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Animal culture describes the current theory of cultural learning in non-human animals through socially transmitted behaviors. The question as to the existence of culture in non-human societies has been a contentious subject for decades, much due to the inexistence of a concise definition for culture. However, many leading scientists agree on culture being defined as a process, rather than an end product. This process, most agree, involves the social transmittance of a novel behavior, both among peers and between generations. This behavior is shared by a group of animals, but not necessarily between separate groups of the same species.

The notion of culture in animals dates back to Aristotle and Darwin, but the association of animals' actions with the actual word "culture" first was brought forward with Japanese primatologists' discoveries of socially-transmitted food behaviors in the 1940s.

History of Animal Culture
Though the idea of 'culture' in animals has only been around for just over half of a century, scientists have been noting social behaviors of animals for centuries. Aristotle was the first to provide evidence of social learning in the songs of birds. Charles Darwin first attempted to find the existence of imitation in animals when attempting to prove his theory that the human mind had evolved from that of lower beings. Darwin was also the first to suggest what became known as social learning in attempting to explain the transmission of an adaptive pattern of behavior through a population of honey bees.

The vast majority of cultural anthropological research has been done on non-human primates, due to their being closest evolutionarily to humans. In non-primate animals, research tends to be limited, and therefore evidence for culture strongly lacking. However, the subject has become more popular recently, and has prompted the initiation of more research into the area.

Primate Culture
The beginning of the modern era of animal culture research in the middle of the 20th century came with the gradual acceptance of the term “culture” in referring to animals. Japan’s leading primatologist of the time, Kinji Imanishi, first used the word with a prefix as the term “pre-culture” in referring to the now infamous potato-washing behavior of Japanese macaques. In 1948, Imanishi and his colleagues began studying macaques across Japan, and began to notice differences among the different groups of primates, both in social patterns and feeding behavior. In one area, paternal care was the social norm, while this behavior was absent elsewhere. One of the groups commonly dug up and ate the tubers and bulbs of several plants, while monkeys from other groups would not even put these in their mouths. Imanishi had reasoned that, “if one defines culture as learned by offspring from parents, then differences in the way of life of members of the same species belonging to different social groups could be attributed to culture.” Following this logic, the differences Imanishi and his colleagues observed among the different groups of macaques may suggest that they had arisen as a part of the groups’ unique cultures. The most famous of these eating behaviors was observed on the island of Koshima, where one young female was observed carrying soiled sweet potatoes to a small stream, where she proceeded to wash off all of the sand and dirt before eating. This behavior was then observed in one of the monkey’s playmates, then her mother and a few other playmates. The potato-washing eventually spread throughout the whole macaque colony, encouraging Imanishi to refer to the behavior as “pre-culture,” explaining that, “we must not overestimate the situation and say that ‘monkeys have culture’ and then confuse it with human culture”. At this point, most of the observed behaviors in animals, like those observed by Imanishi, were related to survival in some way.

The first evidence of apparently arbitrary traditions came in the late-1970s, also in the behavior of primates. At this time, researchers McGrew and Tutin found a social grooming handclasp behavior to be prevalent in a certain troop of chimpanzees in Tanzania, but not found in other groups nearby. This grooming behavior involved one chimpanzee taking hold of the hand of another and lifting it into the air, allowing the two to groom each others’ armpits. Though this would seem to make grooming of the armpits easier, the behavior actually has no apparent advantage. As Frans de Waal explains from his later observations of the hand-clasp grooming behavior in a different group of chimpanzees, “A unique property of the handclasp grooming posture is that it is not required for grooming the armpit of another individual…Thus it appears to yield no obvious benefits or rewards to the groomers.”

Prior to these findings, opponents to the idea of animal culture had argued that the behaviors being called cultural were simply behaviors that had evolutionarily evolved due to their importance to survival. After the identification of this initial non-evolutionarily advantageous evidence of culture, scientists began to find differences in group behaviors or traditions in various groups of primates, specifically in Africa. More than 40 different populations of wild chimpanzees have been studied across Africa, between which many species-specific, as well as population-specific, behaviors have been observed. The researching scientists found 65 different categories of behaviors among these various groups of chimpanzees, including the use of leaves, sticks, branches, and stones for communication, play, food gathering or eating, and comfort. Each of the groups used the tools slightly differently, and this usage was passed from chimpanzee to chimpanzee within the group through a complex mix of imitation and social learning.

Cetacean Culture
Second only to non-human primates, culture in species within the order Cetacea, which includes whales, dolphins, and porpoises, has been studied for numerous years. In these animals, much of the evidence for culture comes from vocalizations and feeding behaviors.

Cetacean vocalizations have been studied for many years, specifically those of the bottlenose dolphin, humpback whale, killer whale, and sperm whale. Since the early 1970s, scientists have studied these four species in depth, finding potential cultural attributes within group dialects, foraging, and migratory traditions. Hal Whitehead, a leading cetologist, and his colleagues conducted a study in 1992 of sperm whale groups in the South Pacific, finding that groups tended to be clustered based on their vocal dialects. The differences in the whales’ songs among and between the various groups could not be explained genetically or ecologically, and thus was attributed to social learning. In mammals such as these sperm whales or bottlenose dolphins, the decision on whether an animal has the capacity for culture comes from more than simple behavioral observations. As described by ecologist Brooke Sergeant, “on the basis of life-history characteristics, social patterns, and ecological environments, bottlenose dolphins have been considered likely candidates for socially learned and cultural behaviors,” due to being large-brained and capable of vocal and motor imitation. In dolphins, scientists have focused mostly on foraging and vocal behaviors, though many worry about the fact that social functions for the behaviors have not yet been found. As with primates, many humans are reluctantly willing, yet ever so slightly willing, to accept the notion of cetacean culture, when well evidenced, due to their similarity to humans in having “long lifetimes, advanced cognitive abilities, and prolonged parental care.”

Avian Culture
Birds have been a strong study subject on the topic of culture due to their observed vocal “dialects” similar to those studied in the cetaceans. These dialects were first discovered by zoologist Peter Marler, who noted the geographic variation in the songs of various songbirds. Many scientists have found that, in attempting to study these animals, they approach a stumbling block in that it is difficult to understand these animals’ societies due to their being so different from our own. This makes it difficult to understand the animals’ behaviors, let alone determine whether they are cultural or simply practical.

However, despite this hindrance, evidence for differing dialects among songbird populations has been discovered, especially in sparrows, starlings, and cowbirds. In these birds, scientists have found strong evidence for imitation-based learning, one of the main types of social learning. Though the songbirds obviously learn their songs through imitating other birds, many scientists remain skeptical about the correlation between this and culture: “…the ability to imitate sound may be as reflexive and cognitively uncomplicated as the ability to breathe. It is how imitation affects and is affected by context, by ongoing social behavior, that must be studied before assuming its explanatory power." The scientists have found that simple imitation does not itself lay the ground for culture, whether in humans or birds, but rather it is how this imitation affects the social life of an individual that matters.

Culture in Other Animals
When moving past birds and into other creatures, the cultural research becomes few and far between. Notable research has been done with black rats, Norwegian rats, guppies, and ants.

Among the cultural studies of rats, the most widely discussed research is that performed by Joseph Terkel in 1991 on a species of black rats that he had originally observed in the wild in Israel. Terkel conducted an in-depth study aimed to determine whether the observed behavior, the systematic stripping of pine cone scales from pine cones prior to eating, was a socially acquired behavior, as this action had not been observed elsewhere. The experimentation with and observation of these black rats was one of the first to integrate field observations with laboratory experiments to analyze the social learning involved. From the combination of these two types of research, Terkel was able to analyze the mechanisms involved in this social learning to determine that this eating behavior resulted from a combination of ecology and cultural transmission, as the rats could not figure out how to eat the pinecones without being “shown” by mature rats. Though this research is fairly recent, it is often used as a prime example of evidence for culture in non-primate, non-cetacean beings.