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The English anthropologist Edward Taylor defined culture as:
“… that complex whole which includes knowledge, belief, art, morals, law, custom, and any other capabilities and habits acquired by man as a member of society”.
Indeed, by this definition, animals would lack culture. However, by other definitions, for example John Bonner's definition,"the transfer of information by behavioural means", many animals, plants and even bacteria possess culture. Most comparative psychologists have preferred to define culture in an equally broad, and arguably less anthropocentric way. McGrew argues that there are four essential features of culture:
- It is socially learned
- It is non-instinctive
- It is normative (many individuals in a group exhibit it)
- It is collective
Therefore, it is not features such as tool use or language which distinguishes culture, but the mode it is learned and transmitted. This broad mode of defining culture does not mean to detract from the social sciences, but stimulate comparative research in animals, and is thus used solely in this context. 
Other researchers have used the terms proto-culture, sub-culture or "culture" in quotation marks to term non-human culture, however now most researchers simply call culture in animals culture. van Schaik and colleagues proposed a taxonomy of four different types of cultures based on the presumed complexity of innovation that is involved.  These cultures include: (i) labels (arbitrary learnt associations such as food preferences or predator recognition with no innovation) (ii) signals (arbitrary innovations of displays such as song dialects in birds) (iii) skills (rare innovations which may be adaptive and presumed to be complex to socially learn), (iv) symbols. These labels for different cultures have been criticised as arbitary and vague.
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.
Culture from the Evolutionary Perspective
Memes and Cultural Evolution
Richard Dawkins argues for the existence of a "unit of cultural transmission" called a meme. This concept of memes has become much more accepted as more extensive research has been done into cultural behaviors. Much as one can inherit genes from each parent, subsequently individuals acquire memes through imitating what they observe around them. Memes are an extremely important factor when it comes to creation of culture. The more relevant actions (actions that increase ones probability of survival), such as architecture and craftwork are more likely to become prevalent, enabling a culture to form. The idea of memes as following a form of Natural Selection was first presented by Daniel Dennett. It has also been argued by Dennett that memes are responsible for the entirety of human consciousness. He claims that everything that constitutes humanity, such as language and music is a result of memes and the unflinching hold they have on our thought processes.
There was a hypothetical simulation conducted in which a population selects different memes . It shows both the positive and negative side effects of these hypothetical imitations. One of the main disparities between humans and animals is that humans have a much higher capacity for imitation. Also concluded was that the use of memes is responsible for the large brain size of humans. By mapping how long each of these memes will take to become an actual cultural change the conclusion of the study was that it is possible for human culture to have evolved using this model.
Culture has had a renewed interest in the evolutionary sciences due to the insight that not only that culture (or "memes") can evolve, but that culture can effect natural selection on genes, and genes can effect culture. From this perspective evolution (particularly hominin evolution) can be seen as an interplay between two processes: genetic and cultural evolution.
One highly illustrative example is natural selection on the ability for dairy processing. For most adults around the globe, consuming milk makes them ill, due to the lack of lactase activity after childhood.. A single nucleotide polymorphism in genes is known to keep the activity levels of this enzyme high throughout life. There is a significant correlation between populations which have a history of dairy farming and the ability to consume milk throughout life.  Analysis on the gene shows that the onset of selection was around 5,000-10,000 years ago, around when we know dairy farming began, as shown by high levels of cattle remains in neolithic sites. </span>
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. 
Social Learning and Cultural Transmission in Animals
- See main article: Social learning in animals
By definition, social learning is essential for culture. Forms of social learning is evident in many animal taxa. Various forms of social learning exist, from complex bodily imitation to relatively simple stimulus enhancement, which only involves drawing one's attention to an object.
Not all forms of social learning exist in all animals, however culture can exist with also relatively basic forms of social learning. For example, in Zebra Danio Fish, a new predator which attacks a Danio will cause an alarm reaction from the shoal . This leads to individuals who were not attacked associating the new predator with the alarm reaction, leading to cultural differences in predator knowledge, through little more than basic stimulus-assocation responses.
Genetic Vs. Cultural Transmission
Genetic transmission, like cultural transmission is a means of passing behavioral traits from one individual to another.The main difference is that Genetic transmission is the transfer of behavioral traits from one individual to another through genes which are transferred to an organism from its parents during the fertilization of the egg. As can be seen, genetic transmission can only occur once during the lifetime of an organism. Thus, genetic transmission is quite slow compared to the relative speed of cultural transmission. In cultural transmission, behavioral information is passed through means of verbal, visual, or written methods of teaching. Therefore, in cultural transmission, new behaviors can be learned by many organisms in a matter of days and hours rather than the many years of reproduction it would take for a behavior to spread among organisms in genetic transmission.
Until recently, teaching was a skill that was thought to be uniquely human. Now, as research has increased into the transmission of culture in animals, the role of teaching among animal groups has become apparent. Teaching is not merely limited to mammals either. Many insects, for example have been observed demonstrating various forms of teaching in order to obtain food. Ants, for example, will guide each other to food sources through a process called "tandem running," in which an ant will guide a companion ant to a source of food. It has been suggested that the "pupil" ant is able to learn this route in order to obtain food in the future or teach the route to other ants. There have been various recent studies that show that cetaceans are able to transmit culture through teaching as well. Killer whales are known to "intentionally beach" themselves in order to catch and eat pinnipeds who are breeding on the shore. Mother killer whales teach their young to catch pinnipeds by pushing them onto the shore and encouraging them to attack and eat the prey. Because the mother killer whale is altering her behavior in order to help her offspring learn to catch prey, this is evidence of teaching and cultural learning. The intentional beaching of the killer whales, along with other cetacean behaviors such as the variations of songs among humpback whales and the sponging technique used by the bottlenose dolphin to obtain food, provide substantial support for the idea of cetacean cultural transmission.
Imitation is often misinterpreted as merely the observation and copying of another's actions. This would be known as mimicry, because the repetition of the observed action is done for no other purpose than to copy the original doer or speaker. In the scientific community, imitation is rather the process in which an organism purposefully observes and copies the methods of another in order to achieve a tangible goal. Therefore, the identification and classification of animal behavior as being imitation has been very difficult. Recent research into imitation in animals has resulted in the tentative labeling of certain species of birds, monkeys, apes, and cetaceans as having the capacity for imitation. For example, a Grey parrot by the name of Alex underwent a series of tests and experiments at the University of Arizona in which scientist Irene Pepperberg judged his ability to imitate the human language in order to create vocalizations and object labels. Through the efforts of Pepperberg, Alex has been able to learn a large vocabulary of English words and phrases. Alex can then combine these words and phrases to make completely new words which are meaningless, but utilize the phonetic rules of the English language. Alex's capabilities of using and understanding more than 80 words, along with his ability to put together short phrases, demonstrates how birds, who many people do not credit with having deep intellect, can actually imitate and use rudimentary language skills in an effective manner. The results of this experiment culminated with the conclusion that the use of the English language to refer to objects is not unique to humans and is arguably true imitation, a basic form of cultural learning found in young children.
Language is another key indicator of animals who have greater potential to possess culture. Though animals do not naturally use words like humans when they are communicating, The well-known parrot Alex demonstrated that even animals with small brains, but are adept at imitation can have a deeper understanding of language after lengthy training. A bonobo named Kanzi has taken the use of the English language even further. Kanzi was taught to recognize words and their associations by using a lexigram board. Through observation of its mother's language training, Kanzi was able to learn how to use the lexigrams to obtain food and other items that he desired. Also, Kanzi is able to use his understanding of lexigrams to decipher and comprehend simple sentences. For example, when he was told to "give the doggie a shot," Kanzi grabbed a toy dog and a syringe and gave it a realistic shot. This type of advanced behavior and comprehension is what scientists have used as evidence for language based culture in animals.
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 "cultural" behavior in chimpanzees, a particular form of social grooming (the grooming-hand-clasp), was reported in 1978. McGrew and Tutin compared two longitudinal studies of chimpanzees in Tanzania, and showed that only the group in the Mahali Mountains and not those in the Gombe (where famously Jane Goodall conducted research since 1960), demonstrated this form of social grooming. Other aspects of this behaviour made it interesting; it seemed to be performed by many members of the group, no infants performed the behavior, subordinate individuals would hold the others wrist while grooming more often, and importantly its social function seemed to rule out environmental explanations of behavior. 
A handful of studies have since systematically compared several groups of primates for behavioral diversity which could indicate culture. Andrew Whiten and colleagues published a seminal paper on chimpanzee behavioural diversity, identifying 39 behavioral patterns which potentially have a cultural origin across 7 groups.  The criteria used to seperate cultural from non-cultural behaviour included whether the behavior was: normative, habitual, present in some but not all groups and no ecological explanation of behavior identified. Many behaviors which were suggested to be cultural, were excluded by this criteria (26 out of 65), such as drumming the buttress of a tree (present in all groups) or one group very infrequently had individuals who would comb their hair with a stem. Similar studies have been published in other species, such as orangutans where 19 cultural variants have been identified across 6 groups .
However, some have questioned whether the evidence presented in primates controls for genetic and environmental differences. Differences between species could be accounted for by differences in genes caused by selection or genetic drift, or alternatively differences in the environment such as in the size or species of trees could create non-cultural differences between groups. Some maintain that the "gold-standard" of evidence in culture does not exist in primates, as it does in animals like fish. 
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."
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
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.
Imitation Explains the Propagation, not Stability of Animal Culture
Researchers at the De´partement d’Etudes Cognitives, Institut Jean Nicod, Ecole Normale Supe´rieure acknowledged a difficulty with research in social learning. To count acquired behavior as cultural, two conditions need must be met: the behavior must spread in a social group, and that behavior must be stable across generations. Research has provided evidence that imitation may play a role in the propagation of a behavior, but these researchers believe the fidelity of this evidence is not sufficient to prove stability of animal culture.
Other factors like ecological availability, reward-based factors, content-based factors, and source-based factors might explain the stability of animal culture in a wild rather than just imitation. As an example of ecological availability, chimps may learn how to fish for ants with a stick from their peers, but that behavior is also influenced by the particular type of ants as well as the condition. A behavior may be learned socially, but the fact that it was learned socially does not necessarily mean it will last. The fact that the behavior is rewarding has a role in cultural stability as well. The ability for socially-learned behaviors to stabilize across generations is also mitigated by the complexity of the behavior. Different individuals of a species, like crows, vary in their ability to use a complex tool. Finally, a behavior’s stability in animal culture depends on the context in which they learn a behavior. If a behavior has already been adopted by a majority, then the behavior is more likely to carry across generations out of a need for conforming.
Animals are able to acquire behaviors from social learning, but whether or not that behavior carries across generations requires more investigation.
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- Detailed article on defining culture
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- ↑ Bonner, J.T. (1980) The Evolution of Culture in Animals. Princeton University PRess
- ↑ McGrew, W. C. (2004) The Cultured Chimpanzee: Reflections on Cultural Primatology. Cambridge: Cambridge University Press.
- ↑ Laland, K.N. & Hoppitt, W. (2008) Do animals have Culture? Evolutionary Anthropology, 12, 150-159.
- ↑ McGrew, W.C. & Tutin, C.E.G. (1978) Evidence For a Social Custom in Wild Chimpanzees? Man,13, 234-251.
- ↑ van Schaik, C.P. et al. (2003) Orangutan Cultures and the Evolution of Material Culture. Science, 299, 102-105.
- ↑ 6.0 6.1 6.2 6.3 Laland, Kevin N. and Bennett G. Galef, eds. The Question of Animal Culture. Cambridge, Mass: Harvard UP, 2009.
- ↑ 7.0 7.1 7.2 7.3 Holdcroft, David, and Harry Lewis. Memes, Minds, and Evolution. Philosophy 75.292 (2000): 161-182.
- ↑ 8.0 8.1 Higgs, Paul G. The Mimetic Transition: A Simulation Study of the Evolution of Learning by Imitation. Proceedings: Biological Sciences 267.1450 (2000):1355-1361
- ↑ Laland, K.N. & Brown, G.R. (2011) Sense and Nonsense: Evolutionary Perspectives on Human Behaviour. Oxford: Oxford University Press, p.177-178.
- ↑ Bersaglieri, T. et al. (2004) Genetic signatures of strong recent positive selection at the lactase gene. American Journal of Human Genetics, 74, 1111-1120.
- ↑ Heyes, Cecelia M. and Bennett G. Galef, Jr., eds. Social Learning in Animals: The Roots of Culture. San Diego: Academic Press, 1996. </li>
- ↑ Suboksi, M.D. et al. (1990) Alarm Reaction in Acquisition and Social Transmission of Simulated-Predator Recogntion by Zebra Danio Fish (rachydanio rerio). Journal of Comparative Psychology, 104, 1, 101-112. </li>
- ↑ Bonner, John Tyler (1980) The Evolution of Culture in Animals. Princeton University Press, Princeton </li>
- ↑ Caro, T. M. & Hauser, M. D. 1992. Is there teaching in non-human animals? Quarterly Review of Biology, 67, 151–174 </li>
- ↑ 15.0 15.1 Hoppitt WJ, GR Brown, R Kendal, L Rendell, A Thornton, MM Webster, and KN Laland. Lessons from Animal Teaching. Trends in Ecology & Evolution (Personal Edition). 23. 9 (2008): 486-93. </li>
- ↑ 16.0 16.1 16.2 16.3 16.4 16.5 16.6 Rendell L, and H Whitehead. Culture in Whales and Dolphins. The Behavioral and Brain Sciences. 24. 2 (2001): 309-24. </li>
- ↑ Hurley, S. L., and Nick Chater. Perspectives on Imitation From Neuroscience to Social Science. CogNet. Cambridge, Mass: MIT Press, 2005. </li>
- ↑ Pepperberg, Irene M. Grey Parrots Do Not Always 'parrot': the Roles of Imitation and Phonological Awareness in the Creation of New Labels from Existing Vocalizations. </li>
- ↑ 19.0 19.1 19.2 19.3 Hillix, William A., and Duane M. Rumbaugh. Animal Bodies, Human Minds: Ape, Dolphin, and Parrot Language Skills. Developments in primatology. New York: Kluwer Academic/Plenum Publishers, 2004. </li>
- ↑ 20.0 20.1 Huffman, Michael A., Charmalie A.D. Nahallage, and Jean-Baptiste Leca. Cultured Monkeys: Social Learning Cast in Stones. Current Directions in Psychological Science 17 (2008) 410-414. </li>
- ↑ McGrew, W.C. & Tutin, C.E.G. (1978) Evidence For a Social Custom in Wild Chimpanzees? Man,13, 234-251. </li>
- ↑ Whiten, A. et al. (1999) Cultures in Chimpanzees. Nature, 399, 682-685. </li>
- ↑ van Schaik, C.P. et al. (2003) Orangutan Cultures and the Evolution of Material Culture. Science, 299, 102-105. </li>
- ↑ Laland, K.N. & Hoppitt, W. (2003) Do Animals Have Culture? Evolutionary Anthropology, 12, 150-159. </li>
- ↑ Sargeant, Brooke L., and Janet Mann. From Social Learning to Culture: Intrapopulation Variation in Bottlenose Dolphins. The Question of Animal Culture. Ed. Kevin N. Laland and Bennett G. Galef. Cambridge, Mass: Harvard UP, 2009. 152-73. </li>
- ↑ 26.0 26.1 West, Meredith J., and Andrew P. King. Social Learning: Synergy and Songbirds. Social Learning in Animals: The Roots of Culture. Ed. Cecelia M. Heyes and Bennett G. Galef. San Diego: Academic P, 1996. 155-78. </li>
- ↑ Terkel, Joseph. Cultural Transmission of Feeding Behavior in the Black Rat (Rattus rattus). Social Learning in Animals: The Roots of Culture. Ed. Cecelia M. Heyes and Bennett G. Galef. San Diego: Academic P, 1996. 17-48. </li>
- ↑ Galef, Bennett G. Culture in Animals? The Question of Animal Culture. Ed. Kevin N. Laland and Bennett G. Galef. Cambridge, Mass: Harvard UP, 2009. 222-246. </li>
- ↑ Claidiere, N., & Sperber, D. (2010). Imitation explains the propagation, not the stability of animal culture.Proceedings of The Royal Society B: Biological Sciences, 277(1681), 651-659. doi: 10.1098/rspb.2009.1615 </li></ol>
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