PRIMATE ECOLOGY AND SOCIAL STRUCTURE
Volume 1: Lorises, Lemurs and Tarsiers
Robert W. Sussman
Department of Anthropology
Washington University
1999
PEARSON CUSTOM PUBLISHING
CONTENTS
PREFACE...ix
PART I...1
Chapter 1 Ecology: General Principles...3
The Ecosystem...3
The Components of Ecosystems...5
Energy Transfer Within Ecosystems...6
The Pyramid of Energy Transfer...8
Ecological Interactions...10
Interactions Between Abiotic Components and Producers...10
Biomes and the Process of Succession...12
Some Features of a Climax Community...13
Leibig's "Law" of the Minimum...14
Interactions Between Populations of Consumers...20
Interactions Between Animals Occupying Different Trophic
Levels: Predator-Prey Interactions...20
Interactions Between Animals at the Same Trophic Level:
Competition and Coexistence...22
Coevolution...25
Social Groups, Social Structure and Social Organization...27
Bibliography...31
Chapter 2 The Taxonomy and Evolution of Primates...39
Principles of Animal Taxonomy...39
What Is a Primate?...45
The Origin of Primates: The Arboreal Theory of Primate
Evolution...49
The Origin of Primates: A Paleontological Approach...50
The Origin of Primates: Terminal Branch Feeding
Insectivores...51
Were the Earliest Primates Visually Oriented Predators?...53
A New Theory of Primate origins: Coevolution with
Angiosperms...55
Bibliography...63
PART II THE ECOLOGY OF PROSIMIANS...71
Introduction...71
Chapter 3 Lorisiformes...73
Introduction...73
The Lorisids of Gabon: A Case Study...74
Spatial Localization...75
Diet...80
Defense Against Predation...83
The Ecology of Other Species of Lorisiformes...85
The Social Behavior of the Lorisiformes...90
Summary of the Ecology and Social Behavior of
Lorisiformes...99
Bibliography...101
Chapter 4 The Nocturnal Lemuriformes...107
Introduction...107
The Ecology of the Nocturnal Lemuriformes-A Case Study in
Western Madagascar...114
The Ecology of Other Species of Nocturnal Lemuriformes...124
Predation...130
The Social Organization of the Nocturnal Lemuriformes...139
Reproduction...138
Summary of the Ecology and Social Behavior of Nocturnal
Lemuriformes...139
Bibliography...141
Chapter 5 The Ecology of the Diurnal Lemuriformes...149
Introduction to Nocturnal Vs. Diurnal Primate
Adaptations...149
Introduction to the Ecology of the Diurnal Lemuriformes...152
The Ecology of the Diurnal Lemurs of Western Madagascar...154
A Study of Two Sympatric Species of Lemur...154
Vertical and Horizontal Ranging
Patterns and Intergroup Spacing...157
Diet and Foraging Behavior...164
Activity Cycles...167
Social Structure and Organization...167
Summary...172
Propithecus verreauxi: One Species Living in Two Different
Habitats...173
Methodology...173
Vertical and Horizontal Ranging...174
Diet and Foraging Behavior...177
Activity Cycles...179
Social Structure and organization...179
Other Diurnal Species of the Occidental Vegetation Zone...182
The Ecology of the Diurnal Lemuriformes of the Oriental
Zone...192
The Frugivorous Lemurs of the Eastern Rain Forest...192
Folivorous Lemurs of the Eastern Rain Forest...200
Bamboo and Grass Eaters of the Eastern Forests...206
Predation...209
Summary of the Ecology and Social Behavior of Diurnal
Lemuriformes...210
Archaic Coevolutionary Pollinator Relationships...212
Infanticide in Lemur catta...213
Female Dominance, Reproductive Stress, Seasonality, and Adult
Sex Ratios...214
Bibliography...215
Chapter 6 Tarsiiformes...231
Introduction...231
The Ecology of Tarsius...236
Locomotion and Habitat Preferences...236
Activity Cycles and Sleeping Sites...240
Diet and Feeding Behavior...241
Predation and Competition...242
The Social Behavior of Tarsiiformes...243
Acoustic and Olfactory Communication...246
Reproduction and Infant Development...247
Summary of the Ecology and Behavior of the Tarsiiformes...249
Bibliography...250
Chapter 7 Summary Chapter: Patterns of Variability, Diversity
and Conservation Among Prosimians...257
Activity Cycles...258
Habitat and Locomotion...258
Diet...260
Predation...262
Social Organization...263
Conservation...266
Bibliography...268
INDEX...271
Preface
In this book I review the literature on free-ranging nonhuman
primates. Initially I attempt to characterize these animals
as they exist in their least disturbed state. When relevant,
behavior in disturbed situations and captivity is compared to
that occurring in more undisturbed contexts. By doing this,
it is possible to gain a better understanding of the reasons
primates behave as they do and the ways they fit into the
communities in which they live. I realize this is an
idealistic approach, since few localities inhabited by
primates escape some level of human disturbance.
Research in field primatology during the late 195Os and early
196Os was mainly descriptive natural history, with few
comparative and quantitative, or problem-oriented studies.
However, by the 197Os, primatology moved into a problem-
oriented phase (see Sussman 1979, Smuts et al. 1987, Smuts
and Fedigan in press). Problem-oriented studies, focus mainly
on determining relationships between behavior and morphology,
ecology and social structure, and community interactions, and
were a major component of field primatology during the 197Os
and 198Os. More recently, many field primatologists are
attempting to formulate and test theories that have developed
out of classical sociobiology; such as those related to kin
selection, inclusive fitness, reciprocal altruism, and sexual
selection (see below). In many cases, the theories and
studies are elegant and elaborate but the data are meager
(e.g., see Casti 1989, Allen 1993, Sussman in press).
The philosopher of science, F.S.C. Northrop (1965) suggests
that any healthy scientific discipline goes through three
stages during its development. The first stage involves the
analysis of the problem. The second is a descriptive natural
history phase. Finally, there is a stage of postulationally
prescribed theory, in which fundamental theories are tested.
Although there is a movement in field primatology towards
this final stage of inquiry, in fact, even today many primate
species have not been studied in detail, and the most studied
species normally are known only from a few localities. The
range of variation in the behavior and ecology of most free-
ranging primates is still unknown, and basic natural history
remains a necessary component of the subdiscipline. As stated
by Northrup (1965:37-38):
In fact, if one proceeds immediately to the deductively
formulated type of scientific theory which is appropriate to
the third stage of inquiry, before one has passed through the
natural history type of science with its inductive Baconian
method appropriate to the second stage, the result inevitably
is immature, half-baked, dogmatic and for the most part
worthless theory.
Although, in the following chapters, I will discuss many of
the theories currently popular in primatology, these cautions
must be born in mind.
Primates make up only a small part of the communities in
which they are found. In fact, plants and insects typically
are the most diverse and numerous organisms in these
communities. In order to understand primate behavior it is
necessary to study the ecological communities and ecosystems
in which they live, the broad biogeographical context in
which they are found, and their phylogenetic history
(Schoener 1988, Brooks and Mclennan 1991, Losos 1996). In the
first chapter, I discuss the concept of an ecosystem and
describe some of the general types of interactions that occur
between the various components of ecosystems. Primates are
highly social animals and the way animals space themselves in
the environment and reproduce is related to other aspects of
their ecology. The study of these relationships is referred
to as socioecology. In the final pages of Chapter 1, I
discuss some of the terms and theories used in socioecology.
Many of these concepts are further explored in later
chapters.
In Chapter 2, I discuss the taxonomy of primates, their
morphological characteristics, and the origins and
evolutionary history of the earliest primates. After these
two introductory chapters, I review the literature on the
galagos and lorises (Chapter 3), nocturnal Malagasy lemurs
(Chapter 4), diurnal Malagasy lemurs (Chapter 5), and
tarsiers (Chapter 6). The general organization of each of
these review chapters is similar in order to facilitate
comparisons. In the final chapter, cross-taxonomic
comparisons are made of each of the topics discussed in the
review chapters. In a second volume, I will review the
literature on New World monkeys, and in a third volume that
on old World monkeys and apes. I have used earlier versions
of this book for the past 2O years in teaching courses on
primate ecology and social behavior. Revisions of earlier
versions have benefited from the input of many students over
this time. A few of my colleagues have read various versions
of the book in its entirety and I appreciate their comments
and suggestions. These include John Buettner-Janusch, Paul
Garber, Terry Gleason, Charles Hildebolt, Jane Phillips-
Conroy, Alison Richard, Ian Tattersall, and Mildred Trotter.
Besides the above, for specific chapters, specialists in the
topics covered have generously agreed to read the volume and
offer their suggestions. Jonathon Losos and Peter Raven
offered their input on Chapter 1, Glenn Conroy, Tab Rasmussen
and Peter Raven on Chapter 2, Leanne Nash, Kimberly Nekaris
and Tab Rasmussen on Chapter 3, Sylvia Atsalis on Chapter 4,
and Myron Shekelle on Chapter 6. I thank all of these people
and 8 anonymous reviewers for the assistance although I take
responsibility for the final product. l also thank those who
provided photographs and Terry Gleason for his assistance in
getting this volume into final form. Finally, I thank Brett
Nachman for copy editing and Wayne Spohr and Jerry Brennan of
Pearson Education for their
assistance.
REFERENCES
Allen, G.E. 1993. In Search of the Grail: Biology, Social
Science, and the Meaning of Human Nature. Contemp. Psych.
38:455-458
Brooks, D.R., Mclennan, D.A. 1991. Historical Ecology:
Examining Phylogenetic Components of Community Evolution. Pp.
267-28O. In Species Diversity in Ecological Communities:
Historical and Geographic Perspectives. R.E. Ricklefs; D.
Schluter, Eds., Chicago: University of Chicago.
Casti, J.L. 1989. Paradigms Lost New York, William Morrow.
Losos, J.B. 1996. Phylogenetic Perspectives on Community
Ecology. Ecology 77:1344-1354.
Northrop, F.S.C. 1965. The Logic of the Sciences and
Humanities. Cleveland, Meridian.
Schoener, T.W. 1988. Ecological Interactions. Pp. 255-297. In
Analytical Biogeography: An Integrated Approach to the Study
of Animal and Plant Distribution. A.A. Myers; P.S. Giller,
Eds., London, Chapman and Hall.
Smuts, B.B., Cheney, D.L., Seyfarth, R.M., Wrangham, R.W.,
Strnhsaker, T.T., Eds.
1987. Primate Societies. Chicago, University of Chicago.
Smuts, B.B., Fedigan, L.M., Eds. In press. Primate
Encounters. Animals, Scientists
and Science. Chicago, University of Chicago.
Sussman, R.W. 1979. Primate Ecology. Problem oriented Field
Studies. New York,
Wiley.
Sussman, R.W. In press. The Piltdown Man: Father of American
Field Primatology.
In Primate Encounters: Animals, Scientists and Science. B.B.
Smuts; L.M. Fediaan. Eds.. Chicago. Universitv of Chicavo.
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