Primate Ecology: Studies of Feeding and Ranging Behavior in Lemurs, Monkey and Apes

627 Pages·1977·13.13 MB·English

by  T.H. Clutton-Brock (Eds.)

Primate Ecology:
Studies of feeding and ranging
behaviour in
lemurs, monkeys and apes
Edited by
Τ. H. CLUTTON-BROCK
School of Biological Sciences, University of Sussex,
Brighton, England
1977
ACADEMIC PRESS
London · New York · San Francisco
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ACADEMIC PRESS INC. (LONDON) LTD
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London NW1
US Edition published by
ACADEMIC PRESS INC.
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New York, New York 10003
Copyright © 1977 by ACADEMIC PRESS INC. (LONDON) LTD
All Rights Reserved
No part of this book may be reproduced in any form by photostat, microfilm, or
any other means, without permission from the publishers
Library of Congress Catalog Card Number: 77-77368
ISBN: 0-12-176850-3
Printed in Great Britain by Robert MacLehose and Company Limited
Printers to the University of Glasgow
Contributors
DAVID J. CHIVERS Sub-Department of Veterinary Anatomy, University
of Cambridge, Tennis Court Road, Cambridge CB2 1QS, England
Τ. H. CLUTTON-BROCK School of Biological Sciences, University of
Sussex, Biology Building, Falmer, Brighton BJV1 9QG, Sussex, England
R. I. M. DUNBAR Department of Psychology, University of Bristol, 8-10
Berkeley Square, Bristol 8, England
D. FOSSEY Karisoke Research Centre, B.P. 105, Ruhengeri, Rwanda
ALAN G. GOODALL Biology Department, Paisley College of Technology,
High Street, Paisley, Strathclyde, Renfrewshire PA 1 2BE, Scotland
A. H. HARCOURT Sub-Department of Animal Behaviour, University of
Cambridge, Madingley, Cambridge CB3 8AA, England
PAUL H. HARVEY School of Biological Sciences, University of Sussex,
Biology Building, Falmer, Brighton BN1 9QG, Sussex, England
C. M. HLADIK Museum National d'Histoire Naturelle, Equipe de Recherche
sur les Prosimiens, Laboratoire d'Ideologie Generale, 4 Avenue du Petit Chateau,
91800 Brunoy, France
DOROTHY B. KLEIN Department of Ζοο1ο&> University of Alberta,
Edmonton, Alberta T6G 2E1, Canada
LEWIS L. KLEIN Department of ^oology, University of Alberta, Edmonton,
Alberta T6G 2E1, Canada
WARREN G. KINZEY Department of Anthropology, The City College of
The City University of New York, New York NT10031, USA
D. G. LINDBURG Department of Anthropology, University of California,
Los Angeles, California 90024, USA
ν
vi CONTRIBUTORS
Zoological
JOHN F. OATES New York Society, 630 Fifth Avenue, New
York 20, USA
J. I. POLLOCK Department of Anthropology, University College London,
Gower Street, London WC1E 6B T, England
A. RICHARD Department of Anthropology, Yale University, Box 2114,
New Haven, Connecticut 06520, USA
PETER S. RODMAN Department of Anthropology, University of California,
Davis, California 95616, USA
C. C. SMITH Division of Biology, Kansas State University, Manhattan,
Kansas 60506, USA
R. W. SUSSMAN Department of Anthropology, Washington University,
St Louis, Missouri 63130, USA
PETER WASER Department of Biological Sciences, Purdue University, West
Lafayette, Indiana 47907, USA
R. W. WRANGHAM Sub-Department of Animal Behaviour, University of
Cambridge, Madingley, Cambridge CB3 8AA, England
Preface
After reading its title, the ecologist with little experience of primate
field research may be surprised at the contents of this book. All the 17
field studies which it includes are concerned with the ecology or
behaviour of particular social groups of primates rather than of whole
populations. None of them touches on population dynamics in any
depth and only two investigate energetic aspects of feeding behaviour.
Instead, they describe behavioural aspects of ecology: activity pattern­
ing, food selection and ranging behaviour.
Differences between ecological research on primates and research
on other vertebrates have developed for two reasons. First, interest in
primate ecology has been stimulated by investigation of the adaptive
significance of social systems. Early attempts to investigate their
functional aspects of social behaviour relied on correlations between
particular kinds of social system and gross ecological variables such as
habitat or diet type (e.g. Crook and Gartlan, 1966). Subsequent
critiques (Struhsaker, 1969; Clutton-Brock, 1974a; S. A. Altmann,
1974; Wilson, 1975) pointed out that social behaviour varies widely
between species allocated to the same ecological category and
challenged the view that interspecific differences were closely related
to ecological variation at this level. This, in turn, led to the develop­
ment of detailed research in those aspects of primate ecology most
likely to reveal the adaptive significance of differences in social
behaviour: food selection, the availability and dispersion of supplies,
ranging behaviour, variation in grouping patterns and antipredator
behaviour.
A second, and perhaps more important, reason for the divergence
between primate studies and those of most other vertebrates, is the
difference in accessibility of ecological information. Most primates
cannot be trapped with ease; it is often both impracticable and un­
desirable to consider killing adequate samples of animals; and the
majority of species have relatively long lifespans. Difficulties are en­
hanced by the fact that only small numbers of individuals can usually
be sampled since unhabituated animals can rarely be observed
vii
viii PREFACE
consistently. Consequently, it is seldom possible to collect the kind of
information required for detailed research on population dynamics.
In contrast, because individual animals can usually be recognized
and their behaviour followed (if intermittently) over considerable
time periods, it is often possible to examine a range of questions not
accessible to most studies of other vertebrates. For example, it is
frequently feasible to assess interindividual variation in food selection,
to describe sequences of food choice and to measure ranging patterns.
While it is important that future studies should investigate more
traditional ecological questions, the main contribution to vertebrate
ecology that primate studies can provide may lie primarily in those
areas of behavioural ecology to which the observation conditions are
best suited.
Between 1972 and 1974, a regular discussion group of primate
field workers developed in England. We met for one-day workshop
sessions on particular aspects of primate ecology or social organiza­
tion—on food selection, on ranging behaviour, on social structure
and on grouping patterns. Our primary interest lay in the functional
significance of interspecific and intraspecific variation in aspects of
behaviour and ecology. Attempts to construct or test generalizations
about variation at either level required comparison of the results of
different studies, and a regular complaint at our meetings was the lack
of comparability between studies. Though many of us had completed
similar work on different species, our interests varied and it was usually
impossible to make as wide a range of comparisons as the data could
have allowed. At one of these meetings, the idea arose of producing a
symposium on primate ecology, and it was agreed that we should draw
up an outline of the topics which an ideal chapter should cover and
invite selected workers in the field to write similar chapters on different
species. We did this, and the final list of topics included feeding heights
and sites, activity patterns, food selection (by part and species), home
range size and utilization, day range length and group size. For all these
measures, we wished to compare the ways in which they varied between
populations, between times of day, between sex/age categories and
between species. Each author was invited to prepare a chapter which
covered as many of these topics as possible. Though none of the 17 field
study chapters has covered the whole range of topics, the book allows
comparison of most topics across a considerable range of species.
As in previous studies, accurate comparisons are complicated by
methodological differences between studies. Sampling methods,
sample distributions, study duration and analytical techniques all
vary widely and past experience shows that such variation can some-
PREFACE ix
times produce major discrepancies between estimates (see Appendix I).
At first, it seemed possible that these differences would obscure
important relationships and might invalidate quantitative comparisons.
Two points are relevant to this problem. First, methodological variation
is more likely to affect absolute values than relative ones, so that even
where the former cannot be compared, it may still be possible to
compare intraspecific trends. Second, the evidence shows that where
firm predictions can be made about relationships between different
aspects of feeding and ranging behaviour at an interspecific level,
significant associations can be demonstrated in most cases where a
reasonably large sample of species can be used (see Milton and May,
1976; Clutton-Brock and Harvey, in press). We believe that this
justifies attempts at broad comparisons, though results must be
interpreted with great care (see Chapter 19).
An alternative view, expressed both at our meeting and elsewhere,
is that comparisons should not be made until measuring techniques
have been standardized. Not only does the empirical evidence indicate
that this is an unduly pessimistic opinion but there is another reason
why this position is impracticable. Although a greater degree of
standardization will probably emerge in future studies, differences in
observation conditions and in the focus and duration of field studies
will always be sufficient to prevent full standardization of recording
techniques. Consequently, quantitative comparisons between species
will always be limited to aspects of behaviour and ecology where
differences are sufficiently robust to override the effects of varying
methodology.
The first 17 chapters are arranged in the taxonomic order of the
species concerned. The final two chapters survey some of the generali­
zations emerging from comparison of inter- and intraspecific differ­
ences in feeding and ranging behaviour. The aim of these is to suggest
areas of particular interest where research can be usefully developed
rather than to provide definitive answers concerning interrelationships
between behaviour and ecology.
King's College, Cambridge Τ. H. CLUTTON-BROCK
May 1977
Acknowledgements
I am extremely grateful to Paul Greenwood for editorial assistance and
for compiling the Subject Index; to H. A. Clutton-Brock for help in
proof-checking; to Anthony Watkinson and Anne Matthews of
Academic Press for their encouragement, enthusiasm and tolerance, and
to the Director of the Nature Conservancy (Scotland), Martin Ball
and Jeremy Barr for permitting the contributors to hold a workshop
meeting to discuss generalizations arising from studies of different
species in Kinloch Castle, Isle of Rhum.
χ
1
Feeding Behaviour of
Lemur catta and Lemur fulvus
R. W. S U S S M AN
Department of Anthropology, Washington University, St Louis, Missouri,
USA
1. Introduction . . . . . . . . . .. 1
2. Study areas . . . . . . . . . .. 4
2.1. Antserananomby . . . . . . . .. 4
2.2. Tongobato 9
2.3. Berenty 10
3. Sampling methods . . . . . . . . .. 11
3.1. General methodology. . . . . . . . . 11
3.2. Data collection on feeding behaviour . . . . .. 12
4. Feeding behaviour . . . . . . . . .. 13
4.1. Feeding techniques . . . . . . . . . 13
4.2. Feeding heights and feeding sites . . . . . . . 15
4.3. Diurnal activity patterns and percentage of time spent feeding . . 17
4.4. Species composition of the diet . . . . . .. 20
4.5. Parts of plants eaten . . . . . . . .. 26
4.6. Drinking 28
5. Feeding behaviour and social behaviour . . . . . . 29
6. Predation . . . . . . . . . . . 31
7. Summary and discussion . . . . . . . . . 31
Acknowledgements . . . . . . . . .. 36
1. INTRODUCTION
In this paper I will describe the diet and foraging behaviour of Lemur
catta and Lemurfulvus (Figs 1 and 2). These species were studied between
September 1969 and November 1970, and filmed during July and August
1974. The focus of the study was to compare the behaviour and ecology
2 R. W. SUSSMAN
FIG. 1. Lemur catta.