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Culture in the Animal Kingdom
Sunday, 01 May 2005

Mark Harrison explores the evidence for cultural behaviour in animals.

Culture has historically been thought of as a uniquely human characteristic: the Oxford English Dictionary defines culture as “the civilization, customs, and artistic achievements of a people, especially at a certain stage of its development or history”. However, recent discoveries have confirmed that, like the use of tools and a highly developed brain, culture is fairly common in the animal kingdom. Yet studies reveal that it is not just restricted to our closest relatives.

Culture is important to biologists — anthropologists in particular — because in order to exhibit culture an animal must be capable of socially transmitted learning, traditionally considered a hallmark of high intelligence. Revealing the extent of culture in the animal kingdom, especially in the primate lineage, may help us to understand how and why human culture and intelligence evolved. The Oxford English Dictionary’s definition precludes the possibility of culture in animals, and so a better definition for biologists might be that culture is “a group- or population-typical behaviour pattern shared by members of a community and relying on socially learned and transmitted information”.

If an animal is capable of social learning, once discovered, a useful behaviour will spread through the population until it reaches an unsurpassable geographical barrier, such as an ocean or mountain range. Behaviour seen on one side of the barrier will therefore differ from that on the other side. But just observing differences in behaviour between populations, not necessarily caused by differences in social learning ability, could be due to genetic or ecological factors. For example, if a tool is only present at the site where it is in use, it is difficult to say for certain whether that use is cultural, or whether it would be used at both sites if it were present. Only when ecological and genetic explanations can be discounted can a behavioural variant confidently be considered as cultural. Hence culture is easiest to identify when it involves non-essential behaviours, for example grooming techniques, as these are less likely to be affected by ecological factors. The ability to learn from others is useful because it allows the bearer to adapt to changes more quickly than solely through genetic adaptation or individual learning.

A sensible place to start looking for culture in the animal kingdom is in our closest living relatives, the great apes. It has long been established that great apes in captivity are capable of learning a wide variety of behaviours including problem-solving, tool use and even sign language. Despite many years of debate the existence of cultural behaviour in the wild has only really been confirmed in the past six years. Information collected by Andrew Whiten and colleagues from seven sites, totalling 151 years of observation, revealed that an ecological explanation could be discounted — and hence a cultural explanation accepted — for 39 different behaviours found to vary between the sites, including forms of tool use, grooming techniques and courtship behaviour. This is consistent with our genetic similarities to chimpanzees, and the numerous observations of tool use by chimpanzees in the wild, including fishing for termites and cracking nuts with hammers.

Carel van Schaik and colleagues looked at cultural behaviours in orangutans, which rarely use tools in the wild and are largely solitary. Long-term observations from six research sites in Borneo and Sumatra demonstrated a similar pattern of cultural behaviour in wild orangutans to that seen in chimpanzees: blowing raspberries when making their nest for the night, opening up tough foods with sticks, and building roofs on their nests to protect themselves from the rain. With the exception of tool use during feeding, cultural variations in great ape feeding behaviour have yet to be documented. This is because small differences in the availability and nutritional content of food items between sites make it difficult to preclude ecological explanations. Current studies by the author and a number of other researchers aim to take these small ecological differences into account and reveal the true extent of cultural variation in orangutan feeding behaviour.

Sceptics have pointed out that in virtually all these cases it is impossible to completely discard an ecological or genetic explanation. Much of the available evidence for culture in primates remains based on plausibility arguments. For example, chimpanzees in one area may have a greater supply of fruit, their preferred food, and so may not use hammers to crack nuts even if both the nuts and hammers are available to them. This possibility is hard to exclude, but the studies on orangutan feeding behaviour mentioned above may go some way towards achieving this. Furthermore, the long life span of many primates — an estimated 58 years for orangutans in the wild — hinders the study of transmission of traditions between generations. For these reasons, the best candidates for the demonstration of culture in animals actually come from non-primates.

Whales and dolphins also exhibit cultural behaviour. Male humpback and bowhead whales sing to attract females, and the songs of different males within a population at any given time are virtually identical, despite the fact that the population of thousands may span thousands of kilometres. These songs vary widely and continuously, both within and between breeding seasons. The speed and geographic breadth of these changes makes an ecological or genetic explanation highly unlikely. Off the coast of Argentina, killer whales swim ashore to catch seals. This behaviour is very risky as beached individuals will die if they can’t return to the ocean. Killer whale mothers have been observed teaching their young this technique, actively pushing juveniles up onto the beach, directing them towards prey and helping them back into the sea afterwards. This satisfies all the conditions of teaching and social learning and so can be considered cultural.

The best evidence yet for culture comes from studies on fish, in which one can conduct direct and rigorous experimental tests. For example, Helfman and Schultz’s translocation experiments in the early 1980s — on the elegantly named ‘French grunt’— showed that translocated fish adopt the schooling and migratory behaviour of the group to which they were introduced, rather than that from where they came. The same study also showed that control fish translocated into the same area, but with resident ‘teacher fish’ removed, did not adopt this behaviour. These experiments completely preclude both genetic and ecological explanations. The confirmation of cultural behaviour in fish provides excellent support for its presence in primates, given the latter species’ higher level of intelligence. There is also a hint that the mechanism of cultural transmission of behaviours may be simpler than was originally thought.

It is worth noting that the definition of culture used by biologists in such studies is much stricter than that typically used for humans. Many examples of human behaviour would not be considered cultural under the criteria used by biologists. The use of medicinal plants by forest Indians, for example, can be explained on ecological grounds, as these plants are not available to all human populations. We know that the knowledge of which plants to use for medical treatment is passed on by social learning in humans, but sceptics are unwilling to accept a cultural explanation for their behaviour, even in animals such as chimpanzees, who have a proven ability for social learning in captivity. Were animals to be judged by the same standards used for humans, we would find animal culture to be even more widespread than currently realised.

Some might argue that the degree of complexity in human culture is so great that animal and human cultures can never be compared. In light of this, many prefer to use the term ‘tradition’ rather than ‘culture’ when discussing behavioural variants in animals. This is a little like saying that because digestion or locomotion differ in humans and chimpanzees, the two processes are not comparable. Whilst it is certainly true that the degree of cultural complexity in humans is much greater than that shown in even the most cultural animals, this raises the interesting question of whether our ancestors were also cultural. Would it be reasonable to assume that stone-age or Neanderthal man were cultural? What about Homo erectus and Australopithecus? Most would agree that these species would have been cultural to some extent, but chimpanzees — separated from us by just 5.5 million years — are considered to be our sister taxa and are almost identical to us genetically. So why not them too? An increasing number of anthropologists and biologists now believe that some animals can indeed be considered cultural and that a broader definition of culture — like that used in this article — is more useful for understanding how and why human culture evolved. It is now beyond doubt that the cultural differences between ourselves and the rest of the animal kingdom are differences of degree rather than kind, which leaves us to ponder once again, what does actually define us as human?

Mark Harrison is a PhD student in the Department of Anatomy

 
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