BIOLOGY AND ENVIRONMENT General Editor: Bernard Stonehouse. THE BIOLOGY OF PENGUINS Edited by BERNARD STONEHOUSE THE BIOLOGY OF MARSUPIALS

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1 EVOLUTIONARY ECOLOGY

2 BIOLOGY AND ENVIRONMENT General Editor: Bernard Stonehouse THE BIOLOGY OF PENGUINS Edited by BERNARD STONEHOUSE THE BIOLOGY OF MARSUPIALS Edited by BERNARD STONEHOUSE and DESMOND GILMORE EVOLUTIONARY ECOLOGY Edited by BERNARD STONEHOUSE and CHRISTOPHER PERRINS

3 EVOLUTIONARY ECOLOGY Edited by BERNARD STONEHOUSE School of Environmental Science University of Bradford and CHRISTOPHER PERRINS Edward Grey Institute of Field Ornithology University of Oxford M

4 The contributors 1977 All rights reserved. No part of this publication may be reproduced or transmitted, in any form or by any means, without permission. First published 1977 by THE MACMILLAN PRESS LTD London and Basingstoke Associated companies in Delhi Dublin Hong Kong Johannesburg Lagos Melbourne New York Singapore and Tokyo Paperback edition first published 1979 ISBN ISBN (ebook) DOI / Typeset by Reproduction Drawings Ltd Sutton, Surrey This book is sold subject to the standard conditions of the Net Book Agreement The paperback edition of this book is sold subject to the condition that it shall not, by way of trade or otherwise, be lent, resold, hired out, or otherwise circulated without the publisher's prior consent in any form of binding or cover other than that in which it is published and without a similar condition including this condition being imposed on the subsequent purchaser.

5 Contributors VAN BALEN, DR J. H- Institut voor Ecologisch Onderzoek, Kemperbergerweg 11, Arnhem, The Netherlands CAVE, DR A. J. - Institut voor Ecologisch Onderzoek, Kemperbergerweg 11, Arnhem, The Netherlands CHITTY, PRoFESSOR D. H.- Department of Zoology, University of British Columbia, Vancouver, Canada FRY, DR C. H.- Department of Zoology, University of Aberdeen, Aberdeen GIBB, DR J.- DSIR Ecology Division, Lower Hutt, New Zealand HARRIS, DR M.P.- Institute of Terrestrial Ecology, Banchory Research Station, Glassel, Banchory, Kincardine shire KLUYVER, PROFESSOR H. N.- Institut voor Ecologisch Onderzoek, Kemperbergerweg 11, Arnhem, The Netherlands KREBS, DR J. R.- Edward Grey Institute of Field Ornithology, Department of Zoology, University of Oxford, Oxford LEVIN, PROFESSOR D. A.- Department of Botany, University of Texas at Austin, Austin, Texas MEDWAY, LoRD -Great Glemham House, Great Glemham, Saxmundham, Suffolk 1 P1 7 1 LP NELSON, DR J. B.- Department of Zoology, University of Aberdeen, Aberdeen NEWTON, DR I.- The Nature Conservancy Council, 12 Hope Terrace, Edinburgh 9 NISBET, DR I. C. T. - Massachusetts Audubon Society, Lincoln, Massachusetts RIANS, PROFESSOR G. H.- Department of Zoology, University of Washington, Seattle 5, Washington 0RIANS, C. E.- Bowdoin College, Brunswick, Maine 0RIANS, K. J.- Overlake School, Redmond, Washington OwEN, DR D. F.- 66 Scraptoft Lane, Leicester PERRINS, DR C. M.- Edward Grey Institute of Field Ornithology, Department of Zoology, University of Oxford, Oxford PYE, PROFESSOR J.D.- Department of Zoology and Comparative Physiology, Queen Mary College, Mile End Road, London E1 4NS RicKLEFS, DR R. E.- Department of Biology, University of Pennsylvania, Philadelphia, Pennsylvania RosEN ZWEIG, DR M. L. -Department of Ecology, University of Arizona, Tucson, Arizona ScHAFFER, DR W. M.- Department of Biological Sciences, College of Liberal Arts, University of Arizona, Tucson, Arizona ScHAFFER, M. V.- Department of Biological Sciences, College of Liberal Arts, University of Arizona, Tucson, Arizona 85721

6 vi Contributors SNow, DR D. W. -Bird Room, British Museum (Natural History), Tring, Hertfordshire TURNER, PROFESSOR B. L.- Department of Botany, University of Texas at Austin, Austin, Texas WATSON, DR A.- Institute of Terrestrial Ecology, Banchory Research Station, Glassel, Banchory, Kincardineshire WYNNE-EDWARDS, PROFESSOR V. C. -c/o Department of Zoology, University of Aberdeen, Aberdeen ZAHAVI, DR A.- University of Tel Aviv, Tel Aviv, Israel

7 Contents Introduction Dr B. Stonehouse and Dr C M Perrins SECTION 1 POPULATION REGULATION AND THE FUNCTIONS OF 3 TERRITORY 2 Society versus the individual in animal evolution 5 Professor V. C Wynne-Edwards 3 Population limitation and the adaptive value of territorial behaviour in 19 Scottish Red grouse Lagopus l. scoticus Dr A. Watson. 4 Natural selection and the regulation of density in cyclic and non-cyclic 27 populations Professor D. Chitty 5 Factors affecting population density in the wild rabbit Oryctolagus 33 cuniculus (L), and their relevance to small mammals DrJ. A. Gibb 6 Song and territory in the Great tit Parus major 47 DrJ. R. Krebs SECTION 2 FEEDING ADAPTATIONS AND ECOLOGICAL SEGREGATION 63 7 Comparative ecology of seabirds in the Galapagos Archipelago 65 DrM P. Ha"is 8 Some relationships between food and breeding in the marine 77 Pelecaniformes DrJ. B. Nelson 9 Coexistence and diversity in heteromyid rodents 89 Dr M L. Rosenzweig 10 Courtship-feeding and clutch size in Common terns Sterna hinmdo 101 Dr l C T. Nisbet SECTION 3 BREEDING ADAPTATIONS AND REPRODUCTIVE RATES Timing and success of breeding in tundra-nesting geese 113 l Newton 12 The evolutionary significance of co-operative breeding in birds 127 DrC H. Fry

8 viii Contents 13 Helpers at the nest in some Argentine blackbirds 137 Professor G. H. Orians, C E. Orians and K. J. Orians 14 The occurrence of time-saving mechanisms in the breeding biology of 153 the Great tit Parus major Dr H. N. Kluyver, Dr!. H. vanbalen and Dr A. J. Cave 15 Latitudinal gradients in clutch size: an extension of David Lack's theory 171 Dr D. F. Owen 16 The role of predation in the evolution of clutch size 181 Dr C M Perrins 17 A note on the evolution of clutch size in altricial birds 193 Dr R. E. Ricklefs 18 Outch size in the Compositae 215 Professor D. A. Levin and Professor B. L. Turner SECTION 4 BEHAVIOUR, ADAPTATION AND TAXONOMIC 223 RELATIONSHIPS 19 Echolocation and the systematics of swiftlets 225 Lord Medway and Professor J. D. Pye 20 Duetting and other synchronised displays of the blue-backed manakins, 239 Chiroxiphia spp. Dr D. W. Snow 21 Reliability in communication systems and the evolution of altruism 253 Dr A. Zahavi 22 The adaptive significance of variations in reproductiv-e habit in the 261 Agavaceae Dr W. M Schaffer and M V. Schaffer Publications by David Lack 277 Author index 285 Subject index 291