Table Of ContentAdaptations of Desert Organisms
Edited by J.1. Cloudsley-Thompson
Springer
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Volumes published in the series
Ecophysiology of the Camelidae Physiological Ecology of North
and Desert Ruminants American Desert Plants
By R.T. Wilson (1989) By S.D. Smith, R.K. Manson, and
J.E. Anderson (1997)
Ecophysiology of Desert Arthropods
and Reptiles Ecophysiology of Small Desert Mammals
By J.1. Cloudsley-Thompson (1991) By A.A. Degen (1997)
Plant Nutrients in Desert Environments Homeostasis in Desert Reptiles
By A. Day and K. Ludeke (1993) By S.D. Bradshaw (1997)
Seed Germination in Desert Plants Ecophysiology of Amphibians
By Y. Gutterman (1993) Inhabiting Xeric Environments
By M.R. Warburg (1997)
Behavioural Adaptations of Desert
Animals Avian Desert Predators
By G. Costa (1995) By W.E. Cook (1997)
Invertebrates in Hot and Cold Arid Carnivore Ecology in Arid Lands
Environments By J. du P. Bothma (1998)
By 1. S0mme (1995)
Ecophysiology of Economic Plants
Energetics of Desert Invertebrates in Arid and Semi-Arid Lands
By H. Heatwole (1996) By G.E. Wickens (1998)
Ecophysiology of Desert Birds Arthropods of Mediterranean-Type
By G.1. Maclean (1996) Ecosystems
By G.P. Stamou (1998)
Plants of Desert Dunes
By A. Danin (1996) Dispersal Biology of Desert Plants
By K. van Rheede van Oudtshoorn and
Biotic Interactions in Arid Lands M.W. van Rooyen (1999)
By J.1. Cloudsley-Thompson (1996)
Spatial Ecology of Desert Rodent Com
Structure-Function Relations munities
of Warm Desert Plants By G.!. Shenbrot, B.R. Krasnov and K.A.
By A.C. Gibson (1996) Rogovin (1999)
Georgy I. Shenbrot Boris R. Krasnov Konstantin A. Rogovin
Spatial Ecology
of Desert Rodent
Communities
With 124 Figures and 38 Tables
Springer
.~~.
D~GeorgyI.Shenbrot
Dr. Boris R. Krasnov
Ramon Science Center
Jacob Blaustein Institute for Desert Research
Ben-Gurion University of the Negev
P.O. Box 194
Mizpe Ramon 80600, Isarel
Dr. Konstantin A. Rogovin
Severtzov Institute of Ecology and Evolution
Russian Academy of Science
Leninsky prospect 33
Moscow, Russian Federation
Cover illustration: Sekeetamys calurus (front cover) and Acomys rus
satus (front and back cover). Photographs taken by Georgy Shenbrot
ISSN 1430-9432
ISBN 978-3-642-64224-1 Springer-Verlag Berlin Heidelberg New York
Library of Congress Cataloging-in-Publication Data
Shenbrot, Georgy I., 1950- Spatial ecology of desert rodent communities / Georgy I.
Shenbrot, Boris R. Krasnov, Konstantin A. Rogovin. p. cm. -- (Adaptations of desert
organisms, ISSN 1430-9432) Includes bibliographical references (p. ) and index.
ISBN -13 :978-3-642-64224-1 e-ISBN-13 :978-3-642-60023-4
DO I: 10.1007/978-3-642-60023-4
1. Rodents--Ecology. 2. Rodents--Ges>graphical distribution.
3. Desert animals--Ecology. 4. Desert animals--Geographical distribution. I. Krasnov,
Boris R., 1955-. II. Rogovin, Konstantin A., 1955-, III. Series. QL737.R6S46 1999 599.35
1756--dc21
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In memory of our teacher, the late Professor Nikolai Naumov
Preface
The authors of this volume were fortunate in being introduced to
zoological studies by their teachers at the Department of Zoology
and Comparative Anatomy of Terrestrial Vertebrates, Moscow State
University. This department has a long tradition of field animal re
search in various life zones. In 1954, Professor Nikolai Naumov was
appointed head of the department. Under his leadership, the depart
ment began intensive ecological studies and has become the alma
mater of several generations of Russian animal ecologists. Another
stimulus for rodent ecology research in the department was created
by the problem of the epidemiology of plague and leishmaniasis in
arid zones, which necessitated detailed studies of rodents, the main
reservoirs of these diseases. Our teachers include the noted Russian
zoologists, namely N.P. Naumov, V.G. Heptner, l.A. Shilov, V.S. Loba
chev, N.N. Vorontsov, L.N. Eliseev, V.E. Flint, l.M. Gromov, V.V. Kuche
ruk, V.E. Sokolov and S.A. Shilova. Almost all of them were connected
in one way or another with the department and we have learned
much from them.
Georgy Shenbrot began his scientific career studying the taxo
nomy and ecology of jerboas. Decades of experience with this desert
taxon stimulated his interest in the general rules governing the orga
nization and functioning of communities of desert rodents. In his
research, he has spent much time in the different arid regions of the
world, from the Gobi to Chihuahuan deserts. Boris Krasnov studied
the behavioral mechanisms that influence the spatial structure of
rodent populations in different landscapes, from the tundra and arc
tic shore of the Chukchii Peninsula to the rainforests of south Viet
nam. His interest in the spatial organization of rodent populations
led him to study the spatial organization of their communities. Kon
stantin Rogovin began his acquaintance with desert animals by
studying the ethology of jerboas and later broadened his scope to
include the behavioral adaptations of desert rodents in general. Thus,
the individual experiences of these authors are complementary. This
interaction has been strengthened by the fact that they have been fri-
VIII Preface
ends since their student days (Krasnov and Rogovin even earlier).
Shenbrot and Rogovin collaborated for 7 years at the Institute of Ani
mal Ecology and Evolutionary Morphology, at the Russian Academy
of Science in Moscow, Russia (now the Institute of Ecology and Evo
lution Problems). For the last 7 years, Shenbrot and Krasnov have
worked together at the Ramon Science Center. This small scientific
unit is located in the heart of the Negev Desert of Israel and is affil
iated with the Jacob Blaustein Institute for Desert Research, Ben
Gurion University of the Negev.
The ideas elucidated in this book have been discussed over the
years with our colleagues. These include (in chronological and geo
graphical order) L.Y. Borkin, I.Y. Pavlinov, D.V. Semenov, A.V. Surov,
1.M. Fokin, A.M. Gilyarov, Y.A. Puzachenko, V.V. Ivanitsky, E.N. Panov,
E.G. Potapova, V.S. Gromov,A.V. Chabovsky, S.V. Popov (Russia), V.M.
Mazin (Kazakhstan), O.V. Mitropolsky (Uzbekistan), J.H. Brown, E.J.
Heske, D.A. Kelt, J.S. Brown (USA), D.W. Morris (Canada), B.J. Fox,
C.R. Dickman (Australia), P. Marquet (Chile), D. Ward, Z. Abramsky,
U. Safriel, A.A. Degen, M. Kam, B.P. Kotler, 1.S. Khokhlova, T. Dayan, Y.
Yom-Tov (Israel), M. and A. Stubbe (Germany), 1. Prakash, M. Idris
(India), G. Diaz (Argentina), J. Nel (South Africa). Technical assis
tance in data collection was provided by (in chronological order) G.
Kulikova, N. Bodyak, 1. Bar-Sagi, and M. Laniado.
We thank John Cloudsley-Thompson for inviting us to write this
book and for taking on the gargantuan task of editing it. His editing
was most helpful and we owe him much for clarifying our statements.
We also thank David Ward, the Director of the Ramon Science Cen
ter. Dave encouraged us to write this book. He made invaluable com
ments on the manuscript and worked hard on improving our English
prose.
Financial support during the writing of this book was provided by
the Ministry of Science of Israel, the Ministry of New Immigrant
Absorption of Israel and the Local Council of Mizpe Ramon (for
Georgy Shenbrot and Boris Krasnov) and by the Russian Academy of
Science and the Russian Foundation of Basic Researches (for Kon
stantin Rogovin).
Ramon Science Center Dr. Georgy Shenbrot
JaQob Blaustein Institute for Desert Research Dr. Boris Krasnov
Ben-Gurion University of the Negev
January 1999
Contents
1 Introduction .................................. .
2 Deserts of the World . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
2.1 Common Features. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
2.2 Regional Features ............................... 6
3 Composition of the Rodent Faunas
of Different Desert Regions ............. . . . . . . . . . . 25
3.1 Biogeographic Classification Using Rodents
as a Key Taxon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
3.2 Rodent Faunas of the World Deserts . . . . . . . . . . . . . . . . 32
4 Geographic Patterns of Faunistic Composition . . . . . . . 63
4.1 Taxonomic Composition ......................... 63
4.2 Taxonomic Diversity. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
4.3 Species Richness ................................ 74
5 Inter-Desert Comparisons of Rodent Faunas. . .. . .. . . 87
5.1 Climatic, Substrate and Vegetation Features Important
for Rodents .................................... 87
5.2 Distribution of Body Sizes Within Regional Faunas ... 94
5.3 Distribution of Locomotory Modes
Within Regional Faunas. . . . . . . . . . . . . . . . . . . . . . . . . . 102
5.4 Distribution of Feeding Modes
Within Regional Faunas. . . . . . . . . . . . . . . . . . . . . . . . . . 110
x Contents
6 Life Forms of Desert Rodents and Convergent Evolution
Among Species of Different Faunas ................ 125
6.1 Life Forms of Desert Rodents. . . . . . . . . . . . . . . . . . . . . . 125
6.2 Convergent Evolution Among Species
of Different Faunas .............................. 143
7 Species-Habitat Relationships
in Desert Environments . . . . . . . . . . . . . . . . . . . . . . . . . . 151
7.1 Methodology of Analysis: Niche Concept
and Description of Niche Dimensions .............. 151
7.2 Habitat Structure and Types of Habitat Specialization 153
7.3 Morphological Correlates of Habitat Specialization ... 172
7.4 Habitat Specialists and Habitat Generalists in Deserts 181
7.5 Habitat Selection as a Mechanism of Species Distribution 193
8 Spatial Organization of Desert Rodent Communities. . 203
8.1 Definitions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203
8.2 The Qualitative Approach to the Analysis
of Community Composition ...................... 204
8.3 Multivariate Description of Spatial Structure. . . . . . . . . 208
8.4 Environmental Factors Structuring a Community. . . . . 210
8.5 Guild Structure and Factors Determining It . . . . . . . . . . 216
8.6 Spatial Patterns of Species Diversity
in the Rodent Communities of Different Deserts. . . . . . 228
8.7 The Role of Interspecific Competition
in Community Structure ......................... 233
8.8 Temporal Dynamics of Spatial Structure ............ 238
9 Conclusions and Perspectives for the Future . . . . . . . . . 243
9.1 Historical and Ecological Components
of Spatial Structure .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 243
9.2 Ecological Concepts and Comparative Approach. . . . .. 244
9.3 The Lack of Empirical Data ....................... 245
References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 247
Appendix: List of Rodent Species Considered
to be Desert-Dwellers ............................ 279
Subject Index ........................................ 287
CHAPTER 1
Introduction
Rodents are conspicuous and important components of the desert biome.
Their populations are frequently great and many of them inhabit large areas.
They also provide the diet for numerous carnivorous and scavenging species.
Furthermore, they provide habitats for many vertebrate and invertebrate ani
mals as a consequence of their burrowing activity. Additionally, they regulate
the aeration and water regime of the soil, and provide a suitable environment
for the estivation and early development of the seeds of numerous plants.
Many general concepts in modern community and behavioral ecology are
based on desert rodents as a main model. The study of spatial structure is one
of the most common approaches to the community ecology of terrestrial ver
tebrates. Comparative descriptive studies of vertebrate, particularly rodent,
communities are essential because they are free of many of the limitations
inherent in experimental studies (Schoener 1983, 1986; Brown et al. 1986;
Diamond 1986; Price 1986; Dueser et al. 1989; Gurevitch et al.1992). They are
also partly free of the difficulties that arise in the construction of null models
designed to test specific hypotheses based on field observations (Bowers and
Brown 1982; Harvey et al. 1983; Simberloff 1983; Gilpin and Diamond 1984;
Connor and Simberloff 1984; Strong 1986; Wiens 1989). Comparisons of com
munity spatial organization in different regions with similar environments,
such as the different deserts of the world, can aid in understanding which
patterns are universal and which are determined by the unique taxonomic
composition and/or evolutionary history of a community. Comparisons of
communities occurring in more or less similar environments, but possessing
different histories and taxa, can provide information about general directions
in the evolution of community organization, as well as identifying the speci
fic features of each community.
Conservation of biodiversity is impossible without understanding the
rules of the natural patterns of species distribution within the landscape.
Understanding the response of species assemblages to environmental factors
has long been a central aim of community ecology (e.g. Arrhenius 1921; Elton
and Miller 1954). Few would deny that animal species live in a characteristic
range of habitats; within this range they are most abundant around their
environmental optima. Species abundance, as well as the composition of bio-
