Essential Ornithology This page intentionally left blank Essential Ornithology Graham Scott Department of Biological Sciences, University of Hull, UK 1 3 Great Clarendon Street, Oxford ox2 6dp 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 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 offi ces 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 © Graham Scott 2010 The moral rights of the author have been asserted Database right Oxford University Press (maker) First published 2010 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 Scott, Graham (Graham W.) Essential ornithology/Graham Scott. p. cm. ISBN 978–0–19–856997–8 (Pbk.) 1. Ornithology. 2. Birds. I. Title. QL673.S275 2010 598—dc22 2010010914 Typeset by SPI Publisher Services, Pondicherry, India Printed in Great Britain on acid-free paper by CPI Antony Rowe, Chippenham, Wiltshire ISBN 978–0–19–856998–5 (Hbk.) 978–0–19–856997–8 (Pbk.) 1 3 5 7 9 10 8 6 4 2 For Lisa, William, and Adam who have been very, very patient, and for Les and Shirley who help them to keep me birding! In memory of my father William Scott. He encouraged my interest in birds and for that I shall be eternally grateful. This page intentionally left blank Contents Preface xi Acknowledgements xiii 1 Evolution of birds 1 1.1 Birds are dinosaurs 2 1.2 Archaeopteryx 3 1.3 The evolution of modern birds 6 When did the modern lineages arise? 6 1.4 The phylogeny of birds 7 Morphological phylogeny 7 Character conservation and convergence 8 Biomolecular phylogeny 9 Putative phylogenies 10 1.5 Adaptive radiation and speciation 14 Darwin’s fi nches 15 Hybrids 16 Summary 22 Questions for discussion 22 Appendix 1 22 2 Feathers and fl ight 27 2.1 Feathers 27 Feather types 28 Contour feathers 28 Down feathers and semiplumes 31 2.2 Feather tracts 31 2.3 Feather colour 33 2.4 Feather damage 33 2.5 Feather maintenance 34 2.6 Moult 35 Moult strategies 37 VIII CONTENTS 2.7 Flight 39 Gliding and soaring 42 Flapping fl ight 42 Respiration and fl ight energetics 44 Flight speeds 48 2.8 The evolution of fl ight and fl ightlessness 49 Summary 50 Questions for discussion 50 3 Movement: migration and navigation 51 3.1 The ecology of migration 52 3.2 Genes and migration 55 3.3 Physiology of migration 59 3.4 The weather and migration 63 3.5 Navigation 64 Navigational cues 65 3.6 Spatial memory 68 Summary 70 Questions for discussion 71 4 Eggs, nests, and chicks 73 4.1 Sex and the gonads of birds 73 4.2 The egg 75 4.3 Egg shell coloration and patterning 80 Camoufl age 80 Egg mimicry 81 Egg recognition 81 Signals of quality 84 Pigments and shell quality 84 4.4 Incubation and the developing embryo 86 4.5 Hatching 90 4.6 Chicks 90 Summary 91 Questions for discussion 92 5 Reproduction 93 5.1 Males and females are diff erent 93 5.2 Mating systems 96 5.3 Courtship and mate choice 98 Resource provision 99 Ornaments and displays 101 Sharing a mate 101 CONTENTS IX 5.4 Bird Song 106 Song learning 107 Functions of song 107 Synchronized singing 110 5.5 Raising a family 112 Begging 113 Imprinting and independence 117 Summary 117 Questions for discussion 118 6 Foraging and avoiding predators 119 6.1 Foraging 119 Finding food and capturing prey 119 Sharing information 120 Foraging fl ocks 122 Do herbivores cooperate? 123 6.2 Optimal foraging 124 Feeding territories 125 6.3 Risk and foraging 127 6.4 Predator avoidance 128 Camoufl age 128 Predator distraction displays 129 Tonic immobility 130 Alarm calls 130 Mobbing 134 Flocks and colonies 134 Summary 137 Questions for discussion 137 7 Populations, communities, and conservation 139 7.1 Populations 139 Life history strategies infl uence population growth 140 Population change 140 7.2 Communities 144 Communities are dynamic 145 Niche divergence 147 Niche shifts, ecological release, and competition 150 7.3 Extinction and conservation 152 Conservation can be a success 154 Summary 155 Questions for discussion 155 Index 157