined Freshwater Science (formerly the North American
scientific hypothesis or goal that underlies the objec Benthological Society) contain scores of other papers
tives ofthe study. A clearly defined scientific question describing devices for collecting aquatic insects, as
then leads to other inquiries about what types of data does the extensive bibliography of Elliott eta/.(1993).
are needed (e.g., qualitative vs. quantitative, abun As noted by Cummins(1962)long ago,the number of
dance vs. biomass, population vs. community), where different samplers used for benthic macroinverte-
to sample (e.g.,the habitat such as a wadeable stream, brates is nearly equal to the number of benthic
deep river, wooded wetland, or stormwater pond, investigations!
and more specific locations such as upstream vs. The classification system used in Table 3A is based
downstream or littoral vs. profundal), and when to primarily on the habitat and community being
sample (e.g., once per year [e.g., spring vs. summer], sampled. Substrate composition, although not out
quarterly, monthly, according to flow regimes). lined in detail, also is an important consideration when
Answers to those questions are important when sampling the benthos (e.g., Minshall and Minshall
developing a plan that includes decisions about sam 1977; Rabeni and Minshall 1977; Lamberti and Resh
pling devices or laboratory processing procedures to 1978; Reice 1980; Minshall 1984), and the equipment
be used. It is also through this process that other data and techniques listed below may require modification
such as water chemistry, algal biomass, benthic depending on the substrate type. For example, an
CPOM (coarse particulate organic matter), etc., are Ekman grab is listed as an appropriate device for
identified as essential and can be added to the sam littoral benthos(IV. A.1. a. in Table 3A); however,the
pling plan. presence of sticks or even small stones could prevent

17
 00

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

Subhabitat Qualitative
Major and or Semi-
Sampling Ecological Quantitative quantitative
Habitat Community Sampler Figure Reference(s) Sampler Figure References

LOTIC HABITATS
1. Shallow Streams, A. Riffles (erosional
Rivers, and Springs zones)
1. Benthos
a. Sediments Surber sampler 3.7 1031, 2764, 3613, 4090, Aquatic net 3.2 6099, 3631, 3721
4289, 5823, 954, 5775,
4808

Mess or modified Mess 3.6 2571, 2902, 6363, 879, Hand screen collector 3.3 6099
sampler 1673

T-sampler 3.8 3655 Artificial substrate 3.23 2764, 3613, 3765,

samplers 5084, 5115, 5823, 954,
6416, 3371, 896, 2901,
6889, 6903

Individual stone, 5378, 2753, 6768, Individual stone sampler 1460, 6768, 3928
bedrock, or rock- 6214, 3928
outcrop sampler
Wilding or stovepipe 3.5 126, 5690, 6409, 6556 Kick sampling 2004, 2829, 4773,
sampler: box-type 3631, 5775, 3486, 6892
sampler
Suction samplers (air-lift 3.40 623, 4384, 6698, 2262, Recolonization 6596
and water pump) 725

Hand collection 3613, 6099

Leaf packs 3.25 4025, 4655, 5394,

624, 1450, 6896

Ellis-Rutter Stream 3.9 3174, 235, 6214 Photographic methods 1246, 1247
sampler
Riffle sampler 1077 Graded sieves 3.33 296, 4197, 6099

Electroshocking 3.29

b, Hyporheic area Implants 3.41 517, 1086, 2125, 2834, Freeze-core samplers 5752, 665, 4837,
(subterranean) 4144, 4781 (including electro- 664, 3579
freezing)
Pump sampling 6897, 6906
Standpipe corer 3.42 1733, 6583, 6595, 6597 Canister sampler 3.41 2125

) ) ))) ) ) ) ) ) ) )j ) ) ) ) I ) ) ) ) ) :i ^
) ) J ) ) ) 3 ) )))) ) )) ) ) ) 3 ) ) ) ) ) ) ) )

Table 3A Continued

Subhabltat Qualitative
Major and or Semi-
Sampling Ecological Quantitative quantitative
Habitat Community Sampler Figure Reference(s) Sampler Figure References

c. Plants Bag sampler 2833,6409 Needham apron net 6099

Surber sampler 3.7 Aquatic net 3.2 6099

Snag sampler
Stovepipe sampler 3.5 126, 6393 1395

Lambourn sampler 2587

d. Drift and Drift net 3.10 208, 1627, 6355, 6596, Aquatic net 3.2 6099
Neuston 6638, 91, 1673, 3822,
(surface) 4808

Plankton net 3.11 1627, 2833

Cushing-Mundle drift 1288, 4195
sampler
Hardy plankton 1624, 2833
indicator type sampler
Surface film sampler 1074
Colonization cages 2514

2. Emerging Adults Surface film sampler 1074 Hand screen collector 3.3 6099

Mundle pyramid trap 3.26 4193 Pan traps 2246, 3335

Stationary screen trap 2342, 2844, 3773 Window traps 984, 5837

Enclosed channels 6343 Light traps 5582

Floating emergence 3501, 3615,4130, 1352

Traps 3.9

B. Pools depositional
zones)
1. Benthos
a. Sediments Ekman grab with pole 3.16 2833 Aquatic net 3.2 299

Wilding or stovepipe 3.5 Graded sieves 3.33 6099

sampler
Single corerwith pole 3.12 1089, 2032, 6227
Suction samplers (air-lift 3.40 623, 725
and water-pump)
Mark-recapture 3.14

(continued)

SAMPLING
 Table 3A Continued

Subhabitat Qualitative
Major and or Semi-
Sampling Ecological Quantitative quantitative
Habitat Community Sampler Figure Reference(s) Sampler Figure References

b. Plants Bag sampler 2833 Needham apron net 6099

c. Drift and See Section I.A.I ,d See Section I.A.I.d.

Neuston

2, Emerging Adults See Section I.A.2 Section I.A.2

II. Large Rivers A. Riffles

1. Benthos
a. Sediments Suction samplers (air-lift 3.40 1648, 2036, 2285, Basket or cylindrical- 3.24 147, 174, 371, 1219,
and water pump) 6175, 1516, 2380, type artificial substrate 1847, 2034, 2337,
4384, 1517 samplers 2587, 2597, 3768,
3771, 4489, 5123, 6395

Suction samplers (air-lift 3.40 1516, 4532, 3651

and water pump)
SCUBA diving 2034, 2035, 2587, 1725 Single or multiple-plate 3.23 1423, 1847, 2424,
samplers 2574, 5123
Grab samplers 82, 1645, 1648 Drag-type samplers 1646, 1648, 6104
b. Drift and Floating drift trap (with 1627,4196
Neuston floats)
Hardy plankton 1624, 2833
indicator-type sampler
2. Emerging Adults Insect emergence traps 3399, 4130, 1352

Floating drift trap (with 4196

floats)
B. Pools
1. Benthos
a. Sediments Ponar grab 3.17 1645, 1648, 3517, 4801 Basket-type artificial 3.24 1847, 2034, 3768, 5123
substrate samplers
Petersen-type grabs 3.19 1645, 3517, 4674 SCUBA diving 1648, 2833

Ekman grab 3.16 1617, 1645, 1648,

2833, 3517
Core sampler 3.12 1648, 1846, 2032, 2118

) ) ) )) ) ) ) ) ) ) ) V ) ) ) ) ) ) ) ) ) ) ))
Exploring the Variety of Random
Documents with Different Content
 When Theo heard where I was going, she said she
would walk with me. We had a pleasant ramble through the
wood and down the Coombe to the village, and were most
hospitably received by good Mrs. Penrose, and entertained
with cakes and cream. Mr. Penrose was well pleased with
the idea, and said he would himself talk with Dinah and find
out her qualifications.

"I should like to be a parson's wife," said Theo, as we

walked homeward.

"You Theo!" I exclaimed, in amazement. "You of all

people."

"Yes, I of all people," she returned gaily. "It seems to

me such a useful, pleasant, quiet life."

"But I thought you did not like quiet," I said. "You

always seem to enjoy company so much."

"Well, so I do; and I like to dress prettily, just as I like

everything to be pretty and neat; but any head is not set on
such matters—no, not so much as Martha's, though she is
so demure. Perhaps not so much as yours is."

"You would make a good parson's wife in many ways, I

am sure of that," said I. "You would make every one like
you."

"I know I am not so very bright," said Theo; "I cannot

sing and play like you, nor read great books like Martha, nor
do any other grand things. But I like to help people enjoy
themselves in their own way, and to comfort them in
trouble if I can."

"I am sure you do," said I. "Janey Lee said the other
day when her child died it was a comfort just to have you
come in."

"Did she? I am very glad," said Theo. "But I don't know

what I did, only to sit by her, and let her weep, and by and
by draw her on to talk of the poor babe and its little pretty
ways. I never can preach to people in trouble. It seems
somehow unfeeling to talk to them of judgments and so on.
No, if I should marry a parson, I should let him do all the
preaching, you may be sure of that. I should content myself
with making his house pleasant, and cooking up messes for
the poor, and making baby things for the lying-in women.
That is my idea of a happy life."

It seemed as if Theo's idea of a happy life was like

enough to be fulfilled. She went on a little visit to her
godmother, any lord's sister, an elderly lady who had a
house near Exeter, where she maintained several young
ladies of reduced circumstances but good family, giving
them a suitable education, and a small dowry whenever
they settled in life.

Here she made the acquaintance of the Dean of Exeter,

a man, of course, a good deal older than herself, but of fine
presence and agreeable manners. He had always been a
good deal of a stickler for the celibacy of the clergy; but it
seems Theo found means to change his mind, for she had
not been at home a week before he followed her, and asked
her of her father in marriage.

It was one of those happy matches to which there

seems no objection on any side. The dean was rich and
greatly respected. He had beside his deanery a cure in the
same parish where my lady Jemima, my lord's sister,
resided, and a beautiful rectory, where in Theo might
concoct sick messes and make baby linen to her heart's
content. She had a small property of her own, and my lord
gave her a portion as to his own daughter.

Mrs. Martha's wedding (which I should have mentioned

in its proper place) was celebrated very quietly, as we were
all in recent mourning for my mother; but my lord was
determined that Theo should have a grand wedding. So she
did, indeed, with all proper ceremony from the first going to
church to the bedding of the bride. Matters of that sort have
greatly changed since that time, and I cannot but think for
the better, though I do hold that weddings should be
celebrated publicly and joyfully, not huddled up as if they
were something to be ashamed of. If matters go on as they
have begun, I expect my granddaughters will jump into a
carriage at the church door, and drive off to get as far as
possible from all their friends.

However; Theo's wedding was public enough. We had

the house full of guests, and among them two whom I had
no wish to see, and beheld with dread as birds of ill omen,
and so indeed they were. These were no other than my
cousin Betty and her husband. They had come to the
neighborhood to visit a cousin of Mr. Lovel's, and my lady
meeting them and learning who they were, thought she
could do no less than invite them to the wedding. My lord
did not look too well pleased when he heard of it, for he had
taken a great dislike to Betty upon their first meeting, but
he could not treat her otherwise than courteously in his own
house. As to Mr. Lovel, he never seemed to me to have any
character, but to be a mere lay figure for the display of
whatever mode in clothes or manners happened to be
uppermost.

Betty had not been one evening in the house before she
began exercising her powers. My lord was praising lip the
institution of marriage, of which he was a great promoter,
and my lady, smiling, called him a match-maker.

"Well, I am a match-maker, I don't deny it," said he.

"Would you be ashamed of it if you were me, cousin Lovel?"

Betty had been sitting rather silent, and I suppose he

meant to include her in the conversation. She answered at
once—

"No, indeed, my lord. It is a good vocation. I am sure I

have always thanked Vevette for betraying me to my
brother, and so bringing my marriage to pass sooner than I
could have done."

She spoke in those clear silver tones of hers, which

always commanded attention, and several people turned to
look at us. As may be guessed, I was covered with
confusion, but I made shift to answer.

"You certainly owe me no thanks, Betty, for what I

never did. I knew nothing of your affairs, and therefore
could not betray them, had I been so inclined."

"Oh, I beg your pardon!" said she, with her mocking,

superior smile. And then presently to me in a kind of stage
aside which every one about us could hear—

"What is the use of keeping up that stale pretence? I

suppose you did what you thought right, and I don't blame
you; but why deny what you and I know to be true?"

To this I made no answer whatever, and my lady

presently called upon me to sing. I by and by saw Betty in
close conference with Mrs. Bernard, and I had no doubt
from the looks Martha cast at me that I was the subject of
their conference.
 The next day brought home my lord's son, whom I had
not yet seen. He had been travelling abroad for some years,
but meeting the news of his sister-in-law's approaching
marriage in London, he had hurried home to be present on
the occasion. He was a fine, grave, soldierly-looking young
man, and very much like Andrew in the face, though taller
and with much more of courtly grace in his manner. He was
warmly welcomed by all, and especially by Mrs. Bernard. I
never saw her soften so much toward any one, and, indeed,
I believe he was the only person she ever really loved. He
was very polite and kind to me, and I naturally liked him
because he was so much like Andrew. He was musical, like
all his family, and we sang together a good deal. One
morning, as we were practising a song together, Betty
peeped into the room. I believe she thought I did not see
her, for she slipped out and presently returned with my
lady, whom I have no doubt she brought on purpose. They
stood listening a few minutes, and then Betty said half
under her breath, and with a sigh—

"Ah, my poor brother, I see his cake is dough; but no

doubt it is all for the best."

We stopped singing at this, and my lady asked me with

some sharpness whether I had been at the school that
morning. I told her no, and she at once thought of errands
for me, both there and at the village, which would keep me
busy all the morning.

"I will walk with you, cousin," said my young lord. "I
want to go down to the Cove and see Will Atkins."

Certainly, my lady had not mended matters for herself

or me. I got rid of my cousin as soon as I could, telling him
that I should be a long time at the school-house, and after
that had some poor people to visit. He was rather unwilling
to leave me, but I insisted, and he had to yield.

Betty staid two days longer, and then went back to

Allinstree, leaving mischief enough behind her. I do believe
my lady meant to be just to me, but it was hard to resist
the force of Betty's constant and artful insinuations, and she
really came to think that I was angling for her step-son. It
was not long, of course, before my lord took up the same
idea, and what was worst of all, my young lord soon showed
that he had no kind of objection to being angled for, and in
fact was very ready and even anxious to be caught.

From this time my life at the castle was not at all

comfortable. I missed the companionship of Theo, of whom
I had grown very fond, though she never filled Rosamond's
place to me. I missed my mother more and more. Besides,
my conscience was not easy. My lord and lady were good
people, as I have said; but the times were times of great
laxity. It was the fashion to profess great abhorrence of the
Puritans and their ways, and immense devotion to the
Church of England, and a good many people showed their
devotion by deviating as far as possible from the ways of
the precisians, as they were called.

We professed to observe Sunday—that is, we all went to

church in the morning, and my lady was very careful to see
that all the servants were present at prayers. But my lord
yawned over a play or romance all the evening when he had
no one to take a hand at cards or tables with, and when we
had company staying in the house the Sunday evening was
as any other. My young lord had taken up the kind of infidel
notions by which, as I said, some young men tried to
appear intellectual at a cheap rate, and he had brought
down some books of Mr. Hobbes with him which he would
fain have had me read; but that I refused. I had been
brought up to a strict observance of Sunday as a day of
worship and of sacred rest, and at first I was shocked at
what I saw. While my mother lived we usually spent our
Sunday evenings together in her own room, but after her
death, and especially after Dinah went away, I was easily
drawn into whatever was going on below stairs, even to
playing at tables with my lord, when he had no one else to
amuse him.

Then my old pleasure in dreams of wealth and

consequence revived. I was something of an heiress,
though my income was wholly dependent upon my lord's
pleasure or discretion till I should be of age, and so I had
plenty of attention. I began again to let the world come into
my mind, and, of course, it soon gained a foothold there
and ruled for the most part supreme.

Now and then, especially when anything strongly

reminded me of my mother, my better self—that self which
loved Andrew—came uppermost, but at such times, I
suffered so much from the reproaches of conscience, that I
strove by every means to stifle its voice. I said to myself
that my father and mother had been brought by the
circumstances in which they were placed to take a gloomy
view of religion and its requirements. That the strictness
which they had inculcated was not needful at present, and
that it tended (a favorite argument this with the devil) to
make religion unamiable. That a man or woman might be a
Christian and yet allow themselves many diversions which
the stricter sort denied.

In fine, my thought was, not how much I could do for

my Lord, but how much of the world I could safely keep for
myself. I was like a man who in time of war, instead of
fleeing to the safe hills in the interior of the country,
chooses to live as near the border as he can for the
advantage of keeping up a trade with the enemy. Instead of
simply shutting my ears to my cousin's infidel reasonings
and declining the subject, I allowed myself to listen to him,
and to be influenced by him to think that so long as a man
lived a good life, forms and doctrines mattered very little,
and I did not ask myself on what this good life was to be
founded.

In short, I grew more and more conformed to the world,

which in the bottom of my heart I had always loved, and in
proportion as I did so, the remembrance of my father and
mother, and of their teachings faded from my mind, I still
loved Andrew enough to reject with considerable vivacity a
proposal made me by young Mr. Champernoun, a
gentleman of the neighborhood, with a good fortune, and I
must say a personable and pleasing man, though grave
beyond his years.

My lord and lady were very much vexed at my refusal,

and used every argument to make me change my
resolution, saying that Mr. Champernoun was a much better
match than Andrew could ever be—which was true so far as
fortune went—and that I should perhaps never have so
good a chance to settle in life again.

"Well, well!" said my lord at last. "Wilful must have her

way. An I had not promised your honored mother never to
force your inclinations in any such matter, I should not use
so much ceremony with you, mistress! You should be made
to do what was best for you, whether you liked it or not."

He could not let the matter rest, but must needs take it
up again when his son was present.

"Vevette is right," said my young lord. "Were I in her

place I would not marry black Basil Champernoun either—a
sour Puritan and precisian whose father was in the favor of
Old Noll as long as he lived. I wonder, my lord, that you
could think of such a thing."

"Aye, aye, you would fain find her a husband, I dare

say; but mind, I will have none of that. If Vevette is flying
at any such game, she may as well come down at once."

"I am not flying at any game that I know of," said I,

feeling my cheeks flame, as what lady's would not.

"Your face tells another tale," returned my lord. "Such

blushes do not come for nothing."

"One may blush for others as well as for one's self," said
I, rising from the tables where I had been playing with my
lord, and in my confusion oversetting the board. And I
betook myself to my own room, nor did I leave it all the
next day, saying that I was ill at ease, which was the truth,
and wished to be quiet. Lewis must needs make matters
worse by coming to my door to inquire for me, and though I
did not see him, but sent him a message by Lucy, my new
little maid, his doing so did not help me with his father and
mother.

When I came down-stairs again, I found my lord had

gotten over his pet and was as gracious as before, but my
lady was very cool to me. She loved Lewis as her own son,
and was ambitious for him. The insinuations of Betty had
not been without their effect, and Mrs. Bernard, who was
settled in the neighborhood, threw all her influence on the
same side.

In short, I was very unhappy, and as I had about that

time an opportunity of writing to my Aunt Jemima in
London, I told her my troubles, and added that I knew not
what to do.
 The result was an immediate invitation from her and my
uncle to come to them in London, and make their house my
home. My uncle also wrote a letter to my lord, which I did
not see, but which I suppose satisfied him, for he made no
objection to my going, and my lady decidedly forwarded it.
Lewis had a great deal to say against it, but it may be
guessed that his arguments had no great weight.

It was settled that I was to travel with Theo and her

husband, who were going up in a week or two, and my lady
was directly in a great bustle to get me ready; now that
there was a chance of getting me off her hands, she was all
kindness once more.

The evening before I was to go to join Theo at Exeter, I

sought out my lady in her dressing-room and asked to
speak with her in private. I thanked her for her kindness to
me, and assured her that I had had no desire to displease
her in any way, and least of all by marrying Lewis. Then as
she gave me a kind though somewhat embarrassed answer,
I ventured to ask her what Betty had said about me. She
would not tell me at first, but presently changed her
purpose, and when I heard the cunning tale which Betty
had imposed upon her, I no longer wondered so much at
her change toward me. It was not only in the matter of the
meeting with Mr. Lovel, that she had misrepresented me,
but she had told my lady that I had avowed to her a settled
purpose to make myself the wife of some great man, and to
that very end had persuaded my mother to break off the
match with Andrew, at the very time that the change in my
fortunes made it likely that I should go to Stanton Court.

I explained the whole matter to my lady from beginning

to end, and she was pleased to say that I had wholly
exculpated myself, and to take shame to herself for being
so ready to believe evil. She kissed me and said she was
sorry I was going away, and bade me always think of
Stanton Court as my home. She had been very generous to
me before, and she now gave me a gold watch and a
beautiful set of pearl ornaments which she had bought in
Exeter. I believe she talked my lord over that night, for the
next day he told me he was sorry I was going away, and if I
would even now give up the plan, I should have a home at
the court as long as I liked, and he would not tease me to
marry any one.

But the die was cast. The step was taken which was the
beginning of a long journey—far longer indeed, than any of
us thought, and I had no mind to turn back.

CHAPTER XVI.
 LONDON.

THE next day I went to Exeter, from which place we

were to set out for London in a few days. I found Theo
living in a noble house, with everything pleasant about her,
and enjoying herself to the full. She had no fancy for the
journey to London, and would, I believe, much have
preferred going to the country rectory, whither Mr. Dean
usually retired in summer. We rode out to see the place,
and truly I did not wonder at her love for it—all about it was
so beautiful. There were no gentlemen's houses very near,
but my Lady Jemima, my lord's sister, lived, as I have said,
in an old mansion which had once been a convent of gray
nuns. The house stood on a rising ground, and was
beautifully embosomed in very ancient timber and a part of
this same wood reached even to the walls of the rectory
itself.

We visited the little village school, taught by a charming

old dame, and where Theo distributed buns, gingerbread,
and comfits with a lavish hand. Then we went into the
house, where all was in order, and where the old
housekeeper and her blooming neat maids welcomed us
with evident pleasure at seeing their mistress.

We also called upon my Lady Jemima, who was as great

a contrast to my own Aunt Jem as could well be conceived.
She was sitting at work among her family of maidens, who
were all busy with their fingers, while one read aloud. There
were six of them, all dressed alike in gray gowns and white
caps with blue ribbons, and I must say they looked very
bright and happy. Lady Jemima was a plain woman, with
none of the family beauty of color, but she had a most
sweet expression, at once benign and commanding. She
sent away her young ladies to walk, and then sat down to
talk with us.

"You have married off the last of your old family, have
you not?" asked Theo.

"Yes, only a month ago, and the child hath done well, I
think. Another has gone to be a governess in the family of a
distant cousin of ours, a rich sugar refiner's wife in Bristol,
and in one way or another, they are all scattered and doing
well for themselves. But my house is nearly full again."

"Not quite full, I hope, for I have a petition to make for

a poor maid, the eldest child of Mr. Brown, the vicar of
Torton," said Theo. And she proceeded to unfold the matter,
saying that the Curate was very poor, with a large family,
and this daughter being lame, was not fit for service.

"Are they so very poor?" asked Lady Jemima.

"They are poorer than they need be, if the wife were a
better manager," replied the dean's lady. "But she hath
been a waiting-gentlewoman to my Lady Saville, and still
sets herself up on her gentility, forsooth, cannot possibly
work with her hands, and talks of how she hath come down
in the world. The aunt, who is a good plain farmer's wife,
with a small army of children, tells me that this maid's
lameness hath come, she verily believes, from working
beyond her strength to make up her mother's deficiencies.
She is her father's greatest comfort, poor man, but he will
willingly spare her for the chance of having her recover her
health."

"Will you send him to see me?" asked Lady Jemima. "I
would talk the matter over with him myself, for no
disparagement to you, Theo," she added with a smile, "you
are one of those softhearted people who think everybody
ought to have everything, and as my means are limited, I
must make a discrimination, and not use them to encourage
idleness or improvidence."

Theo smiled in her turn, and admitted that she was

easily imposed upon. "But I am learning something, I
assure you," said she. "I have found out that all the clean
people are not saints and all the dirty ones reprobates,
which was the notion I at first set out with."

After a little more talk we had dinner with Lady Jemima

and the young ladies, and set out on our way home, calling
at the house of the curate I have mentioned.

Such a house—showing in every corner the results of

sluttishness and improvidence.

The poor man, into whose study we were shown, sat in

a ragged cassock, writing with one hand and holding a
sleeping infant on the other arm, while his lame daughter
was resting upon a rude couch or settle—a hard resting-
place it looked—keeping two more little ones quiet by telling
them a story, though her feverish cheeks and bright heavy-
lidded eyes showed how much she needed rest.

Another girl about twelve was clearing a table of the

remains of what certainly looked like a very scanty meal.
Theo at once took possession of the children, and
distributed some cakes among them, which they devoured
in a way that showed their dinner had still left them with an
appetite. She had also brought new gowns for the elder
girls, at sight of which the somewhat sullen face of the
second girl brightened, and she looked really pretty.

The father said just enough and not too much by way of
thanks, and promised that he would go to see Lady Jemima
next day. Just as we were about going, madame sailed into
the room, having evidently been busy attiring herself in the
remains of her old waiting-gentlewoman's finery. She was
loud in her thanks and praise of the gowns, and equally
loud in her lamentations over the state of her own
wardrobe, a hint of which Theo took no notice.

"I little thought I should live to receive charity," said the

foolish woman; "but when one weds beneath one's station,
there is no knowing what one will come to."

"As to that, I dare say your husband was so much in

love as to think you capable of filling any station," returned
Theo, wilfully misunderstanding her; whereat she tossed
her head, and looked ready to bite, but made no reply.

"I dare say she will make up the gowns for herself," said
Theo, when we had taken leave. "It is a wonderful thing to
see what sort of people little children are sent to, is it not?"

I agreed with her. I may as well say that the woman

flatly refused at first to let Sally go to Lady Jemima,
declaring that her lameness was more than half a pretence
to get rid of work. But the father had his way for once, and
poor Sally, if she did not recover, at least spent her last
days in peace.

In a day or two we went up to London, in the dean's

coach, with outriders, and spare saddle horses for one of us
to ride now and then. It was a toilsome journey—worse by
far than it is now, and that is saying a great deal. More than
once the coach was fairly stuck, and we had to borrow oxen
from the neighboring farmers to drag it out of the mire, and
once we just missed an attack from highwaymen. They
thought our party too strong, it seems, and let us pass, but
a gentleman with whom we had spent the evening before at
an inn, was stripped of all his own and his wife's valuables
and received a severe wound in the arm. However, in spite
of dangers and detentions, we arrived safely in London at
last, and I was left at my uncle's new house in Covent
Garden, whither he had removed at the death of my Aunt
Jean's father, who had left her quite a fortune.

My uncle and aunt were not at home, but I received

every attention from my aunt's waiting-gentlewoman, and
was installed in a pleasant room and treated to a cup of
chocolate. I was glad to go to rest early, as I was very tired
with the journey, and Mrs. Mercer said her lady would not
be at home till quite late. It was long before I could fall
asleep, there was such a noise in the street, but weariness
overcame me at last.

I slept soundly and awoke refreshed, though still

somewhat stiff with the jolting I had endured. I had meant
to begin the day with reading and devotion, but I was
hurried and a good deal in awe of the new waiting-damsel
my aunt had provided for me. I was afraid I should keep my
aunt waiting breakfast, and so went down without any
prayer whatever. Thus I began my new life with a false step.

I found my uncle much changed, and not for the better.

He received me very kindly, as did my aunt, but he looked
haggard, had grown older, and had a hard, worn
expression, as if he lived under the stress of some habitual
excitement. My aunt too looked older, and had lost a good
deal of her beautiful bloom. They both welcomed me kindly,
and any aunt began at once to talk of taking me out to the
theatre and the park so soon as I should be provided with
new clothes. My uncle said very little, and went out
immediately after breakfast. I saw his wife take him aside
and ask him some questions to which, judging from her
face, she did not receive a favorable answer.
 "But the child must have new clothes! I cannot take her
out with me till she is fit to be seen," I heard her say.

"Well, well. I suppose Lord Stanton has sent me some

money by the dean. I shall wait upon him as soon as it is
late enough. Meantime I can spare you this," putting some
gold into her hands. "It is a part of my winnings last night."

"Ah, Charles, if you would but quit gaming," said my

aunt, in a low tone, but not so low but that I heard her.

"How can I, child, when the king sets the example,

unless I withdraw from court altogether, and I suppose you
would not, have me do that?"

"No, you cannot do that," replied my aunt, "but then—"

"Don't trouble thy head about the matter," interrupted

my Uncle Charles. "If I lose one day, why I gain the next.
So it is all even. You will be an old woman before your time,
and have to take to painting, like my Lady Castlemaine—or
to devotion, which I should like still less."

So saying, he kissed her and went away, and she came

back to me with a little line of vexation between her arched
brows.

"Well, well! Men will be men. Come up-stairs, child, and

we will look over your wardrobe and see what you need."

I ventured to say that, my Lady Stanton had provided

me with everything she thought needful.

"Yes, I dare say, according to her notions. But she has

not been in London these seven years, and I dare say she
has not changed the fashion of her dress since that time."
 My new maid had unpacked all my things by this time,
and my aunt, though she criticised unmercifully the fashion
of my gowns and petticoats, yet allowed that Lady Stanton
had been very liberal.

"This may do well enough with a silk petticoat laid with

silver," said she, laying aside what was meant for my best
gown. "But you must have another and some lace whisks
and a hat and riding coat, and Mercer must curl your hair."

"It curls of itself," said I, "but I have always worn a

cap."

"Nonsense, child; what do you want of a cap? Come, I

shall allow yo no free will in this matter of dressing. You
must needs confess that I am the best judge, and be ruled
by me. You shall wear my t'other hat and mantle, and we
will drive to the Royal Exchange and buy you some gloves
and stockings and a fan and so on."

"But, Aunt Jem, I am in mourning," I ventured to say.

"Well, and so am I, child. Don't you see I am all in

black?"

Certainly she was in black, but I should never have

guessed she was in mourning, she wore so much lace and
fine cut work. However, I promised to be guided by her
judgment in all such matters, as indeed was no more than
fitting, seeing I had come to be under her care. We
presently went out in my uncle's coach, and were busy
shopping all the morning. I thought I never could use all the
things my aunt bought for me, and my head fairly whirled
with the excitement of seeing so many new places and
people.
 My aunt was in the very heart of the gayest society
about the court, and many were the salutes she received
from this and that great lady—even from my Lady
Castlemaine herself and another very handsome woman
whom she said was Mrs. Stewart, a great favorite of the
king's. When we had finished our shopping, we went into
the park, and here I saw the king and queen, the latter of
whom I had never beheld before. I thought her very sad-
looking, and remarked upon it to my aunt.

"Yes, poor thing, she is sad enough, and no wonder,

since she is silly enough to love her husband," said my
aunt.

"Do you think it silly for a woman to love her husband,

aunt?" I asked.

"Yes, when he does not love her. But in truth, the queen
is too grave and too devout to please a merry monarch like
King Charles."

"Perhaps she finds comfort in devotion," I ventured to

remark.

"Yes, I dare say. 'Tis the refuge of disappointed wives

and faded widows. Perhaps I may take to it some day—who
knows?"

I thought within myself that my mother always found

comfort in devotion, though she was by no means faded,
and that devotion when it was taken up in that way as a
last resort, was not like to afford any great solace; but I did
not venture to speak my thoughts. I had already learned to
be ashamed of being thought devout.

"And who is that young lady in attendance upon the

queen?" I asked.
 "That! Oh, that is Mrs. Godolphin," was my aunt's reply,
with a curious change of tone. "She is a true saint, if you
please. I do not believe the smile or frown of any or all the
kings in Europe would make her turn a hair's breadth to the
right hand or the left, in any matter of duty or religion. We
used to be great friends when we were young chits together
at school," and she sighed.

"And are you not friends now?" I asked.

"We have never quarrelled, child, if that is what you

mean, but she has gone her way, and I mine. There, we
won't talk of it. See there is the coach of the French
ambassador. Is it not fine? He has some fine lady and
gentleman visiting him from France. I dare say we shall
meet them to-morrow night. But we must be going home to
dinner."

My uncle was not at dinner, being in attendance upon

the Duke of York in some capacity or other. I forget what.
When the meal was over my aunt said she meant to take a
rest, and she dared to say I would like to do the same. I
took the hint and retired to my own room.

Here was a chance for the devotions I had neglected in

the morning, but it may be guessed that I was in no
promising frame for them. However, I read a chapter and
hurried over a few forms, and then spent the rest of the
afternoon reading a French romance I had found on my
table, and in practising upon the harpsicon my uncle had
sent home for me. He was very fond of music, and wished
me above all things to cultivate it and to improve my voice.

In the evening, my aunt entertained a small company of

her friends, and she would have me sing for them. I
received many compliments, both upon my voice and my
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