African Ungulates: A Comparative Review of Their Ethology and Behavioral Ecology

311 Pages·1977·6.281 MB·English

by  Dr. Walter Leuthold (auth.)

Zoophysiology and Ecology
Volume 8
Coordinating Editor D.S. Farner
Editors
W S. Hoar B. Hoelldobler H. Langer M. Lindauer
Walter Leuthold
African Ungulates
A Comparative Review of Their Ethology
and Behavioral Ecology
With 55 Figures
Springer-Verlag
Berlin Heidelberg New York 1977
Dr. WALTER LEUTHOLD
Zurich, Switzerland,
formerly Research Zoologist
and Acting Chief Biologist
Kenya National Parks
ISBN-13: 978-3-642-81075-6 e-ISBN-13: 978-3-642-81073-2
DOl: 10.1007/978-3-642-81073-2
Library of Congress Cataloging in Publication Data. Leuthold, Walter, 1940-. African
ungulates. (Zoophysiology and ecology; v. 8). Bibliography: p. Includes index. 1. Un
gulata-Behavior. 2. Ungulata-Africa. 3. Mammals-Behavior. 4. Mammals-Africa.
I. Title. QL737.U4L48.599'.3'045.76.44535.
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© by Springer-Verlag Berlin Heidelberg 1977.
Softcover reprint of the hardcover I st edition 1977
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Preface
This book has been written mostly within sight of wild African
ungulates, at the research center in Tsavo East National Park,
Vo i, Kenya. While this had many positive aspects, there were
also a few drawbacks. The main one of these is the fact that
Voi is not exactly at the hub of scientific activity, even if
we restrict ourselves to African ungulates. Thus, whereas I
had felt initially that I was sufficiently familiar with ethological
work on these animals to write a useful review, it soon became
woefully evident that this assumption was erroneous. Over
the last few years studies on African ungulates have pro
liferated and results are being published in journals almost
all over the world. My location in East Africa was sometimes
less than ideal with respect to access to the most recent literature,
and I depended to a considerable extent on the goodwill of
colleagues in giving or lending me relevant papers. I am
happy to report that I received a great deal of support and
cooperation in this respect. Nevertheless, I may have over
looked some important papers inadvertently; their authors
should not feel slighted by such omissions, which only reflect
availability of literature to me.
Readers may also notice a considerable geographical bias in
favor of East Africa, at the expense of other African regions,
particularly South Africa. This results mainly from my personal
background, for while I have been fortunate enough to live and
work in East Africa for close to ten years, I have not visited
other parts of Africa. On the other hand it is fair to say that,
at least until quite recently, most field work on African ungulates
was carried out in East Africa:
One or two practical points may be noted here: (1) Cross
references to different sections of the text are m~de by means
of the section numbers given in the Table of Contents and the
respective section headings. (2) Scientific names of the ungulates
mentioned in the text are given in the Systematic Index, along
side the English names. (3) The frequent reference to national
parks and other protected areas has necessitated the use of the
abbreviations "NP" for national park and "GR" for game
reserve.
V
It is a pleasure to acknowledge the valuable cooperation and
assistance extended to me by many colleagues who talked or
corresponded with me, sent me their reprints and reports,
or drew my attention to research workers and literature
previously unknown to me. Over the years, I have benefited
considerably from discussions with numerous colleagues,
among whom I wish to mention specifically Drs. R. D. Estes,
P. J. Jarman, J. B. Sale and F. R. Walther. I am grateful to
Prof. J. D. Skinner (Pretoria) for his valuable services with
respect to literature from Southern Africa, and to Messrs.
R. Underwood (Pretoria) and J. C. Hillman (Nairobi), who
made available unpublished material from their studies on
eland. I particularly wish to thank Dr. M. R. Stanley-Price, for
a time my "next-door neighbor" to the east (some 80 km away),
who not only lent me several unpublished theses in his
possession, but very kindly undertook to read the first draft
of this book and made numerous useful suggestions for its
improvement.
The following kindly provided photographs for inclusion in
this book: Mr. 1. Bernegger, Dr. J. H. M. David, Dr. and Mrs.
I. Douglas-Hamilton, Dr. R. D. Estes, Dr. S. C. 1. Joubert,
Prof. and Mrs. H. Klingel, Dr. H. Kruuk, Mr. and Mrs.
D. L. W. Sheldrick. (Photographs not otherwise identified
are· my own). I am also grateful for the use of several of the
splendid drawings by Dr. Fritz Walther. A number of figures
were drawn by my wife Barbara, who also did the first critical
reading of the text. For permission to reproduce figures
previously published elsewhere I am indebted to Messrs.
A. A. Balkema (Cape Town/Rotterdam), Blackwell Scientific
Publications (Oxford), E. J. Brill (Leiden), IUCN (Morges),
P. Parey (Berlin and Hamburg), Institut Scientifique de
Madagascar (Paris). I am also grateful to the Springer-Verlag,
and to Dr. K. F. Springer personally, for the understanding
shown for my problems.
The Trustees and Director of the then Kenya National
Parks permitted me to write this book while being in their
employment as research biologist in Tsavo National Park.
Mr. M. B. Owaga, Secretary at the Tsavo Research Project,
ably undertook the laborious task of typing the final manu
script.
On the more personal level, I wish to thank my wife and
children for their patience and forbearance shown on the many
evenings and week-ends when I was "unavailable".
Perhaps the greatest debt of gratitude lowe to the one who
first introduced me to the fascinating world of African
VI
ungulates and who played an important part in my professional
and personal education; regrettably, he died prematurely, of
a malicious ailment, some months ago. In grateful recognition,
I wish to dedicate this book to the memory of Helmut K.
Buechner.
January, 1977
Voi, Kenya, and
Zurich, Switzerland WALTER LEUTHOLD
VII
Contents
Introduction 1
Part 1
General Background 3
Chapter 1
African Habitats. 5
Chapter 2
The Ungulate Fauna of Africa. 9
Chapter 3
Ethological Research on African Ungulates 12
A. Historical Review. 12
B. Methods of Study 13
Chapter 4
Conservation 16
Part 2
Comparative Review of Nonsocial Behavior. 19
Chapter 5
Physical Maintenance Behavior 21
A. Feeding Behavior. . . . . 21
I. Food Gathering and Ingestion 23
II. Selectivity . . . . . . . . . 25
III. Development of Food Habits During Ontogeny 26
B. Drinking Behavior . . . 28
C. Urination and Defecation . . . . . . 29
I. Postures. . . . . . . . . . . . 30
II. Spatial and Temporal Distribution 32
III. Associated Behavior. 33
IV. Specific Examples. . . . . . . . 34
D. Comfort Behavior. . . . . . . . . . 35
I. General Review of Ungulate Comfort Behavior 36
II. Allogrooming . . . . . . . . . . . . .. 38
IX
Chapter 6
Behavior Related to the Environment. 40
A. Habitat Selection . . . . . . . . 40
B. Space-Related Behavior . . . . . 41
I. The Concept of Home Range . 41
II. Size of Home Range. . . . . 42
III. Internal Anatomy of the Home Range 46
IV. Functional Aspects of Home Range 50
V. Movements and Migrations 51
C. Time-Related Behavior . . 53
I. Daily Activity Patterns 53
II. Annual Patterns . . . 63
D. Space-Time Systems. . . . 64
I. The Daily Space-Time System 64
II. The Annual Space-Time System. 66
E. Influences of Weather on Behavior 67
I. Sunshine and Temperature . 67
II. Wind 68
III. Rain ......... . 69
Chapter 7
Relations with Animals of Other Species 71
A. Positive Relations. . . 71
I. Ungulate-Ungulate 71
II. Ungulate-Other. 72
B. Neutral Relations 74
C. Negative Relations . . 74
I. Ungulate-Ungulate 74
II. Ungulate-Parasite. 76
III. Ungulate-Predator 76
Part 3
Social Behavior 85
Chapter 8
Expression and Communication 87
A. Visual Signals . . 88
B. Auditory Signals . . . 91
I. Vocalizations. . . 91
II. Nonvocal Signals. 93
III. Combined Visual-Auditory Displays 93
C. Olfactory Signals . . . . . . . . . 94
I. Scents Emanating from the Body 94
II. Urine and Feces as Scent-Bearers 95
x
III. Marking with the Products of Scent Glands 96
IV. Interindividual Marking . . 98
V. Functions of Scent-Marking 99
D. Tactile Signals . . . . . . . . 100
Chapter 9
Individual Recognition . . . . . . . . . . . . . . . 102
Chapter 10
Coordination and Cooperation . . . . . . . . . . . 104
Chapter 11
Leadership . . . . . . . . . . . . . . . . . . . . 107
Chapter 12
Agonistic Behavior. 109
A. Introduction . . 109
B. Weapons and Fighting Techniques 111
C. Threat and Intimidation Displays 116
D. Submission and Appeasement 124
E. Systematic Review . . . . . . 126
Chapter 13
Sexual Behavior 139
A. Precopulatory Behavior 140
I. Male Behavior . 140
II. Female Behavior . 145
B. Copulation. . . . . . 149
C. Postcopulatory Behavior. 150
D. Systematic Review 151
E. Functional Aspects 157
Chapter 14
Maternal/filial Behavior 158
A. Prepartum Behavior. 158
B. Parturition. . . . . 161
I. Time of Day . . 161
II. Position and Behavior During Parturition 162
C. Postpartum Behavior . . . 163
D. The Mother-Young Bond. 167
I. Imprinting. . . . . . 167
II. Spatial Aspects. . . . 168
III. The Sensory Basis for Individual Recognition. 171
IV. Suckling and Associated Behavior .... 171
XI
V. The Following Response. . . . . 176
VI. Maternal Defense. . . . . . . . 177
E. Behavioral Development of the Young. 178
F. Weakening and Dissolution of the Mother-Young
Bond . . . . . . 179
G. Functional Aspects . . . . . . . . . . . . . . . 180
Part 4
Social Organization. . . . . . . . . . . . . . . . . 185
Chapter 15
General Patterns in Social Relations 187
A. Gregariousness, Aggregations and Social Gr\,upings . 187
B. Spatial Relationships Between Individuals in Groups. 189
C. Dominance and Territoriality. 190
I. Dominance . 190
II. Territoriality. . . . . . 192
Chapter 16
Types of Social Organization in African Ungulates 195
A. Nonexclusive Home Ranges . . . . . . . . 195
B. Nonexclusive Home Range but Exclusive Home Site. 207
C. Exclusive Home Ranges in Both Sexes. . . . . . . 210
D. Exclusive Home Range in Males Only: Home Range
Coincides with Territory. . . . . . . . . . . . . 213
E. Only Part of Home Range Exclusive, in Males Only:
Mating Territory . . . . . . . . . . . . 216
F. Territory and Absolute Hierarchy Combined . . . . 222
Part 5
Behavior and Ecological Adaptation. . . . . . . . . . 225
Chapter 17
The Influence of Environmental Factors on the Spatial and
Social Organization . . . . . . 227
A. Movements and Home Range 227
B. Social Organization. . . . . 228
I. Sedentary versus Migratory Populations 228
II. Seasonal Changes Within Populations . 230
III. Influences of the Local Environment on Social
Organization. . . . . . . . . 232
1. Vegetation Structure . . . . 232
2. Rainfall and Degree of Aridity 233
3. Population Density. . . . . 234
XII