Adaptations of Desert Organisms
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1 Adaptations of Desert Organisms Edited by J.L. Cloudsley-Thompson Springer Berlin Heidelberg New York Barcelona Budapest Hong Kong London Milan Paris Tokyo
2 Volumes already published Ecophysiology of the Camelidae and Desert Ruminants By R.T. Wilson (1989) Ecophysiology of Desert Arthropods and Reptiles By J.L. Cloudsley-Thompson (1991) Plant Nutrients in Desert Environments By A. Day and K. Ludeke (1993) Seed Germination in Desert Plants By Y. Gutterman (1993) Behavioural Adaptations of Desert Animals By G. Costa (1995) In preparation Invertebrates in Hot and Cold Arid Environments By L. S0mme (1995) Bioenergetics of Desert Invertebrates By H. Heatwole (1995)
3 Giovanni Costa Behavioural Adaptations of Desert Animals With 56 Figures Springer
4 Prof. Dr. Giovanni Costa Universita di Catania Dipartimento di Biologia Animale Via Androne Catania, Italy Cover illustration: photograph by J. L. Cloudsley- Thompson ISBN-13: DOl: / e-isbn-13: Library of Congress Cataloging-in Publication Data. Costa, Giovanni, 1942~. Behavioural adaptations of desert animals I Giovanni Costa. p. cm. ~ (Adaptations of desert organisms) Includes bibliographical references (p. ) and index. 1. Desert fauna ~ Behavior. 2. Adaptation (Biology) 1. Title. II. Title: Behavioral adaptations of desert animals. Ill. Series. QL 116.C ' 2652 ~ dc This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilm or in any other way, and storage in data banks. Duplication of this publication or parts thereof is permitted only under the provisions of the German Copyright Law of September 9, 1965, in its current version, and permission for use must always be obtained from Springer-Verlag. Violations are liable for prosecution under the German Copyright Law. Springer-Verlag Berlin Heidelberg 1995 Softcover reprint of the hardcover I st edition 1995 The use of registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. Typesetting: Macmillan India Ltd., Bangalore-25 SPIN: /SPS ~ I 0 ~ Printed on acid-free paper
5 Preface From the beginning of my scientific activity, I have been engrossed with the study of animal behaviour. I have always been convinced that animals, as regular inhabitants of specific habitats, evolved winning strategies to adapt to apparently adverse environmental conditions (and more generally to their ecosystems); so it has been natural for me to carry out ecoethological research. I began by studying animals living in sandy/coastal ecosystems and, subsequently, took a great interest in desert animals. In this environment, I expected to find striking adaptations but I must admit that even my fertile imagination was surpassed by what I learnt through both personal experience and the study of the rich scientific literature on the behaviour of desert animals. For this reason I accepted - with pleasure and also with some concern - the invitation of my friend John Cloudsley-Thompson to contribute, with this work of mine, to the series Adaptations of Desert Organisms. I hope that this book will be useful to its readers and will discredit some myths based on incorrect but widespread beliefs, concerning deserts and their faunas. Indeed, many people believe that the desert is a lifeless environment - in everyday language 'desert' is almost synonymous with 'uninhabited'. On the contrary, however, the student of the biology of arid environments knows very well that they are populated by a rich and varied fauna. In connection with this first error, is another, according to which the desert is a hostile habitat: but, since it is true that a desert biotope presents unbearable life conditions for organisms from other ecosystems, the desert promotes its own biocoenosis: in fact, the latter has survived there for a very long time. However, what I have said is true for every environment: almost any organism, if moved from its natural habitat to another, will respond like a "fish out of water". It must be borne in mind that the more selective a biotope is, the more specialized its biocoenosis becomes: plants and animals are strictly linked with their
6 VI Preface habitat, and their particular adaptations are the result of this relationship. This, together with the decisive role of behaviour in animal survival, is what justifies this book. Catania, Spring 1995 G. Costa
7 Acknowledgements I wish to express my gratitude to the editor of this Series, Prof. John Leonard Cloudsley-Thompson. He provided me with advice, suggestions and help. In addition, he undertook the hard task of revising the text and translating my 'dog' English into intelligible language. My thanks are also due to Dr. Dieter Czeschlik, Biology Editorial of Springer-Verlag, for his confidence and boundless patience. I am deeply grateful to Dr. Erminia Conti, my pupil, who assisted me in typing the manuscript and preparing the original illustrations. Moreover, the responsibility for encouraging me to continue working, whenever I wanted to abandon the undertaking rests entirely with her!
8 Contents Introduction Biotope and Vegetation Features.... Definition of Desert.... Factors of Aridity.... Classifications of Deserts.... Tropical and Subtropical Deserts.... Cool Coastal Deserts.... Rain Shadow Deserts.... Interior Continental Deserts.... Polar Deserts.... Sandy/Coastal Areas.... Semi-Desert Areas.... Biotope Features.... Temperature.... Atmospheric Humidity.... Wind.... Characteristics of Desert Vegetation Desert Zoocoenosis General Characteristics Main Animal Groups Protozoans TurbellarianFlatworms Rotifers Nematodes Annelids Gastropods Tardigrades Arachnids Crustaceans Myriapods '" Insects Fishes Amphibians... 34
9 x Contents Reptiles Birds Mammals Thennohygric Regulation.... Timing of Activity Rhythms.... Modulation of Activity Rhythms.... Exploitation of Substratum Resources.... Subterranean Life.... Digging Behaviour.... Looking for Sheltered Microhabitats.... Exploitation of Water Resources.... Exploitation of Body Resources.... Sun-Basking.... Self-Production of Shadow.... Body Orientation.... Locomotor Strategies.... Fog-Basking.... Exploitation of Social Resources.... Self-Protective Mechanisms.... Primary Self-Protection.... Anachoretic Behaviour.... Batesian Mimicry.... Crypsis.... Aposematism.... Secondary Self-Protection.... Retreat.... Escape.... Deimatic Behaviour.... Thanatosis.... Deflection of Attack.... Counterattack.... Social Defense Strategies.... Patterns of Movement.... Locomotor Patterns.... Crawling.... Walking.... Digging.... Swimming.... Orientation Mechanisms.... Primary Orientation.... Secondary Orientation.... Dispersal and Migration
10 Contents XI Exploitation of Food Resources.... Herbivores.... Insects.... Other Invertebrates.... Rodents.... Ungulates.... Other Vertebrates.... Carnivores.... Invertebrates.... Vertebrates Detritivores.... Reproductive Regulation.... Timing of Reproduction.... Partner Location.... Courtship and Mating.... Pair Stabilization; Sexual Groups.... Parental Care and Developmental Regulation.... Social Behaviour.... Sociality in Desert Environments.... Territorial Phenomena.... Learning and Culture lo2 lo3 lo llo References Subject Index
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