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Dragonflies and Damselflies
Dragonflies and
Damselflies
Model Organisms for Ecological
and Evolutionary Research

Edited by

Alex Córdoba-Aguilar
Researcher, Universidad Nacional Autónoma de México

Christopher D. Beatty
Visiting Scholar, Program for Conservation Genomics, Stanford University

Jason T. Bried
Research Scientist, Illinois Natural History Survey, Prairie Research Institute,
University of Illinois at Urbana-Champaign
Great Clarendon Street, Oxford, OX2 6DP,
United Kingdom
Oxford University Press is a department of the University of Oxford.
It furthers the University’s objective of excellence in research, scholarship,
and education by publishing worldwide. Oxford is a registered trademark of
Oxford University Press in the UK and in certain other countries
© Oxford University Press 2023
The moral rights of the authors have been asserted
First Edition published in hardback in 2008
First Edition published in paperback in 2009
Impression: 1
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, without the
prior permission in writing of Oxford University Press, or as expressly permitted
by law, by licence or under terms agreed with the appropriate reprographics
rights organization. Enquiries concerning reproduction outside the scope of the
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address above
You must not circulate this work in any other form
and you must impose this same condition on any acquirer
Published in the United States of America by Oxford University Press
198 Madison Avenue, New York, NY 10016, United States of America
British Library Cataloguing in Publication Data
Data available
Library of Congress Control Number: 2022913669
ISBN 978–0–19–289862–3
DOI: 10.1093/oso/9780192898623.001.0001
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CPI Group (UK) Ltd, Croydon, CR0 4YY
Links to third-party websites are provided by Oxford in good faith and
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 List of Contributors

John C. Abbott Alabama Museum of Natural His- Institute of Entomology, Branišovská 31, České
tory & UA Museums Department of Research and Budějovice, Czech Republic
Collections, Mary Harmon Bryant Hall 357, The Leandro S. Brasil Institute of Biological Science and
University of Alabama, Tuscaloosa, AL 35487, USA Health, Federal University of Mato Grosso, Pontal
Daniel Acquah-Lamptey Philipps-University Mar- do Araguaia, Mato Grosso 78698-000, Brazil
burg, Department of General Ecology–Animal Jason T. Bried Illinois Natural History Survey,
Ecology, Karl-von-Frisch-Straße 8, 35043 Marburg, Prairie Research Institute, University of Illinois at
Germany Urbana-Champaign, 1909 S Oak St (Robert A. Evers
Fernanda Alves-Martins CIBIO-InBIO, Research Laboratory), MC 652, Champaign, IL 61820, USA
Centre in Biodiversity and Genetic Resources, Sebastian Büsse Department of Functional Mor-
University of Porto, Campus de Vairäo, 4485-661 phology and Biomechanics, Institute of Zoology,
Vairäo, Portugal Christian-Albrechts-Universität zu Kiel Am
Giacomo Assandri Area per l’Avifauna Migratrice Botanischen Garten 9 D - 24118 Kiel,
(BIO-AVM), Istituto Superiore per la Protezione e la Germany
Ricerca Ambientale (ISPRA), Via Ca’ Fornacetta 9, Seth Bybee Department of Biology, Brigham Young
I-40064, Ozzano Emilia, Italy University, 4102 LSB, Provo, UT 84604, USA
Celina B. Baines Department of Biology, McGill Uni- Lenize Batista Calvão Laboratório de Ecologia e
versity, 845 Sherbrooke St W, Montreal, Quebec Conservação (LABECO), Instituto de Ciências
H3A 1B1, Canada Biológicas, Universidade Federal do Pará, Av.
Christopher D. Beatty Program for Conservation Perimetral, 2-224 - Guamá, Belém, Pará, 66077-830
Genomics, Stanford University, 327 Campus Dr, Brazil
Room 220C, Stanford, CA 94305, USA Ulises Castillo-Pérez Instituto de Ecología, Univer-
Richard J. Bomphrey Structure and Motion Labo- sidad Nacional Autónoma de México, Circuito
ratory, Department of Comparative Biomedical Exterior s/n, 04510, Cd de México, México
Sciences, Royal Veterinary College, Hawkshead Francesco Cerini Experimental Ecology & Conserva-
Lane, Hatfield AL97TA, UK tion Lab, School of Biological Science, University
Cornelio Andrés Bota-Sierra Alabama Museum of of Bristol, Bristol Life Sciences Building, 24 Tyndall
Natural History & UA Museums Department Avenue, Bristol BS8 1TQ, UK
of Research and Collections, The University of Lenin Chari Department of Zoology and Entomology,
Alabama, Tuscaloosa, AL 35487, USA; Grupo de Life Sciences Building, Rhodes University, African
Entomología Universidad de Antioquia (GEUA), St, Grahamstown, Makhanda, 6139, South Africa
Universidad de Antioquia, Calle 67 No. 53–108, Viola Clausnitzer Senckenberg Natural History
Medellin, 13001 Colombia Museum Görlitz, Department of Entomology, Am
David Boukal University of South Bohemia, Fac- Museum 1, Görlitz 02826 Germany
ulty of Science, Department of Ecosystem Biology, Adolfo Cordero-Rivera Universidade de Vigo, ECO-
Branišovská 1760, České Budějovice, Czech Repub- EVO Lab., Escola de Enxeñaría Forestal, Campus
lic; Czech Academy of Sciences, Biology Centre, Universitario, 36005 Pontevedra, Spain
vi L I S T O F C O N T R I B U TO R S

Alex Córdoba-Aguilar Universidad Nacional Frank Johansson Uppsala University, Department

Autónoma de México of Ecology and Genetics–Animal Ecology, 75236
Charl Deacon Department of Conservation Ecology Uppsala, Sweden
and Entomology, Stellenbosch University, Private Leandro Juen Laboratório de Ecologia e Conservação
Bag X1, Matieland 7602, Stellenbosch, South Africa (LABECO), Instituto de Ciências Biológicas, Uni-
Paulo De Marco Viçosa State University, Department versidade Federal do Pará, Guamá, 66075-110,
of General Biology–Quantitative Ecology, Viçosa, Belém, Pará, Brazil
37570-000 Minas Gerais, Brasil Kari Kaunisto Biodiversity Unit, University of Turku
Amanda Dillon Albany Pine Bush Preserve Com- FI-20014, Turku, Finland
mission, 195 New Karner Rd, Albany, NY 12205, Rassim Khelifa Zoology Department, University of
USA British Columbia, Vancouver, BC, Canada; Bio-
Aleš Dolný Department of Biology and Ecology, Fac- diversity Research Centre, University of British
ulty of Science, University of Ostrava, Ostrava 710 Columbia, Vancouver, BC V6T 1Z4
00, Czech Republic Canada
Rachael Y. Dudaniec Department of Biological Sci- Gabriella J. Kietzka Department of Conservation
ences, 205B Culloden Rd, Macquarie University, Ecology and Entomology, Stellenbosch University,
Sydney, NSW 2109, Australia Stellenbosch University, Private Bag X1, Matieland
Merja Elo Department of Biological and Environ- 7602, Stellenbosch, South Africa
mental Science, P.O. Box 35, FI-40014, University Manpreet K. Kohli Department of Natural Sciences,
of Jyväskylä, Jyväskylä, Finland Baruch College, City University of New York, 17
Ryo Futahashi Bioproduction Research Institute, Lexington Avenue, New York, NY 10010; Division
National Institute of Advanced Industrial Science of Invertebrate Zoology, American Museum of Nat-
and Technology (AIST), Central 6,1-1-1 Higashi, ural History, 200 Central Park West, New York, NY
Tsukuba, Japan 10024, USA

Christine Goforth North Carolina Museum of Nat- Lesley T. Lancaster School of Biological Sciences,
ural Sciences, 11 W Jones St, Raleigh, NC 27601, University of Aberdeen, Aberdeen AB24 2TZ, UK
USA Sarah H. Luke School of Biosciences, University of
Miguel Gómez-Llano Department of Biological Sci- Nottingham, Sutton Bonington Campus, Nr Lough-
ences, 525 Old Main, University of Arkansas, borough, LE12 5RD, UK; 6 Department of Zoology,
Fayetteville, AR 72701, USA University of Cambridge, Downing St, Cambridge
CB2 3EJ, UK
Gregory F. Grether Department of Ecology & Evo-
lutionary Biology, University of California, Los Karen E. Mabry New Mexico State University, 1780 E
Angeles, CA 90095, USA University Ave, Las Cruces, NM 88003, USA

Filip Harabiš Department of Ecology, Faculty of Envi- Milen Marinov Biosecurity Surveillance & Incursion
ronmental Sciences, Czech University of Sciences Investigation Plant Health Team, Diagnostic and
Prague, Prague 16500, Czech Republic Surveillance Services, Biosecurity New Zealand;
Tiakitanga Putaiao
ˉ Aotearoa, Ministry for Pri-
Sönke Hardersen Reparto Carabinieri Biodiversità di
mary Industries Manatuˉ Ahu Matua, 14 Sir William
Verona, Centro Nazionale Carabinieri Biodiversità
Pickering Dr, Burnside, Christchurch 8053 New
“Bosco Fontana”, 46045 Marmirolo, Italy
Zealand
Adam Z. Hasik Department of Biological Sciences,
John Matthews Alliance for Global Water Adapta-
University of Arkansas, Fayetteville, AR 72701,
tion, 7640 NW Hoodview Circle, Corvallis, OR
USA; Mitrani Department of Desert Ecology, The
97330
Jacob Blaustein Institutes for Desert Research,
Ben-Gurion University of the Negev, Midreshet Michael L. May Department of Entomology,
Ben-Gurion 8499000, Israel Cook/Douglass Campus, Rutgers University, 96
Lipman Dr, New Brunswick, NJ 08901
Jaakko J. Ilvonen Finnish Environmental Insti-
tute SYKE, Latokartanonkaari 11, 00790 Helsinki, Shannon J. McCauley Department of Biology, Univer-
Finland sity of Toronto Mississauga, Mississauga, Ontario
L5L 1C6, Canada
 L I S T O F C O N T R I B U TO R S vii

Jenilee Montes-Fontalvo Central University of Maya Rocha-Ortega Departamento de Ecología Evo-

Venezuela, Facultad de Ciencias, Caracas 1040, lutiva, Instituto de Ecología, Universidad Nacional
Distrito Capital, Venezuela Autónoma de México, Circuito Exterior, Ciudad
André Morrill Department of Biology, Carleton Universitaria, 04510 Coyoacán, Distrito Federal,
University, 1125 Colonel By Dr, Ottawa, ON, Mexico
Canada Göran Sahlén The Rydberg Laboratory for Applied
Rosalind L. Murray Department of Biology, Univer- Science, RLAS, Halmstad University, P.O. Box 823,
sity of Toronto Mississauga, Mississauga, Ontario 30118 Halmstad, Sweden
L5L 1C6, Canada Gianandrea Salerno Dipartimento di Scienze Agrarie,
André Nel Institut de Systématique, Evolution, Bio- Alimentari e Ambietali, University of Perugia,
diversité, Muséum national d’Histoire naturelle, 06123 Perugia PG, Italy
CNRS, SU, EPHE, UA, 45 rue Buffon, CP50, F-75005 Michael J. Samways Department of Conservation
Paris, France Ecology and Entomology, Faculty of AgriSciences,
José Max B. Oliveira-Junior Institute of Water Sci- Stellenbosch University, Private Bag X1, Matieland
ences and Technology, Postgraduate Programs in 7602, Stellenbosch, South Africa
Society, Environment and Quality of Life; Biodi- Rosa Ana Sánchez-Guillén Red de Biología Evolu-
versity; Society, Nature and Development, Federal tiva, Instituto de Ecología, A.C., Carretera antigua
University of Western Pará (Unidade Tapajós), a Coatepec 351, Col. El Haya, Xalapa, Veracruz CP
Santarém, Pará, 68040-255, Brazil 91073
Michael A. Patten Ecology Research Group, Nord Melissa Sánchez-Herrera Universidad del Rosario,
University, Steinkjer, Norway Calle 12C Nº 6–25, Bogotá D.C., Colombia; Division
Mary Ann C. Perron Department of Biology, Univer- of Invertebrate Zoology, American Museum of Nat-
sity of Ottawa, 30 Marie Curie Private, Ottawa, ural History, 200 Central Park West, New York, NY,
Ontario K1N 6N5, Canada 10024, USA
Silvana Piersanti Department of Chemistry, Biology Xavier Sánchez-Rivero Departamento de Ecología
and Biotechnology, University of Perugia, Via Elce Evolutiva, Instituto de Ecología, Universidad
di Sotto, 8, 06123 Perugia PG, Italy Nacional Autónoma de México, Circuito Exterior,
Julien P. Renoult CEFE, Univ Montpellier, CNRS, Ciudad Universitaria, 04510 Coyoacán, Distrito
EPHE, IRD, Montpellier, France Federal, Mexico

Bertrand Piney Atlas Entomologique Régional Emily L. Sandall Center for Biodiversity and Global
(Nantes), 120 rue des leptures, 44850 Ligné, France Change, Yale University, New Haven, CT 06520,
USA; Yale University, Department of Ecology and
Stefan Pinkert Center for Biodiversity and Global
Evolutionary Biology, New Haven, CT 06511, USA
Change, Osborn Memorial Laboratories 165
Prospect St, Yale University, New Haven, CT Iago Sanmartín-Villar Universidade de Vigo, ECO-
06520, USA; Yale University, Department of Ecol- EVO Lab., Escola de Enxeñaría Forestal, Campus
ogy and Evolutionary Biology, New Haven, CT Universitario, 36005 Pontevedra, Spain
06511, USA; University of Marburg, Depart- Arnaud Sentis INRAE, Aix Marseille University,
ment of Conservation Ecology, FB 17, Biology, UMR RECOVER, 3275 Route de Cézanne- CS
Karl-von-Frisch-Straße 8, Marburg 35043, 40061, 13182 Aix-en-Provence, Cedex 5, France
Germany Camilla Sharkey Department of Ecology, Evolution
Ângelo Parise Pinto Universidade Federal do Paraná, and Behavior, University of Minnesota, Saint Paul,
Departamento de Zoologia, P.O. Box 19020, MN 55108, USA
81531-980 Curitiba, PR, Brazil Adam M. Siepielski Department of Biological Sci-
Justin Pomeranz Environmental Science Faculty, Col- ences, University of Arkansas, Fayetteville, AR
orado Mesa University, 1100 North Avenue, Grand 72701, USA
Junction, CO 81501, USA Hana Šigutová Department of Biology and Ecology,
Olga Popova Institute of Systematics and Ecology of Faculty of Science, University of Ostrava, 71000,
Animals, Siberian Branch of Russian Academy of Ostrava, Czech Republic
Sciences, Frunze 11, Novosibirsk 630091, Russia John P. Simaika IHE Delft Institute for Water Educa-
Manuela Rebora Department of Chemistry, Biology tion, Westvest 7, 2611 AX Delft, The Netherlands
and Biotechnology, University of Perugia, Via Elce Sabrina Simon Biosystematics Group, Wagenin-
di Sotto, 8, 06123 Perugia PG, Italy gen University & Research, Wageningen, The
Netherlands
viii L I S T O F C O N T R I B U TO R S

Brenda D. Smith Oklahoma Natural Heritage Inven- Julie Verheyen Evolutionary Stress Ecology and Eco-
tory, Oklahoma Biological Survey, University of toxicology, KU Leuven, Charles Deberiotstraat 32,
Oklahoma, 111 E Chesapeake St, Norman, OK 3000 Leuven, Belgium
73019, USA Marina Vilenica University of Zagreb, Faculty of
Robby Stoks Evolutionary Stress Ecology and Eco- Teacher Education, Trg Matice Hrvatske 12, 44250
toxicology, University of Leuven, Deberiotstraat 32, Petrinja, Croatia
Leuven, Belgium Giovanna Villalobos-Jiménez Centro de Investi-
Anton Suvorov Department of Genetics, University gación en Ciencias de Información Geoespacial
of North Carolina at Chapel Hill, 120 Mason Farm (CentroGeo), Circuito Tecnopolo Norte 117, Fracc.
Road, Chapel Hill, NC 27599, USA. Tecnopolo Pocitos, C.P. 20313, Aguascalientes,
Ami Thompson Biology Program, North Carolina Aguascalientes, México
Wesleyan University, 3400 N Wesleyan Blvd, Rocky Simon M. Walker School of Biomedical Sciences, Fac-
Mount, NC 27804, USA ulty of Biological Sciences, University of Leeds,
Nedim Tüzün Evolutionary Stress Ecology and Eco- UK
toxicology, University of Leuven, Deberiotstraat 32, Jessica L. Ware Division of Invertebrate Zoology,
Leuven, Belgium American Museum of Natural History, 200 Central
Khuong Van Dinh Department of Biosciences, Park West, New York, NY 10024, USA
University of Oslo, Blindern 0316 Oslo, Norway Maren Wellenreuther Plant and Food Research,
Yesenia M. Vega-Sánchez Instituto de Investigaciones Nelson, New Zealand, University of Auckland,
en Ecosistemas y Sustentabilidad, Universidad Auckland, New Zealand
Nacional Autónoma de México, Antigua carretera Erin White New York Natural Heritage Program,
a Pátzcuaro # 8701, Col. Ex-Hacienda San José de la SUNY College of Environmental Science and
Huerta, C. P. 58190 Morelia, Michoacán, Forestry, 625 Broadway, 5th Floor, Albany, NY
México 12233, USA
 List of Reviewers

Fernanda Alves-Martins Vincent Kalkman Rosa Ana Sánchez-Guillén

Ron Bassar Rassim Khelifa Romain Sarremejane
David Boukal Gabriella Kietzka Emily Schilling
Wade Boys Oleg Kosterin Tom Sherratt
Pawel Buczyński Lesley Lancaster Adam Siepielski
Francesco Cerini Harvey Lemelin Hana Šigutová
Ryan Cesar Harald Letsch Thomas Simonsen
Andreas Chovanec Natala Matushkina Brenda ”Bee” Smith
Viola Clausnitzer Michael May Frank Suhling
Idelle Cooper Shannon McCauley Anton Suvorov
Joan Damerow Roberto Munguía-Steyer Roy van Grunsven
Charl Deacon Michael Patten Jan van Tol
Aleš Dolný Dennis Paulson Natalia von Ellenrieder
Katie Duryea Mary Ann Perron Jessica Ware
Mark Forbes Edward Pfeiler Phillip Watts
Rosser Garrison Erik Pilgrim Jeff Wesner
Miguel Gómez-Llano David Polcyn Wade Worthen
Filip Harabiš Steve Presley
Pamela Hunt Jakub Prokop
Martin Jeanmougin James Pryke
Frank Johansson Manuela Rebora
Taku Kadoya Volker Rudolf
Göran Sahlén
Contents

Foreword xxiii

1 Introduction to Dragonflies and Damselflies, Second Edition 1

Alex Córdoba-Aguilar, Christopher D. Beatty, and Jason T. Bried

SECTION 1 GENOMICS 5
Edited by Alex Córdoba-Aguilar

2 Genomic insights into micro- and macro-evolutionary processes in

Odonata 7
Maren Wellenreuther, Rachael Y. Dudaniec, and Lesley T. Lancaster
2.1 Introduction 7
2.2 Genomic insights into population processes 8
2.2.1 Dispersal and connectivity 8
2.2.2 Range shifts and other spatial processes 9
2.3 Adaptation and adaptive trait evolution 11
2.3.1 Environmental adaptation 11
2.3.2 Morphological adaptation 12
2.3.3 Life stage-specific adaptation 13
2.4 Genomic variation associated with hybridization and speciation 13
2.4.1 Insights from genome assemblies into species and
order-specific functional traits 14
2.5 Conclusions and future directions 15
References 16

3 Transcriptomic insights into Odonata ecology and evolution 21

Seth M. Bybee, Ryo Futahashi, Julien P. Renoult, Camilla Sharkey, Sabrina Simon, Anton
Suvorov, and Maren Wellenreuther
3.1 Introduction 21
3.2 Color vision 23
3.3 Transcriptomic insight into the eco-evolutionary role of color
variation 25
3.3.1 Ecological significance of color variation within and between
species 25
3.3.2 Evolution of color phenotypes 25
3.3.3 Pigments 26
3.3.4 Structural colors 26
3.3.5 Genes involved in body color formation 27
xii CONTENTS

3.4 Embryogenesis 27
3.4.1 Gene expression during embryogenesis 28
3.5 Phylo-transcriptomics 28
3.6 Future directions 29
3.6.1 Color vision 29
3.6.2 Color 30
3.6.3 Embryogenesis 30
3.6.4 Phylogenomics 31
3.7 Conclusion 31
Acknowledgments 31
References 31

SECTION 2 ORGANISMAL STUDIES 37

Edited by Alex Córdoba-Aguilar

4 Functional morphology in Odonata 39

Sebastian Büsse
4.1 Introduction 39
4.2 Head 41
4.3 Head–thorax articulation 44
4.4 Thorax 44
4.5 Wings 46
4.6 Legs 47
4.7 Abdomen 47
Method boxes 50
References 52

5 The biomechanics of Odonata flight: structure, motion, and function 57

Richard J. Bomphrey and Simon M. Walker
5.1 Flight mechanics 57
5.2 Muscle activation 58
5.3 Wing structure 60
5.4 Flapping wing aerodynamics 62
5.4.1 Leading-edge vortex 62
5.4.2 Stroke plane 63
5.4.3 Planform 63
5.5 Aerodynamic interactions 64
5.5.1 Wing phasing 64
5.6 Flight control and sensing 66
5.6.1 Passive control 66
5.6.2 Active control 66
5.6.3 Predicting sensory inputs 67
5.7 Concluding remarks 68
Acknowledgments 68
References 68
 CONTENTS xiii

6 Odonata immunity, pathogens, and parasites 73

Adam Z. Hasik, Jaakko J. Ilvonen, Adam M. Siepielski, and Rosalind L. Murray
6.1 Introduction 73
6.2 Parasites 74
6.2.1 Viruses 74
6.2.2 Bacteria 74
6.2.3 Gregarines 74
6.2.4 Trematodes 74
6.2.5 Water mites 75
6.2.6 Parasitoids 75
6.2.7 Coinfection 76
6.3 Odonate immunity 76
6.3.1 Overview of insect immunity 76
6.3.2 Components of odonate immunity 76
6.4 Ecology and evolution of immunity and parasites 77
6.4.1 PO and food webs 77
6.4.2 Metacommunity structure 78
6.4.3 Coevolution 79
6.5 Future research directions 79
6.5.1 Genetics 79
6.5.2 Microbiome 79
6.5.3 Climate change 80
6.6 Conclusions 80
Acknowledgments 80
References 80

7 Odonata perception is more than vision 85

Manuela Rebora, Gianandrea Salerno, and Silvana Piersanti
7.1 Introduction 85
7.2 Adult 85
7.2.1 Antennae 85
7.2.2 Mouthparts and gustatory sensilla 90
7.2.3 The ovipositor sensilla: sensing the plant taste and stiffness 92
7.3 Nymph 92
7.3.1 Antennae 92
7.4 Conclusions and future perspectives 95
References 96

8 Thermoregulation in Odonata 101

Ulises Castillo-Pérez, Michael L. May, and Alex Córdoba-Aguilar
8.1 Introduction 101
8.2 Mechanisms of thermoregulation 101
8.2.1 Ectothermy and behavior 102
8.2.2 Ectothermy and color 103
8.2.3 Endothermy 105
8.3 Global change and thermal limits 107
8.4 Global change and body coloration 107
xiv CONTENTS

8.5 Odonate resilience: a link to thermoregulation? 107

8.6 Linking thermoregulation mechanisms to global temperature
changes 108
8.7 Some topics for future thermoregulation research 108
8.7.1 Genetics and physiology of thermoregulation 108
8.7.2 Mechanisms of thermoregulation 108
8.7.3 Trade-offs between thermoregulation and other functions 109
8.7.4 Human awareness via insect thermoregulation risk under
climate change 109
Acknowledgments 109
References 109

SECTION 3 POPULATION ECOLOGY 113

Edited by Christopher D. Beatty

9 Genetic structure, cryptic species, and hybridization: causes and

evolutionary consequences in Odonata 115
Rosa Ana Sánchez-Guillén, Yesenia M. Vega-Sánchez, and Melissa Sánchez-Herrera
9.1 Introduction 115
9.2 Gene flow within species: population genetic structure in odonates 116
9.3 Cryptic species in odonates 118
9.4 Gene flow between species: hybridization in odonates 119
9.5 Conclusions and research directions 124
Acknowledgments 125
References 125

10 Odonata survival: insights from mark–recapture experiments 129

Iago Sanmartín-Villar and Adolfo Cordero-Rivera
10.1 Introduction 129
10.2 The effect of marking 130
10.3 A review of the literature using marking methods with odonates 130
10.4 The effect of sex and age on survival and recapture rates 132
10.5 The effect of female color polymorphism 134
10.6 Individual and environmental covariates 135
10.7 Conclusions and further research 135
Acknowledgments 137
References 137

11 Migration in Anisoptera 141

Michael L. May and John H. Matthews
11.1 Introduction 141
11.2 Migratory case studies in odonates 142
11.2.1 Anax junius 142
11.2.2 Pantala flavescens 144
11.3 Migration and weather 146
11.4 Migration and reproduction 146
11.5 Population studies in migrating dragonflies 147
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Dam, 73; cited, 128; otherwise mentioned, 44, 64
Wingate, Sir Reginald: position of, 185; age of, 197; quoted on possibilities of
irrigation, 235, 236
Wodehouse, General, 166

Yusuf Pasha, 162

Zeila, 153
Zifta Barrage, 87, 88
Zubehr, 150, 152, 155, 157, 163
 THE END

BILLING AND SONS, LTD., PRINTERS, GUILDFORD

 MAP OF
EGYPT and the SOUDAN
1904

London: Stanford’s Geogl. Estabt.

(Large-size)
London: Edward Arnold.
 FOOTNOTES:

[1]1 metre = about 39 inches.

[2]A cubic metre of water equals, roughly, 1 ton.
[3]For the purpose of illustration, it is interesting to compare the
discharge of the Thames at Teddington:
Cubic Metres per
Second.
During June the average discharge for the twenty
35
years ending 1902 was
The average in June, 1903, was 178
The discharge on June 21, 1903, was 387
On February 21, 1900, it was 533
And on November 18, 1894 (greatest on record), it
1,065
was
I have given the discharge in cubic metres per second, the unit
generally in use on the Nile. On the Thames the figures are
usually given in gallons per day, which sounds much more
imposing. If the number of cubic metres per second is multiplied
by about 1,900,000, it gives approximately the number of gallons
per day. But, after all, the discharge of the Thames in June, 1903,
was not so very far below that of the Nile during the same month.
[4]The Egyptian peasant, however, refuses to accept the
prosaic evidence of his eyes about these rats, and, like the stout
conservative he is, prefers to believe the old tradition that they
turn to mud during the flood season. Many a man will gravely
assert that he has himself observed the transformation actually in
progress.
[5]1 Kantar = nearly 100 lbs.
[6]Cf. p. 71.
[7]Estimated. £E1 = £1 0s. 6d.
[8]Estimated.
 Transcriber's note:

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