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RODENTS IN DESERT ENVIRONMENTS
MONOGRAPHIAE BIOLOGICAE
Editor
J.ILLIES
Schlitz
VOLUME 28
DR. W. JUNK h.v. PUBLISHERS THE HAGUE 1975
RODENTS IN DESERT ENVIRONMENTS
Edited by
I. PRAKASH & P . K. GHOSH
DR. W. JUNK h.v. PUBLISHERS THE HAGUE 1975
ISBN-13: 978-94-010-1946-0 DOl: 10.1007/978-94-010-1944-6
e-ISBN-13: 978-94-010-1944-6
© 1975 by Dr. W. Junk b.v., Publishers, The Hague Cover Design M. Velthuijs, The Hague
Softcover reprint ofthe hardcover 1st edition 1975
Zuid-Nederlandsche Drukkerij N.V., 's-Hertogenbosch
I. II.
III. IV. V.
VI.
VII.
VIII. IX.
X. XI.
XII. XIII. XIV. XV.
XVI.
XVII. XVIII.
XIX.
XX.
XXI. XXII.
XXIII.
CONTENTS
Chapters' Contents Preface ..... . Authors' Addresses The Desert as a Habitat The Ecology of Rodents in the northern Sudan The Rodents of the Iranian Deserts . . . . . Comparative Ecological Notes on Afghan Rodents. The Population Ecology of the Rodents of the Rajasthan Desert, India. . . . . . . . . . . . . . . . . . . Outbreaks of Rodents in Semi-Arid and Arid Australia: Causes, Preventions and Evolutionary Considerations. . Observations of Argentine Desert Rodent Ecology, with Emphasis on Water Relations of Eligmodontia typus. . . . La diversite des Gerbillides. . . . . . . . . . . . . Some Observations on Ecological Adaptations of Desert Rodents and Suggestions for further Research work. The Behavior Patterns of Desert Rodents Activity Patterns of a Desert Rodent. Patterns of Food, Space and Diversity . Desert Coloration in Rodents . . . . . The Biology of some Desert-Dwelling Ground Squirrels. Reproductive Biology of North American Desert Rodents Rodent Faunas and Environmental Changes in the Pleistocene of Israel . . . . . . . . . . . . . . . Prehistoric Rodents of the Middle East. . . . . . . . Desert Rodents: Physiological Problems of Desert Life. . Ecophysiology of Water and Energy in Desert Marsupials and Rodents . . . . . . . . . . . . . . . . . . . Thermo-Regulation and Water Economy in Indian Desert Rodents . . . . . . . . . . . . . . . The Physiological Adaptations of Desert Rodents. . . . Nematode Parasites of the Indian Desert Rodents. . . . Ecology of Desert Rodents of the U.S.S.R. (Jerboas and Gerbils) . . . . . . .. Author Index. . . . . . Genus and Species Index. Subject Index . . . . .
VII XIII XVI
1 15 47 59
75
117
155 177
185 189 225 241 269 277 305
331 363 379
389
397 413 445
465 599 607 617
CHAPTERS' CONTENTS
I. The Desert as a Habitat by]. L. CLOUDSLEy-THOMP-SON 1 Introduction 1 Distri bu tion 1 Soils. 3 Climate 5 Vegetation 11 References 12
II. The Ecology of Rodents in the northern Sudan by D. C. D. HAPPOLD 15 Introduction 15 Desert Species. 22 ] ebel Species 30 Riverine Species. 31 Commensal Species 31 Reproduction. 31 Other Adaptations for Life in the Desert 35 Discussion 39 Acknowledgements 44 References 44
III. The Rodents of the Iranian Deserts by X. MISONNE 47 Introduction 47 The Rodents: northern and southern Types. 49 Annual Rhythm 52 Population Densities . 54 Some Peculiarities. 56 References 58
IV. Com.parative Ecological Notes on Afghan Rodents by]. GAISLER 59 Introduction 59 Results. 60 Discussion 69 Summary 72 References 73
V. The Population Ecology of the Rodents of the Rajasthan Desert, India by ISHWAR PRAKASH 75 Introduction . . . . 75
. Population Structure. 81 Ranges of Movements 95 Food . . . . . . . 96 Predator-Prey Relationship. 101 Breeding Season and Litter Size . 103 References . . . . . . . 114
VI. Outbreaks of Rodents in Setni-Arid and Arid Australia: Causes, Preventions, and Evolutionary Con-siderations by A. E. NEWSOME & L. K. CORBETT 117 Introduction . . . . . . . . . . . . 117 Climates in Central Australia. . . . . 119 The Study-Area: Habitats and Rodents. 123 Populations. . . . . . . . 130 Conclusions and Discussion . 143 Acknowledgements 151 References . . . . . . . . 151
VII. Observations of Argentine Desert Rodent Ecology, with Etnphasis on Water Relations of Eligmodontia typus by M. A. MARES. . . . . . . . . . . 155 Introduction . . . . . . . . . . . . . . . 155 The Argentine Monte and some of its Rodents. 156 Methods and Materials. 164 Results and Discussion . 165 Acknowledgements 173 References . . . . . . 174
VIII. La diversite des Gerbillides par F. PETTER. 177
VIII
IX. Sotne Observations on Ecological Adaptations of Desert Rodents and Suggestions for further Re-search work by A. DE VAS 185 Introduction . . . . . . 185 The Desert Environment . 185 Ecological Adaptations. . 186 Need for further Research 188 References . . . . . .. . 188
X. The Behavior Patterns of Desert Rodents by J. F. EISENBERG ............... . 189
Introduction . . . . . . . . . . . 189 The Evolution of Desert Adaptations. 190 A Comparison of Behavior Patterns. . 196 A Comparison of Behavior Patterns in: Meriones, Gerbillus and Perognathus . . . . . 210 Spacing and Communication 217 Acknowledgements 221 References . . . . . . . . 221
XI. Activity Patterns ofa Desert Rodent by N. R. FRENCH 225 Introduction . . . . . . . . . . . . . . . 225 Methods. . . . . . . . . . . . . . . . . 226 The Microdosimeter and the Index of Activity. 227 Variation in Activity. . . . . . . . . . . . 229 Climatological Variables and Animal Activity . 229 Analysis . 232 Discussion 237 Summary 238 References 239
XII. Patterns of Food, Space and Diversity by M. L. ROSENZWEIG, BARBARA SMIGEL & A. KRAFT. 241 Introduction . . . . . . . . . . . 241 Resource Allocation by Seed Selection 242 Habitat Selection in Space . . . . . 251 The Pattern of Local Species Diversity 260 Acknowledgements 266 References . . . . . . . . . . . . 266
XIII. Desert Coloration in Rodents by D. L. HARRISON. 269 Introduction . . . 269 Desert Coloration . 269 Acknowledgements 275 References . . . . 275
XIV. The Biology of so:me Desert-Dwelling Ground Squir-rels by A. C. HA WBECKER . 277 Introduction . 277 Reproduction. . . . . . 279 Food Habits . . . . . . 288 Population Characteristics 294 Habitat Factors . 297 References . . . . . . . 302
IX
xv. Reproductive Biology of North ADlerican Desert Rodents by H. D. SMITH & C. D. JORGENSEN. 305 Introduction . . . . . . . . . . . . . 305 Reproductive Biology: Species Summaries. . . 308 References . . . . . . . . . . . . . . . . 328
XVI. Rodent Faunas and EnvironDlental Changes in the Pleistocene of Israel by E. TCHERNOV . . . . . . . 331 Introduction . . . . . . . . . . . . . . . . . . . 331 The Main Biogeographical Changes in the Near-East since the Miocene . . . . . . . . . . . . . . . . . . . 331 The Composition of the Rodents Faunas in the Pleistocene of Israel . . . . . . . . . . . . . . . . . . . . . 336 The Main Ecological Changes in the Quarternary of Israel . . . . . . . . . . . . . . . . . . . . . . 347 Analysis of the Main Habitats Occupied by the Pleistocene Rodents of Israel . . . . . . . . . . . . . 349 Note on the Origin of the Israeli Desert Rodents 356 References . . . . . . . . . . . . . . . . 361
XVII. Prehistoric Rodents of the Middle East by PRISCILLA F. TURNBULL. . . 363 Introduction . . . 363 Prehistoric Rodents 364 Discussion . . . . 374 Acknowledgements 376 References 376 Addenda. . . . . 378a
XVIII. Desert Rodents: Physiological ProbleDls of Desert
x
Life by K. SCHMIDT-NIELSEN . . . 379 Introduction . . . . . . . . . . . 379 Problems of Temperature Regulation. 380 Water Balance, Intake and Loss. 382 References . . . . . . . . . . . . 388
XIX. Ecophysiology of Water and Energy in Desert Marsupials and Rodents by W. V. MACFARLANE. 389 Introduction . . . . . . . . . . 389 Methods . . . . . . . . . . . . 390 Ecophysiology of Water and Energy 390 Discussion 394 References . . . . . . . . . . . 395
XX. TherDlo-Reguiation and Water EconoDlY in Indian Desert Rodents by P. K. GHOSH. . . . . . . . . . 397
XXI.
XXII.
XXIII.
Introduction . . . Methods ..... Thermo-Regulation Water Economy. Conclusion. References . . .
The Physiological Adaptations of Desert Rodents by L. I. GHOBRIAL & T. A. N OUR. .
Introduction . . . . . . . . . . . . . . . . Water Conservation in Desert Rodents. . . . . Structure and Concentrating Mechanisms in the Desert Rodent Kidney.. .......... . Tolerance to Heat and Thermo-Regulation of Desert Rodents .......... . Reproduction in Desert Rodents. References . . . . . . . . . .
NeDlatode Parasites of the Indian Desert Rodents by S. JOHNSON . . . .
Introduction . . . . . . . . . . . . . . . . Material and Methods. . . . . . . . . . . . Nematode Parasites of the Indian Desert Rodents Discussion References .
Ecology of the Desert Rodents of the U.S.S.R. (Jerboas and Gerbils) by N. P. NAUMOV & V. S. LOBACHEV.
Introduction Jerboas . Gerbils References
Author Index. Genus and Species Index. Subject Index . . . . .
397 397 398 401 410 410
413 413 416
428
434 438 441
445 445 445 445 462 463
465 465 468 524 590
599 607 617
Xl
PREFACE
Ever since BUX.TON published his Animal Life in Deserts in 1923, much individual, collective and organisational effort, both at national and international levels, has gone into the unravelling of the mystique of desert living. Man's interest in the desert must have been a primordial element in his cultural evolution as it was in the womb of deserts that human civilization had dawned and has since been thriving for many millennia. The vision of vast expanses of sun-baked, wind-swept, waterless, forbidding wasteland that seems to occur to most of us the moment we pronounce the word 'desert' is, on the whole, a faithful projection of the reality. But, an important aberration in our thinking that often mars this projection is that the deserts of the world are devoid of any trace of animal life. The situation is, to be sure, quite different and the desert tracts are, if not actually teeming with surface-active animal types, the home of quite a few specialised animal forms and, probably, the last haven of many a vanishing species.
Of all animal types, the rodent would unquestionably be the most numerically heavy tenant of desert lands around the globe. Nature's master plan has guaranteed rodents a place in the sun and they are, apparently, making the most of this arrangement. A study of the biology of deserts would, therefore be, in a large measure, a study of the most important component of the desert biomass, viz. the rodent fauna. It is surprising, however, that with all the recent spate of interest in desert biology, accentuated in no mean measure by the patronage extended by the UNESCO and other world bodies, so little effort has been expended so far in collating the available information on the biology of rodents in desert environments. In our view this has been a major lapse and hence this volume.
Since aridity, rather than great heat or shifting sand masses, truly characterises a desert, both 'hot' and 'cold' (the icy wastes in the Arctic and the Antarctic and at high altitudes on mountains in all latitudes) deserts co-exist on the earth. The scope of this volume has, however, been restricted to 'hot' desert regions. Although the forces causing deserts, or rather aridity, may be several, the present trend of thinking would tend to consider deserts basically as climatic phenomena. But there can also be no two opinion regarding man's role as the principal biotic element in further extending the limits of desert areas. The 'hot' deserts of the world are distributed in two discontinuous belts, one in the Northern Hemisphere and the other in the Southern, more or less centred
XIII
along the Tropic of Cancer and the Tropic of Capricorn, with neither strip deviating by less than 15 degrees or more than 40 degrees from the equator. There are, however, a few patches of desert tract that do not fall into this pattern of distribution.
Although the exact definition of a desert has long been a moot point, the KOPPEN classification of 1918 is generally accepted as the most convenient means of differentiating desert regions from non-desert ones. Temperature and precipitation figures are combined mathematically in the KOPPEN system to establish the boundaries of 'vegetative distributions' for various geographical purposes. KOPPEN defined deserts as having generally high temperatures and under 255 mm (10 in.) of rain annually. According to this estimate, 14 per cent of the earth's 145.6 million sq. km of land are classed as desert. 'KOPPEN steppes', with about 254 to 508 mm (10 to 20 in.) of annual rainfall, and high daily and annual temperature ranges, comprise an additional 14 per cent. Thus the combined desert and steppe areas, all of the arid and semi-arid regions, add up to 41.6 million sq. km. The land areas of the principal deserts of the world are as follows* :
Region
Sahara Desert Australian Desert Arabian Desert Turkestan Desert Great American Desert (includes the Majave, Great Basin, Sonoran, Colorado, Great Salt Lake, Gila and Chihuahuan Deserts of Southwestern North America) Patagonian Desert (Argentina) Thar Desert (India & Pakistan) Kalahari and Namib Deserts (South west Africa) Takla Makan Desert including the Gobi (western China to Mongolia) Iranian Desert Atacama Desert (Peru and Chile)
* Adapted from CLOUDSLEY-THOMPSON, Chapter I in this book.
Square kilomt'l('r (millions)
9.1 3.4 2.6 1.9 1.3
0.67 0.60 0.57
0.52
0.39 0.36
In each of these major deserts of the world, a surprising variety of rodent species has evolved and thrives. Although derived from unrelated stocks, there is often a clearly recognisable resemblance in the morphology, physiology and behaviour of many of these rodent species occupying their respective niches in deserts around the world. Of the two major
XIV
problems of the desert, heat and water, the first can be actively dealt with only at the expense of the second and unavailable commodity. It seems clear now that the rodent's overall strategy of desert survival encompasses both behavioural and physiological adaptations. Much of what follows in this volume is an elaboration of this theme.
As we have emphasised earlier, any worthwhile study of a desert biome must necessarily include a study of its rodent fauna. There seems clearly a case for a proper assessment of the rodent's place in any desert ecosystem. For example, we ought to have quantitative information on the rodent's contributions to the maintenance and aggravation of desertic conditions, its role as a pest of food grains, grasslands and other vegetation and as a carrier of diseases, its relationships with its predator fauna, viz. birds, reptiles and small carnivores of the desert and its susceptibility to chemical and biological control measures. Hopefully, the present volume will provide sufficient background information to generate research interest in these applied aspects of desert rodent biology. We are aware of the deficiencies and limitations of the present volume but such lacunae are, perhaps, inherent in the production of any volume having as multifaceted a scope as this one. Inspite of our best efforts, many species, geographical regions and aspects of research have remained uncovered in this book. Since time was running out fast we decided to plunge into publication without waiting for the manuscripts of several committed authors. Perhaps it will be possible to fill up these gaps in a subsequent edition of the book.
We are thankful to all the contributors to this volume for their excellent cooperation, understanding and patience. We must also thank the Publishers - Dr. W. Junk b.v. and Prof. Dr. J. ILLIES, Editor-in-Chief of the Series for sparing no pains in the production of this volume. Finally, we would like to record our sincere gratitude to Mrs. LAKSHMI 1. PRAKASH for her help at all levels of production of this book.
January 31, 1975 Jodhpur, India
ISHW AR PRAKASH PULAK K. GHOSH
xv
AUTHORS' ADDRESSES
CLOUDSLEy-THOMPSON, j. L., Department of Zoology, Birkbeck College, London, England.
CORBETT, L. K., Division of Wildlife Research, C.S.I.R.O., Alice Springs, N.T., Australia.
DE Vos, A., Food and Agriculture Organisation of the United Nations, Rome. EISENBERG,j. F., National Zoological Park, Smithsonian Institution, Washington D.C.,
U.S.A. FRENCH, N. R., Environmental Resources Centre, Colorado State University, Fort
Collins, Colorado, U.S.A. GAISLER, j., Institute of Zoology, Purkyne University, Brno, Czechoslovakia. GHOBRIAL, L. 1., Middlesex Hospital Medical School, Cleveland Street, London WI,
United Kingdom. GHOSH, P. K., Central Arid Zone Research Institute, jodhpur, India. HAPPOLD, D. C. D., Department of Zoology, University of Ibadan, Ibadan, Nigeria. HARRISON, D. L., Bowerhood House, Sevenoaks, Kent, England. HAWBECKER, A. C., Fresno State College, Fresno, California, U.S.A. JOHNSON, S., Department of Zoology, University of jodhpur, jodhpur, India. JORGENSEN, C. D., Department of Zoology, Brigham Young University, Provo, Utah,
U.S.A. KRAFT, A., Department of Biology, University of New Mexico, Albuquerque, U.S.A. LOBACHEV, V. S., Moscow University, Moscow, U.S.S.R. MACFARLANE, W. V., Waite Agricultural Research Institute, Adelaide, Australia. MARES, M. A., Biology Department, University of Pittsburgh, Pittsburgh Pennsylvania
15260, U.S.A. MISONNE, X., Institut Royal Des Sciences Naturelles de Belgique, Brussels, Belgique. NAUMOV, N., Biological Faculty, Moscow University, Moscow, U.S.S.R. NEWSOME, A. E., Division Of Wildlife Research, CSIRO, Canberra, Australia. NOUR, T. A., Department of Chemistry, University of Alger, Algerie. PETTER, F., Museum National d'Histoire Naturelle, Laboratoire de Zoologie, Mammi
feres et Oiseaux, 55 rue de Buffon, Paris 5e, France. PRAKASH, 1., Central Arid Zone Research Institute, Jodhpur, India. ROSENZWEIG, M. L., Department of Biology, University of New Mexico, Albuquerque,
New Mexico 87106, U.S.A. SCHMIDT-NIELSEN, K., Department of Zoology, Duke University, North Carolina, U.S.A. SMIGEL, BARBARA, Department of Biology, University of New Mexico, Albuquerque,
New Mexico 87106, U.S.A. SMITH, H. D., Department of Zoology, Brigham Young University, Provo, Utah 84601,
U.S.A. TCHERNOV, E., Department of Zoology, The Hebrew University of Israel, jerusalem,
Israel. TURNBULL, PRISCILLA F., Field Museum of Natural History, Chicago, Ill. U.S.A.
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