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It furthers the University’s objective of excellence in research,scholarship, and education by publishing worldwide in Oxford New York Auckland Cape Town Dar es Salaam Hong Kong Karachi Kuala Lumpur Madrid Melbourne Mexico City Nairobi New Delhi Shanghai Taipei Toronto With offices in Argentina Austria Brazil Chile Czech Republic France Greece Guatemala Hungary Italy Japan Poland Portugal Singapore South Korea Switzerland Thailand Turkey Ukraine Vietnam Oxford is a registered trade mark of Oxford University Press in the UK and in certain other countries Published in the United States by Oxford University Press Inc.,New York ©Oxford University Press 2005 The moral rights of the author have been asserted Database right Oxford University Press (maker) First published 2005 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,or under terms agreed with the appropriate reprographics rights organization. Enquiries concerning reproduction outside the scope of the above should be sent to the Rights Department, Oxford University Press,at the address above You must not circulate this book in any other binding or cover and you must impose the same condition on any acquirer British Library Cataloguing in Publication Data Data available Library of Congress Cataloging in Publication Data Data available Typeset by Newgen Imaging Systems (P) Ltd.,Chennai,India Printed in Great Britain on acid-free paper by Biddles Ltd.,King’s Lynn ISBN 0–19–852785–3 978–0–19–852785–5 ISBN 0–19–852786–1 (Pbk.) 978–0–19–852786–2 (Pbk.) 10 9 8 7 6 5 4 3 2 1 In memory of Vida Macrory 1919–2004 This page intentionally left blank Preface This book seeks to capture the contribution that a small freshwater fish—the guppy, Poecilia reticulata,has made to evolutionary ecology. Best known as a popular aquar- ium species and as the subject of an Ogden Nash poem (‘...swans have cygnets, seals have puppies,but guppies just have little guppies’) the guppy has also gained prominence as a model organism in the overlapping disciplines of animal behaviour, ecology,and evolution that together form the subject matter of this monograph. The recent surge of interest in the species has been fuelled both by the recognition that the guppy is uniquely placed to answer topical questions,and by its suitability for inte- grated field and laboratory investigations. Indeed more than 50% of papers on guppy evolutionary ecology have been published in the last decade. However,this work rests on the solid foundation of research laid down over the last century. My book,there- fore,is intended as a tribute to the early investigators who documented the patterns and posed the problems that have stimulated successive generations of researchers. My own research on guppies began when Ben Seghers introduced me to Trinidad. I remain indebted to Ben for sharing his considerable knowledge of guppies with me. Felix Breden, Gary Carvalho, Darren Croft, Christine Dreyer, Lee Dugatkin, John Endler, Doug Fraser, Jim Gilliam, Jean-Guy Godin, Lorenz Hauser, Andrew Hendry,Anne Houde,Michael Kinnison,Astrid Kodric-Brown,Jens Krause,Kevin Laland, Robin Liley, Cock van Oosterhout, John Reynolds, Helen Rodd, Nilla Rosenqvist,David Reznick,and Paul Shaw are just some of the many researchers in the guppy community who have provided advice,encouragement,and ideas over the years. Financial support has come from the Leverhulme Trust, the Natural Environment Research Council,The Royal Society,and the University of St Andrews. I am fortunate in having had excellent graduate students and postdocs and I thank them for their insights into fish behaviour. They include Anette Becher,Anuradha Bhat, Miguel Barbosa, Jonathan Evans, Jennifer Kelley, Siân Griffiths, Lorraine Hawkins,Phil Irving,Anna Ludlow,Kit Magellan,Jerome Masters,Iain Matthews, Alfredo Ojanguren,Charles Paxton,Lars Pettersson,Dawn Phillip,Helder Queiroz, and Stephen Russell. The Life Sciences Department at the University of the West Indies,St Augustine, is my academic home in Trinidad. Mary Alkins-Koo, Peter Bacon, Grace Sirju- Charran,Jake Kenny,Dawn Phillip,Rajindra Mahabir,Indar Ramnarine,and Chris Starr have made me welcome and have generously provided facilities and hospitality. viii • Preface Indar has been unfailingly supportive while Raj’s field expertise has played a crucial role in many investigations. Other friends and colleagues who have helped in diverse ways include John Armstrong,Jerry Coyne,Nalini Dass,Marian Dawkins,Sean Earnshaw,Jeff Graves, Peter Henderson, Felicity Huntingford, Ian Johnston, Constantino Macías-Garcia, Isobel Maynard, Manfred Milinski, Francis Morean, Martin Nowak, Geoff Parker, Linda Partridge,Gerard and Oda Ramsawak,Henry Rae,Mike Ritchie,Peter Slater, and Victoria Soo Poy. Christine Dreyer, Jonathan Evans, Doug Fraser, Jennifer Kelley, Tino Macías- Garcia,and Indar Ramnarine commented on the draft:mistakes and omissions that remain are entirely my own responsibility. Finally,I wish to express my gratitude to Paul Harvey and Bob May for encouraging me to write this book and for reading the manuscript,to Keith Horne for his support, and to Ian Sherman for making it happen. Anne Magurran St Andrews Contents 1 Preview 1 1.1 Evolutionary ecology in the context of this book 3 1.2 A primer of guppy biology 3 1.2.1 Nomenclature 3 1.2.2 Taxonomy and phylogeny 3 1.2.3 Distribution 4 1.2.4 Ecology 7 1.2.5 Reproduction 7 1.2.6 Life-history patterns 8 1.3 Overview of the book 9 2 Ecology of the guppy in Trinidad 11 2.1 What makes Trinidad special? 13 2.2 Predators 13 2.2.1 Fish predators 15 2.2.2 Composition of fish assemblages 24 2.2.3 Avian predators 24 2.2.4 Other vertebrates 27 2.2.5 Invertebrate predators 27 2.3 ...and productivity 29 2.4 Feeding behaviour of the guppy 30 2.5 Parasites 33 2.6 Dynamical aspects of ecology—variation over time as well as space 34 2.6.1 Density 34 2.6.2 Sex ratio 35 2.7 Geographical variation in guppy traits 39 2.8 Conclusions 40
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