International Journal of Primatology, Vol 15, No. 4, 1994

Book Review Primate Social Conflict. E d i t e d by William A. Mason and Sally P.

Mendoza, State University of New York Press, Albany, 1993, vi + 419 pp., $59.50 (hardcover), $19.95 (paperback).

Sociobiological theory and, in particular, game theory have advanced our understanding of the ultimate factors that regulate the nature of agonistic behavior between individuals. However, as Mason and Mendoza correctly point out, social conflict in primate groups does not occur among competing evolutionary strategies, but among interacting individuals in a dynamic system. The authors of the 14 chapters in this book explore the ways in which primates resolve social conflicts. The contributions explore the causes and consequences of conflict in primates, and the proximate factors that regulate social conflict. The book begins with an essay by Mason, who takes to task the notion that status acquisition per se is the cause of conflict and the end point of agonistic interactions. Instead, he argues that hierarchies may be the outc o m e of optimizing the n a t u r e of social interactions such that an individual's satisfaction is maximized while, at the same time, negative consequences of the interaction are minimized. Conflicts in species with large multimale-multifemale social groupings are explored in five chapters. Silk provides an excellent integration of the proximate regulation of social conflicts and cooperation among female primates with Wrangham's ecological model of primate social structure. This approach has considerable heuristic utility. Mendoza addresses the role of social conflict in the development of social relationships. Interactions during the first hours of contact in rhesus triads suggest that conflicts are rule-bound and may function to reduce the potential for injurious interactions during group formation. Echoing Mason's entreaties, she concludes that rank acquisition itself is not the purpose of conflict upon groups formation but that the production of differences in rank appears to be the primary goal. De Waal reviews the study of postconflict behavior, with a 639 0164-0291/94/0800-0637507.00/0 0 1994 Plenum Publishing Corporation

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clear discussion of the concerns about methodological issues in the study of these interactions. Itoigawa describes conflicts in relationships among adult male Japanese macaques. Thirty years of continuous observation yields benefits, though the exposition of the main ideas is not as clear as it could be. Finally, Sapolsky's contribution shows that field behavioral endocrinology has come of age and is producing exciting results. Demographic changes produce natural experiments to test relationships between physiology and dominance, and Sapolsky's chapter reveals the complexity of these relationships in the olive baboon. Conflict between parents and offspring is addressed in two chapters. Mason, Long, and Mendoza describe species differences in mother-offspring relationships in rhesus and bonnet macaques. The patterns are consistent with species differences in social behavior, suggesting unique and species-typical social temperaments. Andrews, Sunderland, and Rosenblum's evaluation o f m o t h e r - o f f s p r i n g relationships u n d e r different foraging demands shows that ecological variables modulate mother-offspring interactions. It is surprising that little or no overt mother-offspring conflict is observed under foraging conditions which dramatically reduce the mother's contact with her offspring. Social and reproductive conflicts in Neotropical primates are addressed in chapters by Menzel, Anzenberger, and Abbott. Menzel describes variations of spatial distribution in two groups of captive titi monkeys and argues that species-specific patterns of spacing may be more strongly influenced by social predispositions than by ecological variables (such as the distribution of food). The chapter is data-lean, but the arguments are amenable to empirical testing. Anzenberger reviews his studies on the behavior of marmoset and titi monkeys during confrontations with unfamiliar conspecifics. Similarities in social relationships in the two taxa are produced by different underlying patterns of attraction to and agonism toward unfamiliars. Abbott provides a broad review of the concomitants of reduced reproductive success in subordinate callitrichid primates, as well as a very readable summary of his own work on the physiology of suppression in marmosets. The contribution by Tiger on the biological antecedents of human aggression is full of rich language and pithy analogies. Unfortunately, the contribution does not match the empirical rigor of the rest of the volume. One would hope that a human social psychologist with interests in evolutionary biology would do for t h e study of human social conflict what David Buss has done for the study of human mate choice. The volume ends with an essay by Lyons, who explores the notion that conflict has beneficial consequences and makes comparisons with the literature from h u m a n developmental and cognitive psychology. In the same way that perceptual

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conflict resolution serves as an effective bridge to more sophisticated stages of Piagetian cognitive development, Lyons argues that intra- and interindividual conflict resolution play an important role in the social development of primates. His essay concludes with some challenging questions that are central to modem cognitive ethology and are, as yet, untested. This volume will serve students of primate social behavior extremely well. Mason and Mendoza have done an excellent job of producing a collection of papers that clearly define the methodologies, theoretical issues, and unresolved questions in the study of primate social conflict.

Jeffrey A. French

Department of Psychology University of Nebraska at Omaha Ohaha, Nebraska 68182-0274

International Journal of Primatology, VoL 15, No. 4, 1994

B o o k Review Getting Here. The Story of H u m a n Evolution. By William Howells, Compass Press, Washington, DC, 1993, xiii + 261 pp., $36.00 (hardback), $10.95 (paperback). Howells has provided us with an up-to-date, light- and even-handed overview of the history of human evolution. In these relatively few pages are presented most of the important "facts" (primary data) and appropriate, usually well-chosen examples of how some such data have been, are being, and, sometimes, might better be viewed, evaluated, and tested against other, complementary databases. He appears, at times, to follow Mark Twain's adage that "there's always a hole in theories somewhere, if you look close enough." It is, and will be (for most) a pleasurable, informative, and often humorous read from an investigator and teacher in human evolutionary studies now halfway through his eighth decade. As a young Navy man, Bill Howells' Mankind So Far (1944) accompanied me on my last Pacific cruise, near the end of WWII, and certainly encouraged my budding enthusiasm toward study of the human past. Hopefully, this volume may have similar impact on others early in their education, as well as professionals of many sorts and walks of life who, I trust, evince some real curiosity about humanity's past. There are 19 chapters in all. The initial three are devoted to evolution and its processes, to lower tetrapods and mammals. Three subsequent chapters outline nonhominid primate evolution. Subsequent chapters (7-19) discuss hominid evolutionary history, according to principal taxa and (more recently) geographic areas, and include an abbreviated (Chap. 10) introd u c t i o n to t h e P a l e o l i t h i c , a n d to t a x o n o m y a n d p h y l o g e n e t i c reconstruction, and a final (Chap. 18) overview of the emerging broader perspective of (several) possible phylogenies and of modern human population relationships. In some ways these constitute an updating of his excellent volume, Evolution of the Genus Homo, published two decades earlier, which often had illustrations (fossil crania) of better quality and rendition than those afforded here. 643 0164-0291~3507.00/0

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Howells repeatedly reveals his critical, often skeptical stance, creative insight, and restrained good humor throughout his exposition. Undaunted, he intrepidly invades the briar patches of hominid phylogenetic controversy, emerging unscathed and unflappable, without even a cladistic scratch. He affords the reader an up-to-date phylogram (Fig. 30), following B. A. Wood, without firm commitment to its taxonomic complexities and implications; but in another place (Fig. 65) he presents a "Splitter's delight" schematic of "the formal names (for species and subspecies) within Homo that might currently be considered acceptable by qualified authorities" which he might himself endorse, though skittish to do so overtly. Schemes of regional continuity in contrast to populational (species) replacement (Fig. 69) in the evolution of genus Homo are offered in conjunction with his earlier (1976) delineation of the Candelabra and Noah's Ark hypotheses, respectively; it is explicit (at least in the text) that he is dissuaded by the former and only uneasily comfortable with the latter. Howells has made significant contributions to present (and past) human population relationships though his extensive, multivariate studies of hominid crania. This database (fully published in 1973 and 1989), drawn from almost 30 modern populations and employing an extensive (n = 57) repertoire of traditional and innovative mensurational parameters, is now an invaluable resource for all students of human variability and population affinity, present or past. The consequences of such studies, and of correlative investigations in population genetics, linguistics, and culture history, are thoughtfully interwoven in his discussion of affinities, dispersals, and origins of modern human groups (Chaps. 15-18). Howells explicitly declaims this volume as a textbook but one, instead, "to tell an interested reader not only the known story but also why the anthropologists think what they think and argue as they do." It succeeds very well toward that purpose. Its major fault, if such that may be, is the lack of any bibliographic sources. However, it constitutes a splendid commentary and pithy evaluation of the state of human evolutionary studies from a highly esteemed, broadly experienced, and uncommonly sage contributor. It surely merits a wide readership, and hopefully some of its contents will be stimulus enough for others to pursue resolution of problems still prevalent in a field of major concern to us all. F. Clark Howell

Laboratory for Human Evolutionary Studies Department of Anthropolog), University of California, Berkeley Berkeley, California 94720

International Journal of Primatology, Vol 15, No. 4, 1994

B o o k Review Biology, Rearing, and Care of Young Primates. By James K. Kirkwood and Katherine Stathatos. Oxford University Press, Oxford, UK, 1992, vii + 154 pp., $75.00 (hardback).

This is a practical manual that seeks to help keepers of captive primates maximize the survivorship, health, and reproductive potential of newborn colony members. The book is concerned in particular with infants that must be partly or wholly hand-raised. Whereas rates of neonatal mortality are high for most wild primates, the authors note our opportunity and responsibility to minimize mortality in animals under our care. The work is presented as an attempt to sift through the scientific literature relevant to care and husbandry and to provide a reference text to assist primate keepers of all kinds to develop high management standards and respond appropriately when problems arise with young, captive-born primates. The volume's 150 pages of core text are organized taxonomically: in separate chapters, the authors review information for each of 18 species. Appropriately, for such a selective approach, representatives of most (9/11) families of primates receive treatment: ruffed lemurs (representing Lemuridae), lesser mouse lemurs (Cheirogaleidae), Senegal bushbabies and Bosman's pottos (Lorisidae), western tarsiers (Tarsiidae), common marmosets, cotton-top and golden lion tamarins (CaUitrichidae), owl monkeys and squirrel monkeys (Cebidae), vervet monkeys, rhesus and stumptailed macaques, savanna baboons, proboscis and black-and-white colobus monkeys (Cercopithecidae), lar gibbons (Hylobatidae), and chimpanzees (Pongidae). Each chapter contains information on a fLxed set of topics: status, subspecies, and distribution; sex ratio; social structure; breeding age; longevity; seasonality; gestation; pregnancy diagnosis; birth; litter size; adult weight; neonatal weight; adult diet; adult energy requirements; growth; milk and milk intake; lactation and weaning; feeding; accommodation; infant management notes; physical development; behavioral development; disease and 645 0164-0291/94/0800-0645507.00/0O 1994 Plenum Publishing Corporation

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mortality; preventive medicine; indications for hand-rearing; and reintegration. The topical headings reveal the innovative and most compelling aspect of the authors' approach: primary focus on daily energy requirements, milk composition, and rates of growth. Drawing from Kirk'wood's expertise in the study of mammalian growth, they present at least one growth curve and plots of estimated milk and energy intake for each species treated. These figures are small and the curves appear to be hand-drawn, but the value of presenting such a large set of parallel data is obvious. Accurate estimates of daily energy requirements, of course, are fundamentally important to anyone who would try to save a primate whose mother could not raise it. Beyond the basic data, Kirkwood and Stathatos provide some perspective on the nature of species differences, introducing concepts such as metabolic weight (Kleiber, 1975) and occasionally discussing predictable relationships between physical and behavioral development and between development and overall life history. With its emphasis on growth, the volume should also attract the attention of researchers. Enough data have accumulated so that much behavioral primatology has begun to be conducted within the framework of life history theory. Kirkwood and Stathatos help to promote future research by providing information on early phases of development for diverse taxa in one location. The book has shortcomings, including some problems with selection of species. Ruffed lemurs (Varecia variegata) for example, constitute an odd, specialized lemurid at best, not a good choice to illustrate general management concerns. Due to their nest-building (vs. infant-carrying), extreme frugivory (vs. leaf-eating), and fission-fusion sociality (vs. group cohesion), optimal management for Varecia differs appreciably from that for most other Lemuridae. From data on the relatively nongregarious ruffed and mouse lemurs, the authors conclude that chances of encountering behavioral problems due to hand-rearing are generally low with prosimians. In fact, imprinting on caretakers and difficult reintegration with conspecifics are challenges almost inevitably associated with intensive hand-rearing of the more gregarious lemurs, e.g., Lemur catta, Eulemur macaco, Hapalemur gr/seus, and also Varecia. One might similarly question the selection of Saimiri and Aotus to represent the Cebidae. Two primate families were omitted from consideration due to rarity in captivity, but at least one-fifth of the volume is devoted to taxa of comparably narrow concern. Finally, given the objective of promoting captive conservation, one might expect somewhat greater emphasis on endangered species. Only 3 of the 18 focal taxa are endangered; 12 are not even threatened. Most readers, however, will be most disappointed by the volume's near-complete lack of general discussion. Though the title forecasts inte-

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gration of information on biology and husbandry, one finds only the 18 chapters of restricted focus, a general three-page introduction, a brief index, and a list of brand-name products. To promote infant-rearing with other species or in different circumstances, a reduction of specific, sometimes marginally relevant detail to allow discussion of emergent patterns and principles would be indicated. The only information tabulated in the text is a listing of families represented by the focal species. One imagines, instead, 10 to 12 well-constructed tables with columns of data under several of the current topical headings. The length of text could have been halved, making pages available for integration. The volume's organization and lack of general discussion preclude complete s u ~ with the authors' objective of saving keepers' time. Those caring for a taxon must read the relevant sections of every chapter to garner leads on solutions to problems they are encountering. Thirst may be discovered as a potential indicator of pregnancy stage in the squirrel monkey chapter. One also finds reference there to a study that showed an effect of protein level in the diet on rate of abortion. The importance of provisioning in a way that ensures intake of vitamin D3 is mentioned in the chapter on common marmosets. Use of vaginal lavages or smears to detect menstruation is first mentioned halfway through the book (vervet monkey), and techniques for training infants to self-feed are under stumptailed macaque. Readers must determine for themselves the likely generality of phenomena, problems, and solutions. How commonly does dietary composition influence ovulation number? How does growth rate generally vary in relation to differences between replacement milk and species-typical milk? Tamarins and squirrel monkeys experience intriguing difficulties with neonatal thermoregulation. Are allomothers potentially important as additional sources of heat? Might this illuminate mothers' and infants' typical approaches to relations with other adults? Do young primates paired with mature members of the opposite sex generally show accelerated maturation? How do species-typical body size, parenting behavior, dispersal patterns, and social structure interact with these phenomena? The bibliographies for most species are limited, and many citations on natural historical phenomena are dated or refer to secondary literature. The authors seem to have foregone direct contact with leading researchers and facilities, which prevented them from including completely up-to-date information with regard to either biology or management in some cases. Amidst the current debate and hard work over the difficult issues of animal welfare, regulation of research, and environmental enrichment, the absence of any perspective on matters of concern in these areas is unfortunate. When immediate physiological needs are easy to meet, but eventual social needs are not, how should the decision whether to hand-raise an aban-

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doned infant generally be approached? Humane killing and high management standards are advocated, but not defined. What areas most often require flexibility in developing standards of care for different taxa? At different institutions? Other omissions were also surprising. Despite the strong focus on growth, there is no mention of its episodic and compensatory aspects. Most young animals, at least in certain phases, show a fascinating potential to achieve normal size, via rapid growth, following a period of difficulty. There is also evidence that infant primates grow faster or longer if they are somehow handicapped, e.g., born late in the season or as the smaller twin. After bursts of rapid growth, individuals commonly undergo periods of relatively slow growth. Values of the immense literature on human growth (Falkner and Tanner, 1986) are not mentioned in the text. Seasonality of reproduction is specifically covered, but no discussion is provided of mechanisms and their implications. The word "photoperiod" does not occur in the book, yet potentially pervasive control of life histories by this phenomenon makes it a central aspect of housing and care. Lemurs, for example, have recently been discovered to grow seasonally (Pereira, 1993). Parallel phenomena concern especially those caring for any of the seasonally-breeding primates. Despite these difficulties, my overriding memory of this book is its standardized illustration of a large data set on the growth rates and energy intakes of diverse infant primates, all in relation to the available data on milk composition. This is very useful. It will help the staffs of primate facilities internationally to advance their management practices. Greater numbers of healthy infant primates should survive, which will be necessary for their species' persistence to be safeguarded most effectively. This was the authors' stated primary objective, and they met it. All keepers of primates would benefit from reading their book. REFERENCES Kleiber, M. (1975). The Fire of Life, Krieger, New York. Falkner, F. and Tanner, J. M. (eds.) (1986). Human Growth, VoL 1, 2nd ed., Plenum, New York. Pereira, M. E. (1993). Seasonal adjustment of growth rate and adult body weight in ringtailed lemurs. In Kappeler, P. M. and Ganzhorn, J. U. (eds.), Lemur Social Systems and Their Ecological Basis, Plenum, New York, pp. 205-221.

Michael E. Pereira Duke University Primate Center 3705 Erwin Road Durham, North Carolina 27705

International Journal of Primatology, VoL 15, No. 4, 1994

B O O K REVIEW Primates of the Americas. Strategies for Conservation and Sustained Use

in Biomedical Research. Edited by Primo Ar~imbulo III, Filomeno Encarnaci6n, Jaime Estupifi~in, Hugo SamaraS, Charles R. Watson, and Richard E. Weller. Batelle Press, Columbus, Ohio, 1993, xv + 314 pp., $34.95 (paperback).

This book is the result of a conference held at Batelle Laboratories in Seattle, Washington, in October 1990. It contains 12 papers. One is a general overview of Neotropical primate conservation. The rest deal with natural populations and sustained cropping, primate reproduction under laboratory conditions, conservation of nonhuman primates and public health, cooperative programs of the Pan-American Health Organization and the World Health Organization, and the utilization of nonhuman primates. There is also a theoretical paper on the economics of primate conservation and another (one page long) on the role of national primatological programs in preserving primates in Latin America. Two papers deal with intercountry cooperation for development of national primatological programs and on the value of "debt for nature" swaps for preserving primates. There is a paper dealing with aspects of wildlife management in Brazil and another about the participation by communities in the protection of primate natural habitats in Bolivia. An important value of the book for Latin American readers is that it contains a version in Spanish of each of the papers and of the technical minutes of the meeting. Most of the emphasis in the book, however, is on the achievements of the Peruvian Primatological Project, leaving one with the impression either that the organizers want this project to become a model to be followed by other Latin-American countries or that the document is a justification for the continued existence of the project. While the book refers to the primates of the Americas, there is little in it that reflects the title. It would have been useful if short contributions from at each of the other Latin American countries--Argentina, Paraguay, 649 0164-0291/94/0800-0649507.00/09 1994 Plenum Publishing Coqmration

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Guatemala, Colombia, Mexico that participated in the meeting had been included regarding specific intracountry information on conservation and use of nonhuman primates in biomedical research. The inclusion of a paper on the "debt for nature" option for primate conservation, at the end of the book, makes it seem to be the "solution" for conservation that needs to be followed by Latin American countries. The final paper is an outline of very general strategies and actions that the authors called "Regional Plan of-Action in Pfimatology for the Americas," but no indication is given about how such an outline bonds with the regional conservation action plans of the Primate Specialist Group of the IUCN for Latin America. The book presents little new information and it seems more like a disconnected collection of papers. At $35.00, it is hardly accessible for Latin American researchers and students.

Alejandro Estrada Estacion de Biologia "Los Tuxtlas" Instituto de Biologia---UNAM Apartado Postal 1 76 San Andres Tuxtla, Veracruz, Mexico