Kalahari Hyaenas: Comparative Behavioural Ecology of Two Species

316 Pages·1990·8.662 MB·English

by  M. G. L. Mills (auth.)

Kalahari
Hyaenas
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Kalahari
Hyaenas
Comparative Behavioural
Ecology of Two Species
M. G. L. Mills
London
UNWIN HYMAN
Boston Sydney Wellington
©M. G. L. Mills, 1990
Softcover reprint of the hardcover 1st edition 1990
This book is copyright under the Beme Convention. No reproduction
without permission. All rights reserved.
Published by the Academic Division of
Vnwin Hyman Ltd,
15/17 Broadwick Street, London Wl V IFP, UK
Unwin Hyman Inc.,
8 Winchester Place, Winchester, Mass. 01890, USA
Allen & Unwin (Australia) Ltd,
8 Napier Street, North Sydney, NSW 2060, AustraIia
Allen & Unwin (New Zealand) Ltd, in association with the
Port Nicholson Press Ltd,
CompusaIes Building, 75 Ghuznee Street, Wellington 1,
NewZealand
First published in 1990
British Library Cataloguing in Publication Data
Mills,M.G.L.
Kalahari hyaenas.
1. Hyaenas
I. Title 11. Series
599.74'427
ISBN 978-94-015-1103-2 ISBN 978-94-015-1101-8 (eBook)
DOI 10.1007/978-94-015-1101-8
Library of Congress Cataloging in Publication Data
Mills, M. G. L.
KaIahari hyaenas: comparative behaviouraI ecology of two species
IM. G. L. Mills.
p. cm.
Bibliography: p.
Includes index.
1. Brown hyena - KaIahari Desert - Behavior. 2. Spotted hyena
KaIahari Desert - Behavior. 3. Psychology, Comparative. 4. Brown
hyena - KaIahari Desert - Ecology. 5. Spotted hyena - Kalahari
Desert - Ecology. I. Title.
QL737.C24M54 1989
599.74'427-dc20 89-16593
elP
Typeset in 10/12 point Linotron Times
by Nene Phototypesetters Ltd, Northampton
Dedicated to
the memory of Stevie
Contents
Foreword page xiii
Acknowledgements xv
1 Introduction 1
1.1 The study 1
1.2 The study area 3
1.3 Methods 7
1.4 The animals 9
1.5 Summary 12
Statistical tests 13
2 Feeding ecology 14
2.1 Food availability 14
2.2 Brown hyaena diet 19
2.3 Spotted hyaena diet 32
2.4 Diets of the other large carnivores and ecological separation
of the predators 46
2.5 The impact of predation on the prey populations 53
2.6 Relations between hyaenas and other carnivorous animals 55
2.7 Summary 68
Statistical tests 69
3 Comparative foraging and feeding behaviour 71
3.1 Activity patterns and day-time resting sites 71
3.2 Foraging group sizes 75
3.3 The use of senses during foraging 78
3.4 Foraging for vegetable matter, birds' eggs and insects 82
3.5 Brown hyaena hunting behaviour 86
3.6 Spotted hyaena hunting behaviour 93
3.7· Feeding behaviour 119
3.8 Summary 127
Statistical tests 128
4 Social structure and spatial organization 130
4.1 Brown hyaena clans 130
4.2 Spotted hyaena clans 135
x CONTENTS
4.3 Nomadic males 146
4.4 Land tenure system 149
4.5 Factors affecting the sizes of social groups and territories 157
4.6 Summary 165
Statistical tests 165
5 Communication patterns and social interactions 167
5.1 Visual and tactile communications and social interactions 167
5.2 Vocalizations 179
5.3 Scent marking 187
5.4 Summary 204
Statistical tests 205
6 The comparative denning behaviour and development
of cubs 207
6.1 Dens 207
6.2 Development of cubs 214
6.3 Sub-adults 219
6.4 Functional considerations of denning behaviour in the
Hyaenidae 221
6.5 Summary 224
Statistical tests 225
7 The individual in hyaena society 226
7.1 Degrees of relatedness between clan members 226
7.2 Brown hyaena society 227
7.3 Spotted hyaena society 241
7.4 Evolutionary trends in the social systems of the two species 258
7.5 Summary 259
Statistical tests 259
8 Relations between, and management considerations
for, brown hyaenas and spotted hyaenas 261
8.1 Relations between brown hyaenas and spotted hyaenas 261
8.2 Management considerations 269
8.3 Summary 273
Appendix A Common and scientific names of species
mentioned in the text 275
CONTENTS xi
AppendixB Estimated numbers of some ungulates in
the spotted hyaena study area 278
AppendixC Ageing criteria of ungulates based on
eruption of teeth in bottom jaws and
tooth wear 280
AppendixD Methods used to measure territory sizes 282
AppendixE Degrees of relatedness between clan
members 285
References 287
Index 299
Foreword
This beast has a stone in its eye, also called an Yena, which is believed to
make a person able to foresee the future if he keeps it under his tongue.
It is true that if an Yena walks round an animal three times, the animal
cannot move. For this reason they affirm that it has some sort of magic
skill about it. (Translated from Latin, from the twelfth century Bestiary
in the University of Cambridge Library. )
Perhaps it is not merely fortuitous that of all the African animals, the
ancients should have selected the hyaena as a vehicle for magical powers.
Of course, there are solid scientific explanations for this; the animals'
nocturnal habits around human habitation, the consumption of people's
mortal remains, the spectacular hermaphroditic appearance, the uncanny
similarity between the calls of a spotted hyaena and the utterances of
deranged humanity. But apart from all rational explanations for the
strange hold of the animal over people, there is a magic about hyaenas
which can only be understood by those who have watched them for some
time. There is a now growing band of us, who came to the African bush
with all our prejudices, with all that 'common knowledge' about hyaenas
which proved so totally wrong, and who just fell for the spell of animals
which were so totally different.
The hyaena family is the smallest one of the carnivores, with a
membership of only three species of proper hyaenas, and the somewhat
aberrant aardwolf. But many more hyaenas existed in days before our
time; we are only seeing the tail end of hyaena glory. Some were huge,
and, for instance, skulls of over 90 cm long have been found in North
Africa. If only we had been there to see the packs of those huge, probably
rhinoceros-sized animals roaming the plains, driving alongside in our
vehicle to spot the details of their hunting behaviour, and comparing them
with all the smaller species!
On a smaller scale, that is just what Gus Mills did with the present-day
remainder of the hyaenas. The two species which now live alongside each
other in the Kalahari provide a beautiful opportunity for a comparison
between the solitary and the gregarious, between a specialist and an
opportunist. At the same time, it takes some superb fieldcraft to realize
that opportunity in the harsh, almost desert conditions. And fieldcraft is
something which Gus has built into him, something which is not likely to
show itself in his scientific writing, which can only be appreciated when
watching the man in his natural habitat, the Kalahari.
Gus knows the southern Kalahari like a hyaena does, at night and by
day; the endless dune ridges, hundreds of miles of sand, scrub and grass; to
xiv FOREWORD
an outsider it is all the same and intensely hostile. This book will not tell of
the thrills and alarms of the midnight cross-country race, following packs of
hyaenas, taking dune tops at speed in the pitch dark, in the hope that the
other side will be a gentle slope. All this with the one aim of keeping up
with the study, with the hyaena ahead, with its way of staying alive. In this
way Gus, and his wife Margie, put an imJtlense investment, in time and
energy, into the acquisition of background knowledge and skills to get the
observations needed for a book like this.
The study is outstanding, especially in the amount of detailed knowledge
amassed over such a long period, of individual hyaenas and their offspring,
and of the offspring of offspring. The interactions within clans, the
differences between the position of individual brown and spotted hyaenas
within their respective social environments, the social and feeding
strategies of each, all this within the overwhelming Kalahari habitat with its
complicated ecology: Gus has proved it to be a goldmine for the behav
ioural ecologist.
The results of the painstaking nightwork reported here will not be the
last study of hyaenas; certainly, more is to come. But it will be a unique
building stone in our understanding of carnivore behaviour in general, and
of hyaena social organization and ecology in particular. It will help in
designing management strategies for a species which needs protection
rather than persecution, the brown hyaena. And in its comparisons with
spotted hyaenas elsewhere, it will show the fascinating flexibility of the
other species, ostensibly a specialist.
Hyaenas do not have to circle their prey three times in order to catch it;
Gus Mills showed that perhaps the ancients were wrong on that. But
through his stunning pictures, his solid data and interpretations, he also
showed that the general idea of the hyaenas behind those medieval
statements was pretty well right: animals of bewitching attraction, with a
profound contribution to make to behavioural ecology.
Hans Kruuk
Banchory, Scotland