MAMMALIAN SOCIAL LEARNING: COMPARATIVE AND ECOLOGICAL PERSPECTIVES.

Hilary O. Box and Kathleen R. Gibson (eds). Cambridge University Press. £60.00 ISBN 0 521 63263 3 (1999)

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It is very refreshing to read a book on social learning that does not focus almost entirely on mechanisms. In this book many of the authors examine social learning within the context of the social and physical environments of their animals and the particular challenges that they face. It includes a wide range of species, some of which have not been discussed in this context before, and for the first time, discussion is included of the social learning of prehistoric hominids.

The book is organised into six parts, with introductory comments by editors. For primatologists, the first and last parts are the most relevant (and therefore elaborated upon here), but one of the beauties of this book is that it places primates into a wider context. In Part 1, New perspectives in studies of social learning, this point is addressed in the opening chapter by Thelma Rowell: "The myth of peculiar primates". This chapter examines the belief that primates are somehow "special" in comparison to all other animals. Rowell suggests that perceived differences are more due to the way in which primates have been studied (small sample sizes meaning that individuals are studied intensively and for long periods), and the type of people who study them (backgrounds in the social sciences). The author suggests that now that the methods and questions asked in primatology are being applied to other species, the idea that primates face unique social and environmental challenges that make them more sophisticated than other animals is being challenged.

The second chapter "New directions in the study of primate learning " is by Barbara King. By using examples from her own studies of baboons, King argues that the dominant approach to the study of social learning, looking at mechanisms, is difficult under field conditions. She suggests that good alternatives to the mechanism-based framework are being developed. She describes a functional approach ("focus on the results of the actions rather than the means by which the results are achieved.") The idea is to assess social information acquisition and donation. After providing several examples, King concludes by emphasising not that one approach is better than another, but that it is time to move beyond a "mechanism-dominated framework".

The next chapter "Temperament and socially mediated learning among primates" by Hilary Box follows on well from the previous one. Box also advocates a functional approach and points out that information is not passed in neat intact packages but is interpreted and transformed by individuals. Infant monkeys acquire information but do so in different ways. Box suggests that one way in which individual difference in social learning could be explored is through temperament. Her review concludes that "the organisation of behaviour among individuals influences opportunities for, and potential use of, information about the environment that is acquired in the context of interactions with other individuals".

Richard Silby contributes the final chapter in the first part of the volume, "Evolutionary biology of skill and information transfer". He explains how genetic and life history considerations are related to the evolution of the ability to transmit and receive information, and considers the conditions under which mechanisms promoting the transfer of skill and information are favoured by natural selection. Overall, Part 1 contains the majority of the theoretical considerations and is great reading for anyone looking for fresh ideas as a fair number of under-explored areas are outlined in this chapter.

Part 2 "Social learning among species of terrestrial herbivores, contains five chapters. Karen Higginbottom and David Croft's chapter "Social learning in marsupials" is an excellent introduction to these fascinating animals in which no-one has explicitly examined social learning. "The social context for learning and behavioural development among wild African Elephants" by Phyllis Lee and Cynthia Moss is based on the long-term study of 381 elephant calves born between 1976 and 1985 at Amboseli, Kenya. David Klein in his chapter on "Comparative social learning among arctic herbivores: the caribou, muskox and arctic hare" describes how differences between these three species exist in their requirements and opportunities for social learning. It is interesting to see how information can be gained by animals who receive such minimal parental care in Robyn Hudson, Benoist Schaal and Ágnes Bilkó's chapter "Transmission of olfactory information from mother to young in the European rabbit". Finally it is pleasing to see Donald Broom's chapter "Social transfer of information in domestic animals" included with the "real" animal ones, and to consider the practical implications of social learning.

Part 3, Rats, bats and naked mole-rats: Animals with information centres, starts with "Exploring the dynamics of social transmission with rats" by Kevin Laland. He outlines the drawbacks of the current most popular methods used to investigate social learning and describes the results of a number of experiments using a different paradigm &endash; the transmission chain approach. In "Social influences on foraging in bats" Gerald Wilkinson and Janette Wenrick Boughman outline the reasons why social learning may be found in bats and test a model which predicts the frequency of social learning at an information centre where animals can acquire information about food choice and location. Chris Faulkes' chapter "Social transmission of information in a eusocial rodent, the naked mole-rat (Heterocephalus glaber)" suggests that social learning plays a part in reproductive suppression, foraging and the recognition of colony members.

Part 4, Social learning among species of terrestrial carnivores, begins with "Opportunities for social learning in bears" by Barrie Gilbert. Although bears are not obvious candidates for the study of social learning, their long juvenile period, high level of attachment to their mothers, specialisation of foraging tactics within local populations and mimicry of parental behavioural traits, suggest that social learning is important to these animals. Andrew Kitchener reviews evidence from field observations, experimental studies of domestic cats and the behaviour of released captive animals in his chapter "Watch with mother: a review of social learning in the Felidae". Taken together, these sources strongly suggest that the ability to recognise appropriate prey and hunt efficiently is socially learned. This is followed by Jan Nel's chapter "Social learning in canids: An ecological perspective". Nel describes how varied canid species are and suggests that the opportunity for, and usefulness of social learning should vary between species.

The fifth part is on Dolphins and whales: Communication and foraging in aquatic environments. James Boran and Sara Heimlich begin with "Social learning in cetaceans: Hunting, hearing and hierarchies". There are numerous interesting examples in this chapter, especially feeding techniques and the practice of beaching in the pursuit of sea lions and elephant seals. Young whales have been seen practising this behaviour while being observed, and sometimes rescued by adults. The actions of the adults suggest that teaching behaviour may be taking place. This chapter also provides examples of cultural traditions including the case of a school of dolphins who co-operate with fisherman off the coast of Brazil. It is suggested that one reason why females of some species live well beyond reproductive age could be the cross-generational transfer of information. Vincent Janik's chapter on the "Origins and implications of vocal learning in bottlenose dolphins" details these animals' vocal capacities in relation to the aquatic environment and examines a number of hypotheses that attempt to explain the evolution of social learning in this context.

The final part, The great ape &endash; human adaptation: culture and the cognitive niche, begins with Dick Byrne's "Cognition in great ape ecology: Skill-learning ability opens up foraging opportunities". Byrne describes differences between great ape and monkey food processing skills, asks whether these differences might reflect more powerful, cognitively complex mechanisms of social learning, and finally examines what we understand about the mechanisms in the acquisition of natural feeding behaviour. Kathleen Gibson's chapter "Social transmission of facts and skills in the human species: Neural mechanisms" argues that the human brain indirectly contributes to social learning capacities by requiring birthing and developmental conditions that provide opportunities for the intergenerational transmission of information. Gibson examines the idea that there must have been strong selective pressures that favoured the abilities that come with brain enlargement.

"Cultural learning in hominids: A behavioural ecological approach" by Stephen Shennan and James Steele concentrates on the manufacture of stone tools and (for non-human primates) uses Boesch's (1993) assertion that the acquisition of this behaviour "entails processes of true imitation guided by frequent maternal intervention". Shennen and Steele argue that the same must have been true for early hominids. The model outlined in this chapter suggests that parent-offspring transmission was the predominant route for the transfer of skills. This suggests a link between genetic and cultural transmission. The final chapter by Steven Mithen, "Imitation and cultural change: A view from the Stone Age, with specific reference to the manufacture of handaxes", uses evidence from the Lower Palaeolithic to challenge the idea that the absence of cultural evolution in animals is due to the absence of imitation. The idea is that imitation allows cultural traditions to build on each other, increasing technical complexity. The author argues that handaxes are difficult to make yet the high degree of similarity between those found means that the manufacturing process was learned by imitation. However, the same type of handaxe is found in the archaeological record for more than 1 million years, so cumulative cultural change is absent; only appearing 50,000 years ago when some other factor came into play. The author concludes that those who are trying to understand the relationship between social learning and cultural change should not ignore the archaeological record.

Many of the chapters are based upon papers presented at the tripartite meeting of the Zoological Society of London, The Mammal Society and the Primate Society of Great Britain meeting in 1996. The editors have done an excellent job in compiling material on such a wide diversity of taxa, and by providing this database of comparative information. Overall, this book will broaden your horizons on social learning. It would be great if another edition comes out in the future to see how many of the questions posed have been pursued &endash; and what the answers are.