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This review appeared in Volume 2 (1) of The Semiotic Review of Books.

Chimpanzees' Lives

by Anne Zeller

Through a Window. By Jane Goodall. 1990. Houghton Mufflin Co. Boston ISNB 0-395-50081-8. Pp. 268

East of the Mountains of the Moon. By Michael P. Ghiglieri 1987. Free Press, Collier Macmillan, New York ISBN 0-02-911580-9. Pp. 315.

Metaphors are an eloquent evocation of meaning, and as such are important to semioticians. Goodall's metaphor in Through the Window is very much intended to reverberate with the phrase "The eyes are windows to the soul". Her recent publication on the chimpanzees of the Gombe Stream Reserve takes up where In the Shadow of Man left off in bringing to our attention the lives, social relationships and deaths of her study population. She speaks clearly in the first chapter on the multiple functions of "windows", as areas of access to knowledge, vision, sense, interpretation and belief. Of all animals chimpanzees should be the most transparent to human observation since we are so closely related to them. Through observing chimpanzees many primatologists feel that we can gain a better understanding of the nature of animals and of where the human/animal boundary lies. Before Goodall began her epic research project in 1960 animals were not permitted the human attributes of self recognition, planning, tool use, deception, or a host of other behaviours which are now recognized, at least in chimpanzees. This level of complex behaviour, and the ramifications it has for human understanding of the nature of our own behaviour patterns Is only gradually making an impact on how we view other animals. The fact that there is now a large body of carefully gathered data to support interpretations of mental complexity in free ranging chimpanzees is in a large measure due to Goodall's work.

In this record of her research, Goodall lets us see more of her own life, her own intellectual evolution, and commitment to the study of our closest relatives whereas many of her earlier works focus intently on chimpanzees life, this book gives us glimpses of why the author was able to see things which were hidden from the better educated, more channelled vision of those who had ideas firmly in mind before they went out to observe. By reversing this process and observing in the absence of theory Goodall opened a whole new vista of what capabilities chimpanzees might possess. In this work she extends her observation on free living chimpanzees by references to research on captive ones and compares for example, the tool use of termiting chimps, with the rake users reported by Wolfgang Kohier back in 1924. Also, although all the wild chimpanzees communicate in their own code, she sees levels of complexity and individual use patterns. Long continued practice and familiarity allow her to recognize individual voices and call patterns, which although similar, are used in different circumstances and send distinguishable messages. This level of complexity, and learning required to master the code allows her to accept the idea of chimps using American sign language as a medium of communication with humans. They have high levels of memory, spatial abstraction, affective input, and the ability to suppress inappropriate gestures which are necessary components in utilizing a code as abstract as human symbol system. The adaptive source for this level of intellectual ability is not only the chimpanzee communication system, but their needs for a spatial map of resources, long term memories and individual recognition, because of the fission/fusion nature of chimpanzee society, as well as the need to socially manipulate and even deceive others when they are too strong to be intimidated. Chimpanzees have repeatedly been observed suppressing appropriate but inadvisable vocalizations and facial gestures in both captivity and the wild in order to mislead other individuals. As Goodall says, you can study the form, results and adaptability of complex mental activities in captivity, but the actual function and adaptive nature of such behaviours is revealed through research in their natural habitat. As examples supporting this claim Goodall's descriptions of hunting behaviour and cooperation clearly indicate the complex levels of communication which occur among chimpanzees. One animal may spot an infant monkey from 30 or 40 yards away and begin a silent stalk. Other chimpanzees deploy themselves in a circle covering possible exits in trees or on the ground. Sometimes the young prey is saved if adults of its own species see the chimpanzees and rush in, alerting the intended victim in time, and distracting the hunters. In other cases the prey often leaps away from the hunter right into the hands and teeth of another chimpanzee. Goblin, a young adult male, was seen to catch two infant monkeys which he gave to his mother and sister and then get a third one for himself. The ability to suppress vocalizations of excitement, to interpret the suddenly silent and stealthy movements of another as an indication of the presence of prey, and to act accordingly, and the willingness to share the results all distinguish chimpanzee hunting as a conceptual as well as physical behaviour. The level of anticipation, of self restraint, and understanding of the consequences of actions revealed in hunting behaviour also shows up in the social manipulations which individuals undertake to gain status in the group. Figan showed the ability to learn through observation and take advantage of opportunities. Mike, a previous alpha male had made a dramatic rise in rank by terrifying other animals with the noise made by empty kerosene tins. Figan, even as an adolescent was seen practising with the abandoned tins when others were not in sight. After the tins were removed Figan gradually developed a strong relationship with his older brother Faben, who he had dominated after Faben lost the use of one arm due to polio. Eventually Figan and Faben joined forces to fight and Intimidate other young adult males, who then became nervous around the brothers. Figan also began to challenge the alpha male Mike, who was aging, by ignoring his displays. Mike was eventually defeated, and Figan, by gaining the assistance of other chimpanzees was able to move into the power vacuum. He would not have been able to hold the highest rank against larger older males without the constant assistance of Faben his brother, who backed him up at every challenge. Eventually the other males became so intimidated by the pair that Figan alone was able to win the bluff displays and fights, thus securing the highest rank in the group. He maintained this position while his brother was still in the troop but shortly after Faben disappeared the other male chimpanzees began to challenge him. However, he retained his position for nearly four years until he injured his hand and was successfully challenged by a young male who had been his constant companion during most of the time. The whole pattern of learning through observation, manipulating social situations, and taking advantage of another's misfortunes are applied to the various life situations in which chimpanzees find themselves.

The chapters of this book explore the chimpanzee mind, the life histories of several individuals, including their rise to power, the intricacies of sexual negotiation, the progress of inter community violence, the learning of hunting, and the intense personal relationships between individuals, revealed in chapters on mother-son dynamics, mother-offspring relationships, and the devastating effects of orphaning on young animals. The last three chapters and two appendices change the tone somewhat by referring to the state of chimpanzees in captivity and their long term chances of survival in the wild. These are matters of immediate concern, since during her 30 years of study the chimpanzee population of Tanzania has dropped to 1/4 of its previous levels, and this in a country which does not hunt the animals for food, or keep them for research purposes. Other areas of Africa have lost even larger proportions of their chimpanzee populations due to hunting, capture for sale, and deforestation for purposes of cultivation in overcrowded countries.

The importance of this book, for those who are not interested In the copious statistical and scientific material in her 1986 work, The Chimpanzees of Gombe, is the opportunity to meet non humans, who hope, love, hate and fear, who plan and deceive, who engage in power struggles, who adopt their orphans in some cases and cannibalize infants in others. These are animals with very complex mental and social behaviours. The material is not experimental, and the observations sound anecdotal, until you realize that they are backed by thousands of hours of observation and hundreds of examples which Goodall has winnowed into a book that reads simply, but is full of profound implications for how we view ourselves and other animals. Probably the most influential aspect of Goodall's work is the individualizing of animals, whose life histories are followed in such detail that the reader can distinguish the personalities of one animal from another. Many of these individuals have become familiar entities to primatologists and lay readers alike and their lives as individuals emphasize the lessons in social relationships that Goodall is endeavouring to present.

Michael Ghiglieri's book on chimpanzees, titled East of the Mountains of the Moon is aimed at the educated public, in the same genre as Goodall's but with a more pronounced and emphasized theoretical bent. It is an account of the personal side of research on Ugandan Chimpanzees in the Kibale Nature Reserve between 1976 and 1981. Rather than being a statistical and scientific report of findings (this was published in 1984 as The Chimpanzees of the Kibale Forest) it is a record of the difficulties and small personal triumphs of trying to study unprovisioned, unhabituated chimpanzees in a region called Ngogo in the centre of the Kibale Forest Reserve. Ghiglieri's study was specifically designed to ascertain whether the sociobiological explanation of male chimpanzees cooperating with each other due to kin relations required for communal protection of their range from non-group chimpanzees could be supported by observations. In order to provide evidence to support this hypothesis he was interested in observing the relations of males with each other, and with females, their ranging, feeding and socializing patterns, and their response to stranger. In addition, the relations of females with each other and with males, and their ranging, feeding and socializing patterns were useful in suggesting comparisons between chimpanzees related by kin bonds (males) and those affiliated by alternative social factors (females). The particular focus of investigation was whether the intergroup aggression seen in Goodall's Tanzanian chimpanzees was a normative, genetically based behaviour, or an abnormal one, due to particular circumstances. Several people, including Tom Strubsaker who invited Ghiglieri to come to this research station, initially felt that the problems involved in habituating chimpanzees without provisioning them at all were too difficult to overcome. It was important to Ghiglieri not to provision the animals because there was mounting evidence from Goodall's and Nishida's work on Tanzanian chimpanzees that even limited provisioning could alter social relations and ecological influences on behaviour. Eventually he began to stake out the large fig trees where both chimpanzees and monkeys came to eat ripe fruit and gradually over a period of months came to habituate and know over 40 chimpanzees of a community numbering about 55 individuals.

First, he studied pant hoot vocalizations, in a effort to discover who gave them and when in order to ascertain what their function might be. Like Goodall, he discovered that calls by individual animals could be distinguished by humans, and thus presumably by chimpanzees, and that many calls were given in the presence of large fruit resources. This evidence of altruism reinforced Ghiglieri's preconceived interpretation of the nature of social bonds among chimpanzees.

His results suggested that males in the Kibale forest spent most time feeding and travelling together and made most of the long distance vocalizations. Females preferred to travel and eat with females, which is a different pattern than the one observed at Gombe and the Mahale Mountains where females appeared to prefer to socialize with males. The list of reasons Ghiglieri advances to explain why the Kibale females preferred to associate with females did not seem to explain this difference in behaviour, because he did not indicate the way that these reasons could account for one pattern of behaviour at Kibale and the opposite in the Tanzanian chimpanzees.

Other sources of information drawn in to support the hypothesis of kin bonding as a basis for territoriality are the author's interpretations of meeting Ethiopian tribes people on a rafting trip down the Omo river, and of lion behaviour observed in Queen Elizabeth Park in Uganda. Two of the three Ethiopian tribes he encountered along the Omo river were under some level of population pressure from increased growth and from owning live stock in an area rife with tse-tse fly. The middle tribe had retained an older lifestyle without cattle, but were being pushed from both sides, by groups requiring more land for grazing and safe waterholes away from the fly and crocodile infested river.

"As I learned more about the strife between the Bodi and Mursi, I saw more parallels between these warring tribes and aggressive communities of chimpanzees reported from Tanzania. The war between the Mursi and Bodi was primarily over territory...Sociobiologically the ecological and psychological rationales for war between the Bodi and Mursi or war between two communities of chimpanzees was basically identical, as was their inescapable dependence on solidarity and mutual support among themselves" (pg. 140).

Similar conclusions were drawn from observations of lions, in which larger groups of males were more successful in defending a hunting territory and thus providing females with a place to hunt for themselves and their cubs. The need for adolescent male cubs to move away from their natal pride in a group, and to maintain what Ghiglieri calls kinship solidarity was considered vital, if this particular group of males was ever to force its way into the ownership of a territory. Yet, several females of a pride usually give birth, and several males breed, so there is no certainty of genetic relationship between these group dwelling males. A point which emphasizes this is the observation that 44% of the newly forming male groups were in fact, non-kin coalitions which worked together to establish and maintain a territory. Thus, while the value of territory maintenance by male lions is not under dispute, and provides them with sexual access to the females living in the territory, the argument that it is kin relationship which holds them together is only valid a little more than half the time. It is more probable that a group of male chimpanzees is more closely related than a group of male lions, but the necessity for a genetic basis for such cooperative territory holding groups does not seem to be strengthened by the lion example. In fact the level of social interaction, and the effort put into the development and maintenance of social relations seem to argue more strongly for the need to establish relationships with little underlying basis, but great reproductive value, than it does for the existence of a genetic predisposition underlying the occurrence of same sex support in chimpanzees.

In order to assess the strength of cohesion Ghiglieri counted occurrences of males travelling together, responding to pant hoots, feeding together and grooming together. To test whether grooming had a predominantly social or hygienic function, he divided the body into self accessible and self inaccessible parts and observed the frequency of grooming in each category by other animals. By far the largest proportion of grooming, 95.7% was done on the less accessible parts, which the author interpreted as an exchange of services by animals, a fairly pragmatic function, rather than a social cement. Yet the need to exchange grooming,--or at least the desire for it--seems to dominate social relations, and as Ghiglieri says "fosters a sense of wellbeing and, sometimes, near euphoria" (pg. 112).

In other words it seems to be a behaviour with a high level of social importance. One major factor which distinguishes males from females is the greater amount of time males spent travelling each day. The males tend to cover more of the range, and thus serve as notice that it is occupied, as well as alerting other animals to undiscovered sources of fruit. One major ecological factor emphasized by Ghiglieri is the unlikely nature of chimpanzee diet. Larger animals tend to eat more common food, but chimpanzees prefer ripe fruit, which is a scarce and patchily distributed resource. When trees are in fruit chimpanzees disperse themselves in it as widely as possible, to reduce competition, and females seldom alert males to its presence, although they will follow the males pant hoots indicating a fruit tree. Single female chimpanzees may be forced out of a fruit tree by several male colobus monkeys acting together, and since Ghiglieri estimated that there were almost 200 monkeys (of 5 species) per chimpanzee, as well as non primate fruit eaters, the supply never lasted long. He questioned whether this use of resources was actually competition, in the sense of affecting the chimpanzees reproductive potential, and concluded that wild chimpanzees do not mature as quickly as captive ones, and thus food scarcity does affect them. Also, red colobus will eat figs at a much greener stage than chimpanzees, and eat only a fraction of what they pick, which reduces the level of resources available for chimpanzees. Thus females pregnant or with young, usually must travel some distance every day to find enough to eat, and do not have extra energy to keep up with the males on their longer daily range. Therefore females tend to travel together, eat together (so they have less competition than from males, and some protection from monkeys) and groom together. If they are related their positive behaviours are only apparent altruism according to sociobiological theory, because the behaviours are reinforced by genetic bonds of inclusive fitness (inclusive fitness is the genetic benefits to your genome of aiding relations). Friendship is a positive relationship between unrelated individuals.

However Triver suggests that friendship and mutual support also have a genetic basis labelled reciprocal altruism. If there is a genetic basis for such behaviour, the factor of interest is why it is only expressed towards certain animals. There is a distinct difference between the behaviours of animals who have "friend" relationships and those who are ignored or who are actively antagonistic. At any rate, females do tend to have preferred relationships and they are reinforced twice as often as positive social interactions between males and females in the Kibale chimpanzees.

Ghiglieri concludes that communities are made up of both sexes in complementary roles, which serve to foster the chimpanzees reproductive strategies. He sees the males as a kin group breeding as a communal unit, even though they would prefer to monopolize estrus females. He makes this comment about their preference, even though the most common mating pattern he reports is serial mating by a number of males, which he interprets as subjugating the priority of monogamous mating in order to maintain male group bonds and solidarity. Probably due to the size of the reserve Ghiglieri did not actually observe the territorial conflict suggested in his hypothesis, but argues that several of the supportive factors indicating the importance of male relationships outlined above such as males preference for males as grooming partners, males travelling together, tolerating each other's mating activities, long distance contact communication, food calls, and minimum of dominance interactions over food all support the interpretation that males have a cohesive social unit focussed on improving reproductive success of the whole kin related group. Ghiglieri repeatedly suggests that male territorial defense is important, even if rare in order to provide protection of the fruit resources for female and young, as well as safety and sexual access to females. He ties this interpretation to human behaviour quite explicitly.

"Primitive hunting and gathering societies the world over-exhibit, or once exhibited, patterns of exogamy, hunting and sometimes ecology, plus territorial defense and warfare basically identical in form and function to that of chimpanzees."(pg. 259).

The last chapter and epilogue of the book is concerned with the threat of poaching and deforestation to the rapidly declining population of chimpanzees in the wild. During the course of the book several times he refers to the inroads of poaching on primates as well as on large game in general. There is also a chapter on forest destruction around the world, with specific references to Africa, especially in light of political exploitation in Uganda after Idi Amin took power and set many conservation laws aside.

The book provides a fascinating picture of chimpanzee life in a different situation than Gombe reserve and emphasizes the importance of studying more than one population of a species in order to gain insights into behaviour. Particularly with chimpanzees who have such a large learned component to their behaviour the differences between the Ngogo population and the Kangawara group, only 10 miles away exemplify the potential for tradition and culture to influence chimpanzee behaviour. Grooming styles, foraging patterns, association units, and use of the resource base all differ between groups as close as 10 miles apart, as well as more distant populations. Aspects of behaviourial differences are also indicated in the comparison of Ugandan chimpanzees with those of Goodall's study population in Tanzania. Some of these differences may also be due to differing ecological situations, such as the frequency of fruit trees or human cutting of forests which attenuates ranges and removes the potential for contact with outside groups. In comparing these two books a number of differences are evident. Ghiglieri fits his research into a more explicit political, economic and theoretical framework than does Goodall. He has chapters on the forest, on anti-poaching patrols, and on his adventures in other parts of Africa which contribute to his conclusions, as well as referring to the effects of political turmoil in Uganda on wildlife and forest conservation. Goodall also refers to the effects of the raid by Zairian rebels on her research station, which resulted in closing it to foreign students and the changes this made in her data collection, but continues to focus more on the information derived from the chimpanzees themselves, than the difficulties encountered in collecting it. This difference in focus is, to me, the most pervasive distinguishing characteristic of these two books.Goodall lets the lives of the animals speak directly to the reader, although she does give some clues for interpreting them, while Ghiglieri speaks for the animals in an attempt to tell us what they are doing and why. Due to his sociobiological theoretical basis (in the Preface he refers to the book as a case study in sociobiology) he presents a rather one sided interpretation of data as I have tried to suggest in several places. For instance, he quotes E.O. Wilson's definition of dominance and uses it without referring to the current level of controversy surrounding the term and its actual function (see Strum 1987). His book also contains no pictures of the chimpanzees or the habitat which would have aided the less experienced reader to visualize the animals and their setting. The portraits of individuals in Goodall's book help to emphasize their personalities and the influence that this important factor has on their social relationships. Ghiglieri names and describes his animals but they are not as distinctively individualized as those who have a visual reality to the reader. Of course Goodall has the advantage of 30 years of field experience and photo archives compiled by trained photographers to help amass a visual resource on which to draw.

Their differing visions of chimpanzees are also exemplified in the chapter titles chosen. Goodall's book has chapter headings of Figan's Rise, Change, Power, War, Sons and Mothers, Gilka, Melissa, Sex, Love and Our Shame. These provide quite a difference ambience than Ghiglieri's chapter titles of Killer Apes, the Creature that Ate the Forest, It's a Jungle Out There, and Return of the Killer Apes. In fact, "killer apes" is a common way in which he refers to chimpanzees throughout the book, even though he sees very few episodes which might be hunting behaviour and no inter-troop conflict at all. This predisposition to emphasize the theoretical approach that he is investigating seems questionable, even though he supports his hypothesis indirectly to his own satisfaction. Although reproductive fitness is undoubtedly the primary goal of evolution, in animals as complex and intelligent as chimpanzees (and humans) the potential for individual solutions to common problems has the potential to modify the expression of behaviours even if they do have an underlying genetic base. Rather than enter a debate about the potential for sociobiological theory to explain chimpanzee behaviour, let me suggest that these two books complement each other in extending our understanding of the complex factors underlying chimpanzee lives.

References

Ghiglieri, M. 1984. The Chimpanzees of Kibale Forest. New York: Columbia University Press.

Goodall, J. 1986. The Chimpanzees of Gombe Patterns of Behaviour. Cambridge MA: Belknap.

Strum, S.C. 1987. Almost Human. New York: Norton Press.

vanLawick Goodall J. 1971. In the Shadow of Man. London: Collins.

Anne Zeller is a primatologist who is an Associate Professor in the Department of Anthropology at the University of Waterloo (Canada). She has recently been conducting fieldwork in Indonesia (Kalimantan) on the adult-infant social relations in Macaca fascicularis (crab-eating macaque). She is the author of a study of visual and vocal communication in Macaca sylvanus in Primates: recherches actuelles. J.J. Roeder and J.R. Anderson (eds.) Paris: Masson (1990). Among her other publications are: "Comparison of component patterns in threatening and friendly gestures in Macaca sylvanus of Gibraltar", in Current perspectives in Primate Social Dynamics. D.M. Taub and F.A. King (eds.) New York: Van Nostrand Reinhold Press, 1986. (487-504), "Primate Facial Gestures: A Study of Communication". Intemational Journal of Human Communication (1980) 13,4: 565 606, and "Speaking of Clever Apes". Recherches Semiotiques/Semiotic Inquiry (1982) 2,3: 276-308.


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