Introduction to the Study of Animal Populations Introduction to the Study of Animal Populations H.G.ANDREWARTHA Department of Zoology, The University of Adelaide, Adelaide, South Australia SECOND EDITION CHAP MAN AND HALL LTD I I NEW FETTER LANE' LONDON EC4P 4EE First published 1961 by Methuen & Co. Ltd. Reprinted 1966,1968 Second edition 1970 Reprinted 1971 Published by Chapman & Hall Ltd. 11 New Fetter Lane, EC4P 4EE Reproduced and printed in Great Britain by Redwood Press Limited, London & Trowbridge ISBN-13: 978-0-412-11020-7 e-ISBN-13: 978-1-4613-3324-1 DOl: 10.1007/978-1-4613-3324-1 © 1961,1970, H. G. Andrewartha Softcover reprint of the hardcover 1s t edition 1961 All rights reserved. No part of this publication may be produced, stored in a retrieval system or transmitted in any form or by any means, electronic mechanical, photocopying, recording, or otherwise, 7.vithout the prior written permission of the publisher. Contents PREFACE TO SECOND EDITION page xi PREFACE TO FIRST EDITION xii PART I: THEORY The history and scope of ecology I. 1.0 INTRODUCTION 3 1.1 POPULATION ECOLOGY: THE STUDY OF THE DISTRIBUTION AND ABUNDANCE OF ANIMALS 6 1.2 THE BROAD BASES FOR POPULATION ECOLOGY 10 I.3 HOW TO WRITE ABOUT ECOLOGY 14 1.4 FURTHER READING 16 2. Environment 2.0 INTRODUCTION 17 2.1 THE IDEA THAT ENVIRONMENT IS DIVISIBLE INTO FIVE COMPONENTS 19 2.2 FURTHER READING 20 3. Components of environment,. resources 3.0 INTRODUCTION 22 3.1 RELATIVE SHORTAGES 25 3.11 Extrinsic relative shortages 25 3.12 Intrinsic relative shortages 30 3.121 Outbreaks of pests 31 3.2 ABSOLUTE SHORTAGES 32 3.21 Territorial behaviour in relation to re.iources 36 v Vi CONTENTS 3.211 Territorial behaviour in insects 37 3.212 Territorial behaviour in vertebrates 40 3.3 THE DISTRIBUTION AND ABUNDANCE OF RESOURCES 4. Components of environment; mates 4.0 INTRODUCTION 54 4.1 SHORTAGE OF MATES 54 4.2 THE PREVALENCE OF SPARSENESS 57 4.3 ADAPTATIONS THAT INCREASE THE CHANCE OF FINDING A MATE WHEN NUMBERS ARE FEW 58 5. Components of environment; predators and pathogens: aggressors 5.0 INTRODUCTION 60 5.1 PREDATORS 61 5.11 The 'biological control' of insect pests 61 5.12 Predators of vertebrates 67 5.2 PATHOGENS 69 5.21 The biological control of rabbits by myxomatosis 69 5.22 The biological control of insect pests 72 5.23 The activity of pathogens 75 5·3 AGGRESSORS 75 6. Components of environment; weather 6.0 INTRODUCTION 85 6.1 TEMPERATURE 86 6. I I The influence of temperature on speed ofd evelopment 86 6. II I The speed of development at constant .tempera- tures 87 6.12 The lethal influence of temperature 89 6.13 The limits of the tolerable zone 95 6.131 The influence of acclimatization on the limits of the tolerable zone 96 6.14 Behaviour in a gradient of temperature 98 6.15 Adaptations to temperature 99 6.2 MOISTURE 101 6.z1 Behaviour in relation to moisture 104 CONTENTS' vii 6.22 Physiological mechanisms for conserving water IIO 6.221 'Water-balance' in aquatic animals IIO 6.222 Conservation of water in terrestrial insects, ticks and snails 112 6.223 Conservation of water in terrestrial mammals 118 6.3 LIGHT 124 6.31 The influence of light in synchronizing life-cycles with each other 124 6.32 The influence of light in synchronizing the life-cycle with the season of the year 125 7. Components of environment; malentities 7.0 INTRODUCTION 129 7.1 BROWNING'S DEFINITION OF HAZARDS 130 7.2 MALENTITIES 130 8. Components of environment; more about the ecological web 8.0 INTRODUCTION 133 8.1 SOME ADVANTAGES IN BEING ONE OF MANY 134 8.2 INTERACTIONS IN THE ECOLOGICAL WEB MAY BE IMPORTANT 136 8.3 INTERACTIONS IN THE ECOLOGICAL WEB MAY RAMIFY 140 8.4 ECOLOGICAL BARRIERS AS PART OF THE ECO- LOGICAL WEB 143 9· Theory; the numbers of animals tn natural populations 9·0 INTRODUCTION 144 9.01 The meanings of 'common' and 'rare' 144 9.1 THE CONDITIONS OF 'COMMONNESS' AND 'RARE- NESS' IN LOCAL POPULATIONS 145 9.1I The conditions of commonness in local populations 145 9.12 The conditions of rareness in local populations 146 9· I 3 The way in u'hich weather keeps a local population rare relative to food and other resources 147 viIi' CONTENTS 9.2 THE CONDITIONS OF 'COMMONNESS' OR 'RARE- NESS' IN NATURAL POPULATIONS 148 9.21 General conclusions 155 9.22 Negative feed-back to density 156 9.221 The conditions of negative feed-back in natural populations 158 PART II: PRACTICAL COURSE Methods for estimating density, patterns of dis 10. tribution and dispersal in populations of animals 10.0 INTRODUCTION 165 10.1 THE MEASUREMENT OF RELATIVE DENSITY 165 10.2 THE MEASUREMENT OF ABSOLUTE DENSITY 170 10.21 Counting the whole population 170 10.22 The use of quadrats 170 10.23 The method of capture, marking, release and re- capture 172 10.3 THE MEASUREMENT OF 'AGGREGATION' IN NATURAL POPULATIONS 178 10.31 A test for randomness: the Poisson distribution 179 10.4 DISPERSAL 181 IO.41 Dispersal by drifting 181 IO.4II Dispersal of the bean aphis, Aphis fabae 181 10.412 Dispersal of wingless insects, mites and spiders 181 10.413 The dispersal of locusts 182 10.414 The dispersal of strong-flying insects by winds near the ground 183 10.415 The dispersal of marine species that drift with currents 184 10.42 Dispersal by swimming, walking or flying 184 10.421 The measurement of dispersal 185 10.43 Dispersal by clinging 189 10·44 The special importance of dispersal zn insect predators 189 10.45 The general importance of dispersal 190 10.5 CLASS EXPERIMENTS ON THE ESTIMATION OF DENSITY, DISTRIBUTION AND DISPERSAL 10.51 DenSity CONTENTS IX 10.51 I To compare the relative densities of two popu- lations of Paramoecium 191 10.512 To estimate the number of Tribolium in a box of flour 194 10.513 To estimate the number of Triboliu11l using stratified sampling 196 10.514 To estimate the number of Helix by capture, marking, release and recapture 197 10.515 To estimate the trends in the density of a population of Tl'lbolium using a trellis diagram 198 10.52 Patterns of dzstribution 204 10.521 To compare the distribution of Saissetia with the Poisson distribution 204 10.522 To compare the pattern of distribution of two colonies of Paramoecium 206 10.53 The measurement of dispersal 2II 10.531 To measure the rate of dispersal of Paramoecium 2II 10.532 To measure the rate of dispersal of Helix 217 I I. Physiological responses to temperature 11.1 THE INFLUENCE OF TEMPERATURE ON SPEED OF DEVELOPMENT 220 I I.I I To compare the rate of egg-laying of flour beetles M'er a range of temperature 220 I I .12 To measure the influence of temperature on the rate of egg-production by Thrips imaginis 225 11.2 THE LETHAL INFLUENCE OF EXTREME TEMPER- ATURE 229 11.21 To estimate the duration of exposure to 5°C re- quired to kzll Calandra granaria 229 11.22 To estimate the duration of exposure to 5°C re quired to kill the flour beetles Tribolium castaneum and T. confusum 234 Behaviour in relation to moisture and food 12. 12.1 MOISTURE 243 12.11 To measure the response of flour beetles to a gradient of moisture 243 12.2 FOOD 249 12.21 To measure the response of egg-laying females of Pieris to a variety of common garden plants 249 x . CONTENTS 12.22 To measure the response of egg-laying females of Pieris to different species of Brassicae 252 12.23 To measure the response of Helicella virgata to food and shelter 255 APPENDIX 261 BIBLIOGRAPHY AND AUTHOR INDEX 263 INDEX 277 Preface to the Second Edition In revising this book I have tried to bring the theory of environment up to date in the light of certain important criticisms that have appeared since 1961, especially in papers by T. O. Browning and D. A. Maelzer, and in the light of experience gained while using the book as a text for an undergraduate course in population ecology in the University of Adelaide. As a consequence the order in which the argument is pre sented has been altered. Some new material has been introduced to expand the discussion of certain topics, especially resources, pathogens, aggressors and territorial behaviour. But the general approach to the subject and the general theory remains very much the same as in the first edition. I am grateful to Professor F. Fenner and Dr F. N. Ratcliffe and to Cambridge University Press for permission to reproduce Fig. 5.04; to Professor D. O. Chitty and the Ecological Society of Australia for permission to reproduce Fig. 5.05 (with minor modifications); Fig. 3.03 has been modified from a figure in a paper by H. G. Andrewartha and T. O. Browning first published in the Journal of Experimental Biology. Adelaide, 1970 H. G. A. xi
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