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The Socioecology of Primate Groups J. Terborgh; C. H. Janson Annual Review of Ecology and Systematics, Vol. 17 (1986), 111-136. Stable URL: bttp//links jstor.org/sici?sic!=0066-4162%281986%2917%3C 111%3ATSOPG%3E2,0.CO%3B2-K Annual Review of Ecology and Systematics is currently published by Annual Reviews ‘Your use of the ISTOR archive indicates your acceptance of JSTOR’s Terms and Conditions of Use, available at hhup:/www.jstororg/about/terms.hml. JSTOR’s Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use ofthis work. Publisher contact information may be obtained at hup:/www jstor-org/journals/annrevs.huml Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the sereen or printed page of such transmission, STOR is an independent not-for-profit organization dedicated to creating and preserving a digital archive of scholarly journals, For more information regarding JSTOR, please contact support @jstor.org. hupslwwwjstor.org/ Fri May 6 12:48:19 2005, Copyright © 1986 by Annual Reviews Inc. All rights reserved THE SOCIOECOLOGY OF PRIMATE GROUPS J. Terborgh Department of Biology, Princeton University, Princeton, New Jersey 08548 C. H. Janson Department of Ecology and Evolution, State University of New York, Stony Brook, New York 11794 INTRODUCTION Of all vertebrates, primates are among the most amenable to study in the wild. ‘They are mostly large diurnal creatures that can readily be habituated to close ‘observation, often to the point that the observer can be spatially integrated into the group under study. Consequently, primate behavior and feeding can be observed in extraordinary detail. Variation in facial features permits individual recognition in many species; this opens the door to studies of the ‘ontogeny of behavior, dominance structure of groups, dispersal, and even lifetime reproductive success. Sometimes individuals or whole groups can be ‘captured repeatedly, making possible precise measurements of growth rates, tooth wear, etc, and the taking of fecal and blood samples for the analysis of parasite loads and genetic relationships. Few nonprimates offer such a multi- Plicity of advantages to the student of animals in the wild. Within the past two years there have appeared a plethora of volumes devoted entirely or in part to the socioecology of primates; these books review the subject far more thoroughly than is possible in the space allotted here (9, 60a, 87, 122, 129, 138). We thus do not attempt a broad treatment of the subject but instead focus on the development of theory pertaining to the possible selective links between environmental variables and the social orga- nization of primate groups. This subject is touched upon in all the works ‘mentioned above, but in none of them is it given the in-depth treatment we think is due um (0066-4 162/86/1120-0111$02.00 112 TERBORGH & JANSON We begin by briefly tracing the historical development of ideas pertinent to ‘our central theme. This recapitulation offers not simply a perspective on the later discussion but a splendid example of how science progresses through a succession of phases and fads. We then delve in greater depth into several recent attempts to enunciate the possible polarity of cause and effect rela- tionships between environmental variables and primate social variables. Special attention is given to the most recently proposed hypothesis for the adaptive basis of primate groups. Finally, we review empirical evidence relevant to this and other proposed hypotheses of primate group organization ‘and examine, to the extent possible with existing data, the predictions and corollaries of these hypotheses. As implied in our ttle, the review focuses on the primate literature with only passing references to papers outside this area HISTORICAL BACKGROUND ‘The quest to understand the adaptive basis of the varied social systems of primates has been a major impetus in primatology since its emergence as a discipline. Even before the frst field studies of wild primates in the 1930s and 1940s, Zuckerman (176) proposed on the basis of observations of captive baboons and other species that Sexual bonds provided the social glue that promoted the cohesiveness of primate groups. A decade later, following his Pioneering studies of wild howler, spider, and rhesus monkeys, orangutans, and gibbons, Carpenter (10) rejected Zuckerman’s thesis, pointing out that the ‘composition of wild primate groups remains stable even in species in which sexual activity is infrequent or seasonally limited. The First Phase: The Classificatory Approach ‘The study of primate social groups then became a discipline without a theory ‘until the 1960s when seminal papers on the adaptive origins of avian mating systems by two ornithologists, Crook (36) and Orians (111), brought home to primatologists the value of comparative analysis. However, the avian models did not seem to apply to the situation of most primates: More than 80% of bird species are monogamous, while fewer than 15% of primates are; relatively few birds live in cohesive social groups; multimale and harem groups are practically unknown in birds, while leks are unknown in primates; and pethaps most critically, the rationale for understanding avian social systems was based on control of resources (territories, nesting sites, etc), while control of resources was not an obvious component of primate social systems. Nonetheless, the idea that sense could be made out of the welter of existing primate social systems by casting them in a comparative framework was contagious. Crook & Gartlan (39) followed with the first serious attempt to organize primate social systems in such a framework in @ paper that provided an

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