Animal culture - Knowledge

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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.

789:, 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. 1296:) where they took fish native to a specific schooling site and transported them to other sites. In this species of fish, the organism uses distinct, traditional migration routes to travel to schooling sites on coral reefs. These routes persisted past one generation and so by relocating the fish to different sites, Helfman and Schultz wanted to see if the new fish could relearn that sites' migration route from the resident fish. Indeed this is what they found: that the newcomers quickly learned the traditional routes and schooling sites. But when residents were removed under similar situations, the new fish did not use the traditional route and instead use new routes, suggesting that the behavior could not be transmitted once the opportunity for learning was no longer there. 1034:. 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. 906:. Among studies of rat culture, 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 606: 1128: 573:

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.

549: 242:. Certain individuals are especially concerned with the analysis of studies connecting "identity, collective memory, social classification, logics of action, and framing." Views of what exactly culture is have been changing due to the convergence of sociological and psychological thought on the subject by the 1990s. Culture is specific to region and not just one umbrella definition or concept can truly give us the essence of what culture is. Also referenced is the importance of symbols and rituals as cognitive building blocks for a psychological concept of shared culture. 679:

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.

1269:) exist in red- and black-headed subtypes, and these subtypes have been shown to have different levels of boldness (measured by the time taken to explore new areas, and other similar tests). Experiments placing black-headed birds (known to be less bold) in the company of red-headed birds (known to be more bold) resulted in the black-headed bird performing "bolder" behaviors, and red-headed birds became "shyer" in the presence of black-headed ones. The experimenters hypothesized that this individual-level conformity could lead to stronger social cohesion. 630:

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 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.

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rated the behavior as either customary – occurring in all individuals within that population; habitual – not present in all individuals, but repeated in several individuals; present – neither customary or habitual but clearly identified; absent – instance of behavior not recorded and has no ecological explanation; ecological – absence of behavior can be attributed to ecological features or lack thereof in the environment, or of unknown origin. Their results were extensive: of the 65 categories of behavior studied, 39 (including

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One of the first signs of culture in early humans was the utilization of tools. Chimpanzees have been observed using tools such as rocks and sticks to obtain better access to food. There are other learned activities that have been exhibited by other animals as well. Some examples of these activities that have been shown by varied animals are opening oysters, swimming, washing of food, and unsealing tin lids. This acquisition and sharing of behaviors correlates directly to the existence of memes. It especially reinforces the

257:. 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, it is suggested that individuals acquire memes through imitating what they observe around them. 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 597:. 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 injection. This type of advanced behavior and comprehension is what scientists have used as evidence for language-based culture in animals. 960: 503:. The experimental setup consisted of an apparatus containing food with two possible methods that could be used to obtain the food. Naïve meerkats learned and used the method exhibited by the "demonstrator" meerkat trained in one of the two techniques. Although in this case, imitation is not the clear mechanism of learning given that the naïve meerkat could simply have been drawn to certain features of the apparatus from observing the "demonstrator" meerkat and from there discovered the technique on their own. 1278: 564:

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. By the 2000s, research into imitation in animals had 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

1068:. Located in the eastern and southern parts of North America, these white-crowned song-birds exhibit learned vocal behavior. Marler and Tamura found that while song variation existed between individual birds, each population of birds had a distinct song pattern that varied in accordance to geographical location. For this reason, Marler and Tamura called the patterns of each region a "dialect": however, this term has since been disputed, as different types of in bird song are much less distinct than 650: 1440: 986: 1137: 969: 1106:) began to attack the bottles, opening the foil or cardboard lids and drinking the cream of the top. It was later shown that this innovative behavior arose independently in several different sites and spread horizontally (i.e. between living members) in the existing population. Later experimental evidence showed that conformity may lead to the horizontal spread of innovative behaviors in wild birds, and that this may in turn result in a lasting cultural tradition. 711:, and defined cultural behaviors as behaviors that are "transmitted repeatedly through social or observational learning to become a population-level characteristic". Eight years later, after "conducting large-scale controlled social-diffusion experiments with captive groups", Whiten et al. stated further that "alternative foraging techniques seeded in different groups of chimpanzees spread differentially...across two further groups with substantial fidelity". 7601: 7117: 1239:) were first analyzed to create a comprehensive network. Then, the order in which individuals learned task-solving behavior from a trained tutor was compared with the network. They not only found that the pattern of learning reflected the network that they had built, but that different types of social connections (such as "affiliative interactions" and "aggressive interactions") characterized different rates of information transmission and observation. 1115:). During the 20th century, individuals in this population began to non-fatally wound the backs of swimming whales with their beaks, feeding on the blubber and creating deeper lesions in areas that were already wounded. Aerial photographs showed that gull-induced lesions on local whales increased in frequency from 2% to 99% from 1974 to 2011, and that this behavior was not observed in any other kelp gull populations other than two isolated incidents. 1411:

constitutes unlearned responses to different selection pressures , but it is also necessary to consider the possibility of genetic variation precipitating different patterns of learning." Gene-culture coevolution, much like the interaction between cultural transmission and environment, both serve as modifiers to the original theories on cultural transmission and evolution that focused more on differences in the interactions between individuals.

5852: 1321: 22: 422:. Cultural transmission can also vary according to different social learning strategies employed at the species and or individual level. Cultural transmission is hypothesized to be a critical process for maintaining behavioral characteristics in both humans and nonhuman animals over time, and its existence relies on innovation, imitation, and communication to create and propagate various aspects of animal behavior seen today. 7141: 527:", 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. By the early 2000s, various 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 428:, when defined as the transmission of behaviors from one generation to the next, can be transmitted among animals through various methods. The most common of these methods include imitation, teaching, and language. Imitation has been found to be one of the most prevalent modes of cultural transmission in non-human animals, while teaching and language are much less widespread, with the possible exceptions of 7129: 727:

choosing which individuals they should imitate in order to increase their own fitness. This type of behavior is very common in human culture as well. People will seek to imitate the behaviors of an individual that has earned respect through their actions. From this information, it is evident that the cultural transmission system of chimpanzees is more complex than previous research would indicate.

1428:), naïve fish preferred taking a long, energetically costly route to a feeder that they had learned from resident fish rather than take a shorter route. These fish were also slower to learn the new, quicker route compared to naïve fish that had not been trained in the long route. In this case, not only is the social tradition maladaptive, but it also inhibits the acquisition of adaptive behavior. 338:, Whiten created a compilation of results from seven long-term studies totaling 151 years of observation analyzing behavioral patterns in different communities of chimpanzees in Africa (read more about it below). The study expanded the notion that cultural behavior lies beyond linguistic mediation, and can be interpreted to include distinctive socially learned behavior such as stone-handling and 637:, 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. Imanishi introduced the Japanese term 226:

than those who do not have a shared goal. A further definition of culture is, "ocially transmitted behavior patterns that serve to relate human communities to their ecological settings." This definition connects cultural behavior to the environment. Since culture is a form of adaptation to one's environment, it is mirrored in many aspects of our current and past societies.

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imitate the behaviors of the older, higher ranking chimpanzee as opposed to the younger, lower ranking individual when given a choice. It is believed that the older higher ranking individual had gained a level of 'prestige' within the group. This research demonstrates that culturally transmitted behaviors are often learned from individuals that are respected by the group.

1084:. They also showed that white-crowned sparrows only learn songs recorded from other members of their species. Marler and Tamura noted that this case of cultural transmission was interesting because it required no social bond between the learner and the emitter of sound (since all sounds originated from a loudspeaker in their experiments). 1307:), researcher Warner found that individuals chose mating sites based on social traditions and not based on the resource quality of the site. Warner found that although mating sites were maintained for four generations, when entire local populations were translocated elsewhere, new sites were used and maintained. 1389:, it is due more to ecological pressures than cultural ones. In other words, when an animal changes its behavior over its lifespan, this is most often a result of changes in its environment. Furthermore, animal behavior is also influenced by evolved predispositions, or genetics. It is very possible that " 932:

techniques are transmitted. In this habitat, the rats' only source of food is pine seeds that they obtain from pine cones. Terkel et al. studied the way in which the rats obtained the seeds and the method that this strategy was transmitted to subsequent generations. Terkel et al. found that there was

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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,

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This finding confirms not only that nonhuman species can maintain unique cultural traditions; it also shows that they can pass these traditions on from one population to another. The Whiten articles are a tribute to the unique inventiveness of wild chimpanzees, and help prove that humans' impressive

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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

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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

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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 other's armpits. Though this would seem

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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

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are more likely to be assimilated into our everyday lives. Organizations that utilize the four aforementioned aspects of culture are the ones that are the most successful. Therefore, cultures that are better able to involve their citizens towards a common goal have a much higher rate of effectiveness

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Victoria Reyes-García; James Broesch; Laura Calvet-Mir; Nuria Fuentes-Peláez; Thomas W. McDade; Sorush Parsa; Susan Tanner; Tomás Huanca; William R. Leonard; Maria R. Martínez-Rodríguez; TAPS Bolivian Study Team (July 2009). "Cultural transmission of ethnobotanical knowledge and skills: an empirical

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are a specific mechanism of cultural transmission in birds. Information learned in social contexts can allow them to make decisions that lead to increased fitness. A great deal of research has focused on the communication of new foraging locations or behaviors through social networks. These networks

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In a later study one more possible explanation for the transmission of sponging was ruled out in favor of cultural transmission. Scientists from the same lab looked at the possibility that 1.) the tendency for "sponging" was due to a genetic difference in diving ability and 2.) that these genes were

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sp.). It has been previously noted that tool use in foraging, called "sponging" exists in this species. "Sponging" describes a behavior where a dolphin will break off a marine sponge, wear it over its rostrum, and use it to probe for fish. Using various genetic techniques, Krutzen et al. showed that

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The older, higher ranking individual's success in similar situations in the past led the other individuals to believe that their fitness would be greater by imitating the actions of the successful individual. This shows that not only are chimpanzees imitating behaviors of other individuals, they are

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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

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have also been frontrunners in the field of cultural transmission, describing behavioral "traits" as characteristics pertaining to a culture that are recognizable within that culture. Using a quantifiable approach, Cavalli-Sforza & Feldman were able to produce mathematical models for three forms

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and transmission are the two main components of culture, specifically referencing tool making and the ability to acquire behaviors that will enhance one's quality of life. Using this definition it is possible to conclude that other animals are just as likely to adapt to cultural behaviors as humans.

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Brakes, Philippa; Dall, Sasha R. X.; Aplin, Lucy M.; Bearhop, Stuart; Carroll, Emma L.; Ciucci, Paolo; Fishlock, Vicki; Ford, John K. B.; Garland, Ellen C.; Keith, Sally A.; McGregor, Peter K.; Mesnick, Sarah L.; Noad, Michael J.; Sciara, Giuseppe Notarbartolo di; Robbins, Martha M.; Simmonds, Mark

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between distance between sites and 'cultural difference' might reflect the well-established correlation between genetic and geographical distances". The farther two populations of a species are separated from each other, the less genetic traits they will share in common, and this may be one source

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Other research showed that although conformity has a strong influence on behaviors adopted by birds, the local tradition can be abandoned in favor of an analogous behavior which gives higher reward. This showed that while conformity is a beneficial mechanism for quickly establishing traditions, but

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traditions with peers and offspring is much higher than that of one individual spreading some aspect of animal behavior to one or more members. Cultural transmission, as opposed to individual learning, is therefore a more efficient manner of spreading traditions and allowing members of a species to

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in other 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.

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A closely related concept to memes is the idea of evolutionary culture. The concept of evolutionary culture gained greater acceptance due to the re-evaluation of the term by anthropologists. The broadening scope of evolution from simple genes to more abstract concepts, such as designs and behaviors

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show cultural traditions in the construction of their communal nests. These nests vary in shape and size among different groups, even when living in close proximity. This variation is not influenced by genetic factors or environmental conditions, but rather reflects group-specific preferences that

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Chimpanzees have been known to use tools for as long as they have been studied. Andrew Whiten found that chimpanzees not only use tools, but also conform to using the same method as the majority of individuals in the group. This conformity bias is prevalent in human culture as well and is commonly

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Teaching is often considered one mechanism of social learning, and occurs when knowledgeable individuals of some species have been known to teach others. For this to occur, a teacher must change its behavior when interacting with a naïve individual and incur an initial cost from teaching, while an

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Brakes, Philippa; Carroll, Emma L.; Dall, Sasha R. X.; Keith, Sally A.; McGregor, Peter K.; Mesnick, Sarah L.; Noad, Michael J.; Rendell, Luke; Robbins, Martha M.; Rutz, Christian; Thornton, Alex; Whiten, Andrew; Whiting, Martin J.; Aplin, Lucy M.; Bearhop, Stuart; Ciucci, Paolo; Fishlock, Vicki;

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In some cases, conformity-based aggression may benefit individuals who conform to traditions. Researchers used the framework of sexual selection and conformism in of song types of songbirds to model territorial aggression against individuals with non-conforming song types. Their model showed that

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Vocalizations have also been proven to be culturally acquired in killer and sperm whale populations, as evidenced by the distinct vocalization patterns maintained by members of these different populations even in cases where more than one population may occupy one home range. Even within the same

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observation in an attempt to discover how much cultural variation existed between populations of the species. The synthesis of their studies consisted of two phases, in which they (1) created a comprehensive list of cultural variant behavior specific to certain populations of chimpanzees and (2)

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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

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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

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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

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occurs from parents to offspring and is a function which shows that the probability that parents of specific types give rise to an offspring of their own or of another type. Vertical transmission, in this sense, is similar to genetic transmission in biological evolution as mathematical models for

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identification transponders to experimentally manipulate avian social networks: this scanner technology allowed them to restrict access to feeders for some birds and not others. Their data showed that individuals are more likely to learn from those who were able to enter the same feeding area as

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In order to make a case for cultural transmission as the mode of behavioral inheritance in this case, Krutzen et al. needed to rule out possible genetic and ecological explanations. Krutzen et al. refer to data that indicate both spongers and nonspongers use the same habitat for foraging. Using

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plays an important role in cultural transmission in chimpanzees. Victoria Horner conducted an experiment where an older, higher ranking individual and a younger, lower ranking individual were both taught the same task with only slight aesthetic modification. She found that chimpanzees tended to

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with each other and with their young. Differences in cultural transmission across species have been thought to be largely affected by external factors, such as the physical environment, that may lead an individual to interpret a traditional concept in a novel way. The environmental stimuli that

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and whistles. The majority of their vocalizations are repetitions of the same calls, referred to as discrete or stereotyped calls, recorded since the 1960s and passed on by the orcas from generation to generation. A Southern Resident calf only learns the discrete calls used in the pod of their

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Conformity is one mechanism through which innovative behaviors can become embedded in culture. In an experimental setting, tits preferentially adopted the locally popular method of opening a two-action puzzle box even after discovering the other possible way of accessing the food. This formed

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and others to refer to the behavior as "pre-culture" and as being acquired through "pre-cultural propagation". The researchers caution 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

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Culture can also contribute to differences in behavior, but like genes and environments, it carries different weight in different behaviors. As Laland and Janik explain, "to identify cultural variation, not only is it not sufficient to rule out the possibility that the variation in behavior

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experiments where pups of naïve mothers were placed with experienced mothers and vice versa, those pups placed with experienced mothers learned the technique while those with naïve mothers did not. This result suggests that this optimal foraging technique is socially rather than genetically

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Culture, which was once thought of as a uniquely human trait, is now firmly established as a common trait among animals and is not merely a set of related behaviors passed on by genetic transmission as some have argued. Genetic transmission, like cultural transmission, is a means of passing

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Another argument against the "ethnographic method" is that it is impossible to prove that there are absolutely no ecological or genetic factors in any behavior. However, this criticism can also be applied to studies of human culture. Though culture has long been thought to arise and remain

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overlapped. In this study, knowledge about new feeding areas spread through social interactions: more birds visited the new area than the number of birds that discovered the area independently. The researchers noted that information likely travelled faster among members of the same species

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in twelve different acoustic settings and observing effects on their verbal behavior, Marler and Tamura found that sparrows learned songs during the first 100 days of their lives. In this experimental setting, male birds in acoustic chambers were exposed to recorded sounds played through a

536:. 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. 1373:

A popular method of approaching the study of animal culture (and its transmission) is the "ethnographic method," which argues that culture causes the geographical differences in the behavioral repertoires of large-brained mammals. Some researchers argue this downplays the roles that

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the behavior of "sponging" is vertically transmitted from the mother, with most spongers being female. Additionally, they found high levels of genetic relatedness from spongers, suggesting recent ancestry and the existence of a phenomenon researchers call a "sponging eve".

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In wild songbirds, social networks are a mechanism for information transmission both within and between species. Interspecific networks (i.e. networks including birds of different species) were shown to exist in multispecies flocks containing three different types of

173:, the ability to make new sounds through imitation. Most species cannot learn to imitate sounds. Some can learn how to use innate vocalizations in new ways. Only a few species can learn new calls. The transmission of vocal repertoires, including some types of 1247:

Bartlett and Slater observed call convergence (i.e. conformity) in budgerigars introduced into groups with different flock-specific calls than their own. They also found that the original calls of flock members did not change significantly during this process.

143:. Culture is increasingly seen as a process, involving the social transmittance of behavior among peers and between generations. It can involve the transmission of novel behaviors or regional variations that are independent of genetic or ecological factors. 5030:

Tello-Ramos, Maria C.; Harper, Lucy; Tortora-Brayda, Isabella; Guillette, Lauren M.; Capilla-Lasheras, Pablo; Harrison, Xavier A.; Young, Andrew J.; Healy, Susan D. (2024). "Architectural traditions in the structures built by cooperative weaver birds".

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under selection. From a test of 29 spongers and 54 nonspongers, the results showed that the coding mitochondrial genes were not a significant predictor of sponging behavior. Additionally, there was no evidence of selection in the investigated genes.

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Teaching is arguably the social learning mechanism that affords the highest fidelity of information transfer between individuals and generations, and allows a direct pathway through which local traditions can be passed down and transmitted.

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is that, though culture plays a huge role in the progression of animal behavior over time, the genes of a particular species have the ability to affect the details of the corresponding culture and its ability to evolve within that species.

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rarely imitate songs played from a loudspeaker, but they regularly imitate songs of an adult bird after only a few hours of interaction. Interestingly, imitation in zebra finches is inhibited when the number of siblings (pupils) increases.

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aggressors won more frequently when targeting non-conformers (than in un-targeted or random aggression). They also found that alleles for conformity-enforcement propagated more effectively than alleles for tolerance of non-conformity.

346:. The implications of their findings indicate that chimpanzee behavioral patterns mimic the distinct behavioral variants seen in different human populations in which cultural transmission has generally always been an accepted concept. 885:

data, Krutzen et al. found a significant non-random association between the types of mitochondrial DNA pattern and sponging. Because mitochondrial DNA is inherited maternally, this result suggests sponging is passed from the mother.

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diversity yet high diversity in matrilineal whale culture may be attributed to cultural transmission, since learned cultural traits have the ability to have the same effect as normal maternally inherited mtDNA. The divergence of the

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learned from each other to open garbage bins. Bin-opening spread more quickly to neighbouring suburbs than suburbs further away. In addition, birds in different areas developed their own variants for accomplishing the complex task.

523:, which is not limited to mammals. 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 " 270:

makes the idea of evolutionary culture more plausible. Evolutionary culture theory is defined as "a theory of cultural phylogeny." The idea that all human culture evolved from one main culture, citing the interconnectedness of

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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. In 1952, Japan's leading primatologist of the time,

1402:, which asserts that "cognitive, affective, and moral capacities" are the product of an evolutionary dynamic involving interactions between genes and culture over extended periods of time. The concept behind gene-culture 998: 166:' discoveries of socially-transmitted food behaviours in the 1940s. Evidence for animal culture is often based on studies of feeding behaviors, vocalizations, predator avoidance, mate selection, and migratory routes. 4986:

Palacín, Carlos; Alonso, Juan C.; Alonso, Javier A.; Magaña, Marina; Martín, Carlos A. (2011). "Cultural transmission and flexibility of partial migration patterns in a long-lived bird, the great bustard Otis tarda".

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One definition of culture, particularly in relation to the organizational aspect is the utilization of "involvement, consistency, adaptation, and mission." Cultural traits that are indicators of a successful form of

4425: 1019:. 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. 1170:

Juvenile birds that migrate in flocks may learn to navigate accurately through cultural transmission of route choice skills from older birds. Cultural inheritance of migration patterns has been shown in bustards

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In the study of social transmissions, one of the important unanswered questions is an explanation of how and why maladaptive social traditions are maintained. For example, in one study on social transmission in

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Rendell, Luke; Fogarty, Laurel; Hoppitt, William J.E.; Morgan, Thomas J.H.; Webster, Mike M.; Laland, Kevin N. (2011). "Cognitive culture: theoretical and empirical insights into social learning strategies".

177:, can be viewed as social processes involving cultural transmission. Some evidence suggests that the ability to engage in vocal learning depends on the development of specialized brain circuitry, detected in 261:. 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. 1222:(conspecifics), but that individuals did not depend solely on conspecifics for transmission. Another study on army-ant-following birds has also evidenced interspecific transmission of foraging information. 456:. The role of cultural transmission in cultural evolution, then, is to provide the outlet for which organisms create and spread traditions that shape patterns of animal behavior visibly over generations. 274:, has also been presented. There is, however, also the possibility for disparate ancestral cultures, in that the cultures observed today may potentially have stemmed from more than one original culture. 1000: 910:

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.

491:. Imitation is one of the most prevalent modes of cultural transmission in non-human animals, while teaching and language are much less widespread. In a study on food acquisition techniques in 1398:

independent of genetics, the constraints on the propagation and innovation of cultural techniques inevitably caused by the genome of each respective animal species has led to the theory of

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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

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them. Additionally, the existing "paths" of information transmission were altered following segregation during feeding: this was attributed to changes in the population's social network.

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Culture can be defined as "all group-typical behavior patterns, shared by members of animal communities, that are to some degree reliant on socially learned and transmitted information".

201: 734:

The results from the research of Victoria Horner and Andrew Whiten show that chimpanzee social structures and human social structures have more similarities than previously thought.

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of killer whale off Vancouver Island is attributed to differences in diet. The resident ecotype feeds on fish and a little squid, and the transient ecotype feeds on marine mammals.

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Lefebvre, Louis (1995-05-01). "The opening of milk bottles by birds: Evidence for accelerating learning rates, but against the wave-of-advance model of cultural transmission".

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Animal culture can be an important consideration in conservation management. As of 2020, culture and sociality were included in the aspects of the management framework of the

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of other 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

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Sargeant, Brooke L.; Mann, Janet (2009). "From Social Learning to Culture: Intrapopulation Variation in Bottlenose Dolphins". In Laland, Kevin N.; Galef, Bennett G. (eds.).

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Others have been able to predict the pattern information transmission among individuals based on a preexisting social network. In this study, social interactions of ravens (

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Ford, John K. B.; Notarbartolo di Sciara, Giuseppe; Simmonds, Mark P.; Spina, Fernando; Wade, Paul R.; Whitehead, Hal; Williams, James; Garland, Ellen C. (28 April 2021).

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P.; Spina, Fernando; Thornton, Alex; Wade, Paul R.; Whiting, Martin J.; Williams, James; Rendell, Luke; Whitehead, Hal; Whiten, Andrew; Rutz, Christian (8 March 2019).

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Hyland Bruno, Julia; Jarvis, Erich D.; Liberman, Mark; Tchernichovski, Ofer (14 January 2021). "Birdsong Learning and Culture: Analogies with Human Spoken Language".

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in non-humans has been a contentious subject, sometimes forcing researchers to rethink "what it is to be human". The notion of culture in other animals dates back to

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Oblique transmission occurs to offspring from the generation to which their parents belong that is, from adults other than the offspring's parents, such as teachers.

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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

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an optimal strategy for obtaining the seeds that minimized energy inputs and maximized outputs. Naïve rats that did not use this strategy could not learn it from

3144: 519:, or social learning, was a skill that was thought to be uniquely human. However, research continued through the 1990s and beyond documented the existence of 2681: 2436:

Whiten, A.; Goodall, J.; McGrew, W.C.; Nishida, T.; Reynolds, V.; Sugiyama, Y.; Tutin, C.E.G.; Wrangham, R.W.; Boesch, C. (1999). "Cultures in Chimpanzees".

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In 20th century Britain, bottled milk was delivered to households in the early morning by milkmen and left on doorsteps to be collected. Birds such as tits (

436:. Some research has suggested that teaching, as opposed to imitation, may be a characteristic of certain animals who have more advanced cultural capacities. 4808:

Marón, Carina F.; Beltramino, Lucas; Di Martino, Matías; Chirife, Andrea; Seger, Jon; Uhart, Marcela; Sironi, Mariano; Rowntree, Victoria J. (2015-10-21).

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Culture is just one source of adaptive behavior an organism exhibits to better exploit its environment. When behavioral variation reflects differential

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Evidence for cultural transmission has also been shown in wild fish populations. Scientists Helfman and Schultz conducted translocation experiments with

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Further study is being done in the matrilineal whales to uncover the cultural transmission mechanisms associated with other advanced techniques, such as

2016:

Carroll, E. L.; Baker, C. S.; Watson, M.; Alderman, R.; Bannister, J.; Gaggiotti, O. E.; Gröcke, D. R.; Patenaude, N.; Harcourt, R. (9 November 2015).

450:. This process by which offspring within a species acquires his or her own culture through mimicry or being introduced to traditions is referred to as 4184:

Terkel, Joseph (1996). "Cultural Transmission of Feeding Behavior in the Black Rat (Rattus rattus)". In Heyes, Cecelia M.; Galef, Bennett G. (eds.).

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Prior to these findings, opponents to the idea of animal culture had argued that the behaviors being called cultural were simply behaviors that had

5794: 594: 4876: 4235: 3284:"Grey Parrots Do Not Always 'Parrot': The Roles of Imitation and Phonological Awareness in the Creation of New Labels from Existing Vocalizations" 793:

when well evidenced, due to their similarity to humans in having "long lifetimes, advanced cognitive abilities, and prolonged parental care."

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Berdahl, Andrew M.; Kao, Albert B.; Flack, Andrea; Westley, Peter A. H.; Codling, Edward A.; Couzin, Iain D.; Dell, Anthony I.; Biro, Dora.

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Finally, other species of birds have been observed to conform to the personality of other individuals in their presence. Gouldian finches (

4810:"Increased Wounding of Southern Right Whale (Eubalaena australis) Calves by Kelp Gulls (Larus dominicanus) at Península Valdés, Argentina" 758:, has been studied for numerous years. In these animals, much of the evidence for culture comes from vocalizations and feeding behaviors. 2980: 913:

Though this research is fairly recent, it is often used as a prime example of evidence for culture in non-primate, non-cetacean beings.

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Lachlan, R.F.; Janik, V.M.; Slater, P.J.B. (2004). "The evolution of conformity-enforcing behaviour in cultural communication systems".

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gene transmission account for variation. Vertical transmission also contributes strongly to the buildup of between-population variation.

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Bartlett, P.; Slater, P.J.B. (1999). "The effect of new recruits on the flock specific call of budgerigars (Melopsittacus undulatus)".

3842:"Dietary specialization in two sympatric populations of killer whales ( Orcinus orca ) in coastal British Columbia and adjacent waters" 3121: 2018:"Cultural traditions across a migratory network shape the genetic structure of southern right whales around Australia and New Zealand" 4353:

West, Meredith J.; King, Andrew P. (1996). "Social Learning: Synergy and Songbirds". In Heyes, Cecelia M.; Galef, Bennett G. (eds.).

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component, seeing as these actions employed by other animals are all mechanisms for making their lives easier, and therefore longer.

5558: 3003: 2491: 868:

By using a "process of elimination" approach, researchers Krutzen et al. reported evidence of culturally transmitted tool use in

777:. Since the early 1970s, scientists have studied these four species in depth, finding potential cultural attributes within group 3378:

Huffman, Michael A.; Nahallage, Charmalie A.D.; Leca, Jean-Baptiste (2008). "Cultured Monkeys: Social Learning Cast in Stones".

3330: 303:

Though the idea of 'culture' in other animals has only been around for just over half of a century, scientists have been noting

5071:"Pathways of information transmission among wild songbirds follow experimentally imposed changes in social foraging structure" 3840:

Ford, John K B; Ellis, Graeme M; Barrett-Lennard, Lance G; Morton, Alexandra; Palm, Rod S; Balcomb, Kenneth C. (August 1998).

622:, first introduced the idea of "kaluchua" or "pre-culture" in referring to the now famous potato-washing behavior of Japanese 593:

has taken the use of the English language even further. Kanzi was taught to recognize words and their associations by using a

5701: 4513: 4362: 4334: 4218: 4193: 4011: 3955: 3776: 3574: 2726: 2560: 2506: 2411: 1823: 1733: 86: 5608: 6728: 4027:

Michael Krutzen; Janet Mann; Michael R. Heithaus; Richard C. Connor; Lars Bejder & William B. Sherwin (June 21, 2005).

58: 4496:

Riebel, Katharina; Lachlan, Robert F.; Slater, Peter J.B. (2015), "Learning and Cultural Transmission in Chaffinch Song",

2799:

Best, Michael L. (January 1999). "How Culture Can Guide Evolution: An Inquiry into Gene/Meme Enhancement and Opposition".

1839:

Gruber, Thibaud; Luncz, Lydia; Mörchen, Julia; Schuppli, Caroline; Kendal, Rachel L.; Hockings, Kimberley (18 June 2019).

4743:

Aplin, Lucy M.; Farine, Damien R.; Morand-Ferron, Julie; Cockburn, Andrew; Thornton, Alex; Sheldon, Ben C. (2014-12-03).

1042:

The complexity of several avian behaviors can be explained by the accumulation of cultural traits over many generations.

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capacity for culture and cultural transmission dates back to the now-extinct common ancestor we share with chimpanzees.

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Whiten, Andrew; Victoria Horner; Frans B M de Waal (2005). "Conformity To Cultural Norms Of Tool Use In Chimpanzees".

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Despite this hindrance, evidence for differing dialects among songbird populations has been discovered, especially in

65: 5832: 5789: 1360: 917:

may be in part cultural; released ungulates have to learn over generations the seasonal changes in local vegetation.

105: 4901:

Klump, Barbara C.; Martin, John M.; Wild, Sonja; Hörsch, Jana K.; Major, Richard E.; Aplin, Lucy M. (23 July 2021).

5382:

Aplin, Lucy M.; Farine, Damien R.; Morand-Ferron, Julie; Cockburn, Andrew; Thornton, Alex; Sheldon, Ben C. (2015).

5282:

Kulahci, Ipek G.; Rubenstein, Daniel I.; Bugnyar, Thomas; Hoppitt, William; Mikus, Nace; Schwab, Christine (2016).

1206: 365:

of cultural transmission, each of which have distinct effects on socialization: vertical, horizontal, and oblique.

3418:"On the controversy over non-human culture: The reasons for disagreement and possible directions toward consensus" 2687: 839: 821:

nucleotide diversities are about ten times lower than other species of whale. Whitehead found that this low mtDNA

6940: 1338: 1252:

diverging local traditions when different populations were seeded with birds specifically trained in one method.

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fishing for termites using a sharpened stick. Tool usage in acquiring food is believed to be a cultural behavior.

39: 2967: 2848: 2178: 72: 7244: 6379: 3980: 3655: 3480:"Professor Masao Kawai, a pioneer and leading scholar in primatology and writer of animal stories for children" 1342: 1226: 43: 2531: 666:

to make grooming of the armpits easier, the behavior actually has no apparent advantage. As the primatologist

379:, or non-vertical tranmission is cultural transmission that occurs among individuals from the same generation. 7625: 2237:

Denison, Daniel R.; Mishra, Aneil K. (1995). "Toward a Theory of Organizational Culture and Effectiveness".

2147: 193:. The lack of common ancestors suggests that the basis for vocal learning has evolved independently through 7578: 7274: 6945: 6184: 5930: 4135:"Associative Mechanisms Allow for Social Learning and Cultural Transmission of String Pulling in an Insect" 4088:"Genes or culture: are mitochondrial genes associated with tool use in bottlenose dolphins (Tursiops sp.)?" 3597:

A. Whiten; A. Spiteri; V. Horner; K.E. Bonnie; S.P. Lambeth; S.J. Schapiro; F.B.M. de Waal (19 June 2007).

3048: 2637: 2584: 605: 54: 7356: 7027: 3846: 1185: 778: 447: 5799: 5784: 1635: 858: 162:, but the association of other animals' actions with the actual word 'culture' originated with Japanese 7557: 7306: 6470: 6287: 6137: 1504: 1390: 842:

maintain unique, stable dialects separate from each other's, though they are associated and share some

520: 474: 7449: 2580: 1256:

that unhelpful traditions will not necessarily be adhered to in the presence of a better alternative.

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have been discovered to show regional "dialects," a trait that has potential to have a cultural basis.

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or from watching experienced rats. Only young offspring could learn the technique. Additionally, from

7301: 7082: 6515: 5945: 5940: 5935: 4264:"Is ungulate migration culturally transmitted? Evidence of social learning from translocated animals" 3092:"Social learning and traditions in animals: evidence, definitions, and relationship to human culture" 327: 1127: 121:

This article is about learning in non-human animals. For the place of animals in human culture, see

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Culture can be transmitted among animals through various methods, the most common of which include

194: 3143:

Hoppitt, W; Brown, G; Kendal, R; Rendell, L; Thornton, A; Webster, M; Laland, K (September 2008).

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Collective animal navigation and migratory culture: from theoretical models to empirical evidence

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Horner, Victoria; Darby Proctor; Kristin E Bonnie; Andrew Whiten & Frans B M de Waal (2010).

3096: 2958: 1331: 1202: 929: 376: 339: 283: 32: 4209:

Galef, Bennett G. (2009). "Culture in Animals?". In Laland, Kevin N.; Galef, Bennett G. (eds.).

3331:"Newly-acquired pre-cultural behavior of the natural troop of Japanese monkeys on Koshima islet" 1951:

Whitehead, Hal; Laland, Kevin N.; Rendell, Luke; Thorogood, Rose; Whiten, Andrew (3 June 2019).

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However, the presence of social bonds strongly facilitates song imitation in certain songbirds.

7388: 7351: 7341: 7286: 7234: 7179: 6788: 6535: 6404: 6252: 6245: 6210: 5997: 5925: 2849:"Experimental evidence for social transmission of food acquisition techniques in wild meerkats" 2246: 907: 3562: 2944: 1694: 7057: 6975: 6848: 6672: 6550: 6495: 6485: 6475: 6349: 6205: 6157: 6047: 6012: 5889: 5869: 5825: 4962: 4026: 3091: 1499: 1386: 1109:

A spread of new foraging behaviors also occurred in an Argentinian population of kelp gulls (

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behaviors in animals, like those observed by Imanishi, were related to survival in some way.

565: 399: 369: 354: 5384:"Experimentally induced innovations lead to persistent culture via conformity in wild birds" 4745:"Experimentally induced innovations lead to persistent culture via conformity in wild birds" 4378:

Aplin, Lucy M. (2019). "Culture and cultural evolution in birds: a review of the evidence".

79: 7494: 7259: 7219: 7214: 7052: 7022: 6930: 6920: 6858: 6838: 6767: 6560: 6545: 6505: 6399: 6032: 6027: 5460: 5395: 5295: 5139: 4914: 4902: 4877:"Cockies are learning how to bust into bins and their skills are spreading across suburbia" 4821: 4756: 4656: 4598: 4541: 4440: 4277: 4087: 4040: 3806: 3725: 3670: 2445: 2095: 2029: 1964: 1469: 1459: 707:

Whiten et al. further made sure that these local traditions were not due to differences in

548: 7504: 3653: 1597: 398:, is the process and method of passing on socially learned information. Within a species, 8: 7585: 7366: 6913: 6903: 6898: 6828: 6718: 6703: 6590: 6580: 6555: 6490: 6465: 6419: 6359: 6267: 6167: 6147: 6063: 6042: 5982: 3797:

Whitehead, Hal (1998). "Cultural Selection and Genetic Diversity in Matrilineal Whales".

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play in influencing behavioral variation from population to population within a species.

1177:), and the pattern of inheritance was shown to depend on social structures in the flock. 697: 499:), researchers found evidence that meerkats learned foraging tricks through imitation of 155: 6480: 5464: 5449:"Conformity does not perpetuate suboptimal traditions in a wild population of songbirds" 5399: 5299: 5143: 4918: 4825: 4760: 4660: 4602: 4545: 4444: 4281: 4044: 3810: 3729: 3674: 2623: 2449: 2355: 2099: 2033: 1968: 7499: 7291: 7269: 7249: 7062: 6995: 6843: 6803: 6723: 6585: 6565: 6520: 6510: 6500: 6460: 6389: 6364: 6334: 6314: 6309: 6277: 6179: 5977: 5962: 5957: 5952: 5884: 5874: 5855: 5756: 5707: 5628: 5589: 5539: 5491: 5448: 5424: 5383: 5324: 5283: 5218: 5185: 5163: 5103: 5070: 5012: 4938: 4852: 4809: 4785: 4744: 4725: 4562: 4529: 4464: 4403: 4161: 4134: 4115: 4063: 4028: 3915: 3884: 3749: 3693: 3636: 3517: 3453: 3395: 3353: 3229: 3067: 3036: 3008: 2972: 2925: 2871: 2816: 2768: 2654: 2625: 2597: 2469: 2321: 2209: 2129: 2058: 2017: 1993: 1952: 1928: 1903: 1880: 1785: 1777: 1706: 1609: 1559: 1154: 869: 762: 701: 235: 122: 5720: 5624: 5127: 4530:"A mesocortical dopamine circuit enables the cultural transmission of vocal behaviour" 4326: 2083: 1531: 234:

Other researchers are currently exploring the idea that there is a connection between

7630: 7604: 7311: 7279: 7172: 7145: 6990: 6883: 6873: 6708: 6645: 6640: 6540: 6530: 6525: 6344: 6339: 6324: 6319: 6304: 6299: 6282: 6272: 6257: 6230: 6220: 6132: 6125: 6108: 5760: 5748: 5740: 5697: 5680: 5581: 5531: 5496: 5478: 5429: 5411: 5364: 5329: 5311: 5264: 5223: 5205: 5167: 5155: 5108: 5090: 5048: 5004: 5000: 4968: 4942: 4930: 4857: 4839: 4790: 4772: 4717: 4709: 4705: 4674: 4626: 4621: 4586: 4567: 4509: 4456: 4395: 4358: 4330: 4295: 4268: 4214: 4189: 4166: 4107: 4068: 4007: 3951: 3920: 3841: 3822: 3772: 3741: 3698: 3628: 3620: 3570: 3521: 3509: 3501: 3445: 3437: 3399: 3391: 3221: 3167: 3113: 3072: 2917: 2909: 2760: 2722: 2718: 2659: 2556: 2502: 2487: 2461: 2407: 2325: 2201: 2121: 2063: 2045: 1998: 1980: 1933: 1884: 1872: 1819: 1769: 1729: 1710: 1698: 1613: 1601: 1563: 1551: 1076: 1059: 1051: 1023: 976: 818: 693: 688: 654: 533: 412: 331: 330:, contributed to the greater understanding of cultural transmission with his work on 292: 174: 140: 5632: 5593: 5543: 5016: 4729: 4468: 4407: 4119: 3457: 3357: 3233: 2976: 2929: 2875: 2820: 2772: 2213: 2133: 1841:"Cultural change in animals: a flexible behavioural adaptation to human disturbance" 1789: 441: 7564: 7509: 7376: 7209: 7121: 7007: 6960: 6908: 6888: 6863: 6798: 6755: 6713: 6677: 6660: 6650: 6625: 6615: 6610: 6605: 6439: 6394: 6384: 6240: 6225: 6162: 6103: 6068: 6007: 5972: 5967: 5894: 5879: 5818: 5732: 5672: 5620: 5573: 5523: 5486: 5468: 5419: 5403: 5356: 5319: 5303: 5254: 5213: 5197: 5184:

Farine, Damien R.; Aplin, Lucy M.; Sheldon, Ben C.; Hoppitt, William (2015-03-22).

5147: 5098: 5082: 5040: 4996: 4922: 4903:"Innovation and geographic spread of a complex foraging culture in an urban parrot" 4847: 4829: 4780: 4764: 4701: 4664: 4616: 4606: 4557: 4549: 4501: 4448: 4387: 4322: 4285: 4156: 4146: 4099: 4058: 4048: 3984: 3910: 3900: 3855: 3814: 3753: 3733: 3688: 3678: 3640: 3610: 3491: 3429: 3387: 3345: 3213: 3159: 3105: 3062: 3052: 2962: 2901: 2892:

Caro, T. M.; Hauser, M. D. (1992-06-01). "Is There Teaching in Nonhuman Animals?".

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Cetacean vocalizations have been studied for many years, specifically those of the

719: 569: 343: 5577: 5527: 5360: 5186:"Interspecific social networks promote information transmission in wild songbirds" 4452: 4391: 3818: 3057: 2867: 7519: 7459: 7444: 7429: 7414: 7383: 7336: 7097: 7092: 7017: 6878: 6833: 6823: 6750: 6745: 6682: 6655: 6630: 6600: 6595: 6575: 6570: 6354: 6329: 6262: 6120: 4834: 4151: 3683: 3433: 3319:. Developments in primatology. New York: Kluwer Academic/Plenum Publishers, 2004. 2626:"Non-vertical cultural transmission, assortment and the evolution of cooperation" 2290: 1479: 1474: 1464: 1300: 938: 934: 662: 304: 250: 5284:"Social networks predict selective observation and information spread in ravens" 3973:

Call Traditions and Dialects of Killer Whales (Orcinus Orca) in British Columbia

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Compilation of 100+ user submitted definitions of culture from around the globe

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J. Terkel (1996). "Cultural transmission in the black rat: pine cone feeding".

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Cohen, Dor; Lewin-Epstein, Ohad; Feldman, Marcus W.; Ram, Yoav (May 26, 2021).

2197: 1976: 1904:"A deepening understanding of animal culture suggests lessons for conservation" 1494: 1445: 1277: 1265: 1197: 903: 766: 619: 524: 361: 357: 258: 170: 159: 5774: 4972: 4553: 4103: 3715: 3615: 3598: 3217: 2317: 1857: 1840: 1781: 1648: 1140: 649: 556:. Imitation forms the basis of culture, but does not on its own imply culture. 552:

Imitation can be found in a few members of the avian world, in particular the

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The likelihood of larger groups within a species developing and sharing these

7619: 7489: 7479: 7474: 7469: 7398: 7331: 7239: 7102: 7087: 7072: 7047: 7037: 6965: 6925: 6793: 6665: 6429: 6409: 6374: 6294: 6235: 6215: 6200: 6174: 6088: 6037: 5744: 5684: 5585: 5535: 5482: 5415: 5368: 5315: 5268: 5209: 5094: 5052: 5008: 4843: 4776: 4713: 4678: 4399: 3624: 3505: 3441: 2913: 2049: 1984: 1876: 1773: 1702: 1605: 1173: 972: 959: 782: 667: 452: 403: 323: 5736: 5473: 5151: 5044: 4926: 4611: 4290: 4263: 4053: 2107: 1547: 985: 7514: 7464: 7424: 7393: 7346: 7067: 7012: 7000: 6985: 6955: 6868: 6808: 6414: 6098: 6078: 6073: 6022: 5752: 5711: 5500: 5433: 5333: 5227: 5201: 5159: 5112: 5086: 4934: 4861: 4794: 4721: 4571: 4460: 4299: 4240: 4170: 4111: 4072: 3924: 3745: 3702: 3632: 3513: 3449: 3225: 3171: 3117: 3076: 2764: 2663: 2645: 2465: 2387: 2370: 2205: 2125: 2067: 2002: 1937: 1919: 1555: 1509: 1484: 1439: 1289: 1235: 1012: 814: 810: 770: 704:) were found to be habitual in some communities but nonexistent in others. 222: 4630: 4528:

Tanaka, Masashi; Sun, Fangmiao; Li, Yulong; Mooney, Richard (1 Nov 2018).

3826: 3599:"Transmission of Multiple Traditions within and between Chimpanzee Groups" 2921: 7454: 7419: 7077: 6980: 6950: 6635: 6093: 5992: 5899: 2260: 1403: 1088: 1081: 806: 802: 774: 642: 500: 163: 5407: 5307: 5259: 5242: 5029: 4768: 4213:. Cambridge, Massachusetts: Harvard University Press. pp. 222–246. 3905: 3737: 1680: 1037: 633:

The most famous of these eating behaviors was observed on the island of

6142: 5920: 5676: 4085: 3988: 3771:. Cambridge, Massachusetts: Harvard University Press. pp. 152–73. 3596: 3349: 2949: 2606: 2548: 2116: 1867: 1752:

Sapolsky, Robert M. (2006). "Social Cultures among Nonhuman Primates".

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Second only to non-human primates, culture in species within the order

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argues for the existence of a "unit of cultural transmission" called a

239: 4669: 4644: 4587:"Social inhibition of song imitation among sibling male zebra finches" 2945:"Social Learning in Animals: Empirical Studies and Theoretical Models" 2304:

Holdcroft, David; Lewis, Harry (2000). "Memes, Minds, and Evolution".

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Durham, William H (1990). "Advances in Evolutionary Culture Theory".

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The Ape and the Sushi Master: Cultural Reflections by a Primatologist

1158: 1141:"Cockatoos learn how to flip open garbage bins by copying each other" 1110: 1016: 921: 755: 675: 560: 480: 433: 308: 271: 151: 5609:"Social transmission of behavioural traditions in a coral reef fish" 5557:

King, Andrew J.; Williams, Leah J.; Mettke-Hofmann, Claudia (2015).

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Animal Bodies, Human Minds: Ape, Dolphin, and Parrot Language Skills

1320: 21: 7326: 7321: 7296: 7195: 5694:

Social Learning: An Introduction to Mechanisms, Methods, and Models

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Whiten, Andrew (2 November 2019). "Cultural Evolution in Animals".

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Convention on the Conservation of Migratory Species of Wild Animals

182: 136: 5447:

Aplin, Lucy M.; Sheldon, Ben C.; McElreath, Richard (2017-07-24).

4086:

K. Bacher; S. Allen; A.K. Lindholm; L. Bejder; M. Krutzen (2010).

3537:"What Japan's Wild Snow Monkeys Can Teach Us About Animal Culture" 2709:

Jones, Nick A. R.; Rendell, Luke (2018). "Cultural Transmission".

2457: 7361: 5841: 5128:"Social Transmission of a Host Defense Against Cuckoo Parasitism" 2942: 1489: 1375: 1069: 1055: 1031: 857:

Further information on southern resident orca vocal culture:

831: 743: 708: 634: 623: 492: 425: 416: 408: 386: 147: 5559:"The effects of social conformity on Gouldian finch personality" 5381: 4807: 4742: 4006:. Friday Harbor, Washington, USA: Orca Watcher. pp. 80–81. 2148:"Animal Culture Linked to Conservation for the First Time | CMS" 326:, professor of Evolutionary and Developmental Psychology at the 6152: 5281: 4426:"Culturally transmitted patterns of vocal behavior in sparrows" 3938:

Ford, John K.B.; Ellis, Graeme M.; Balcomb, Kenneth C. (2000).

1950: 747: 610: 586: 553: 178: 132: 3995: 3878: 3876: 3839: 512:

observer must acquire skills rapidly as a direct consequence.

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Perspectives on Imitation From Neuroscience to Social Science

3004:"Monkeys Are Adept at Picking Up Social Cues, Research Shows" 2492:"31. From material to symbolic cultures: Culture in primates" 1421: 1281: 1015:, who noted the geographic variation in the songs of various 1008: 590: 190: 3253: 7371: 7164: 5804: 5243:"Interspecific information use by army-ant–following birds" 5069:

Firth, Josh A.; Sheldon, Ben C.; Farine, Damien R. (2016).

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Archived from 4439:(3650): 1483–6. 4430: 4421: 4412: 4411: 4380:Animal Behaviour 4375: 4369: 4368: 4350: 4341: 4340: 4310: 4304: 4303: 4293: 4259: 4253: 4252: 4250: 4248: 4231: 4225: 4224: 4206: 4200: 4199: 4181: 4175: 4174: 4164: 4154: 4145:(14): e1002564. 4130: 4124: 4123: 4083: 4077: 4076: 4066: 4056: 4024: 4018: 4017: 3999: 3993: 3992: 3968: 3962: 3961: 3935: 3929: 3928: 3918: 3908: 3880: 3871: 3870: 3868: 3866: 3854:(8): 1456–1471. 3837: 3831: 3830: 3794: 3783: 3782: 3764: 3758: 3757: 3713: 3707: 3706: 3696: 3686: 3660: 3651: 3645: 3644: 3618: 3594: 3588: 3587: 3585: 3583: 3558: 3552: 3551: 3549: 3547: 3532: 3526: 3525: 3499: 3475: 3469: 3468: 3466: 3464: 3413: 3404: 3403: 3375: 3369: 3368: 3366: 3364: 3335: 3326: 3320: 3313: 3300: 3299: 3297: 3295: 3286:. Archived from 3279: 3273: 3266: 3260: 3259: 3251: 3245: 3244: 3242: 3240: 3203: 3194: 3183: 3182: 3180: 3178: 3149: 3140: 3134: 3133: 3131: 3129: 3110:10.1002/wcs.1196 3087: 3081: 3080: 3070: 3060: 3032: 3026: 3025: 3023: 3021: 2999: 2993: 2992: 2990: 2988: 2970: 2940: 2934: 2933: 2889: 2880: 2879: 2856:Animal Behaviour 2853: 2844: 2838: 2831: 2825: 2824: 2807:(3–4): 289–306. 2796: 2790: 2783: 2777: 2776: 2739: 2733: 2732: 2706: 2700: 2699: 2697: 2695: 2677: 2668: 2667: 2657: 2621: 2612: 2611: 2609: 2578: 2567: 2566: 2545: 2536: 2535: 2530:. Archived from 2519: 2513: 2512: 2496: 2484: 2478: 2477: 2433: 2418: 2417: 2399: 2393: 2392: 2390: 2366: 2360: 2359: 2339: 2330: 2329: 2312:(292): 161–182. 2301: 2295: 2294: 2274: 2265: 2264: 2254: 2234: 2225: 2224: 2222: 2220: 2183: 2174: 2163: 2162: 2160: 2158: 2144: 2138: 2137: 2119: 2078: 2072: 2071: 2061: 2013: 2007: 2006: 1996: 1948: 1942: 1941: 1931: 1898: 1889: 1888: 1870: 1860: 1836: 1830: 1829: 1811: 1794: 1793: 1749: 1740: 1739: 1721: 1715: 1714: 1678: 1661: 1660: 1658: 1656: 1651: 1631: 1618: 1617: 1581: 1575: 1574: 1572: 1570: 1527: 1455:Animal cognition 1448: 1443: 1442: 1366: 1359: 1355: 1352: 1346: 1323: 1315: 1139: 1138: 1130: 1118: 1117: 1002: 971: 970: 962: 950: 949: 930:optimal foraging 915:Animal migration 738:Cetacean culture 720:social structure 657:mother and baby. 570:Irene Pepperberg 394:, also known as 305:social behaviors 111: 104: 100: 97: 91: 89: 55:"Animal culture" 48: 24: 16: 7646: 7645: 7641: 7640: 7639: 7637: 7636: 7635: 7616: 7615: 7614: 7609: 7591: 7545: 7524: 7520:Solly Zuckerman 7460:Karl von Frisch 7445:Richard Dawkins 7430:John B. Calhoun 7415:Patrick Bateson 7403: 7337:Pain in animals 7198: 7193: 7163: 7158: 7107: 7098:Western culture 7093:Welfare culture 7018:Eastern culture 6879:Cultural mosaic 6834:Cultural critic 6824:Cultural center 6772: 6746:Cultural Hindus 6692: 6683:Polyculturalism 6656:Monoculturalism 6631:Culture of fear 6601:Cultural safety 6596:Cultural rights 6576:Cultural racism 6571:Cultural policy 6449: 6355:Cultural system 6330:Cultural memory 6263:Cultural cringe 6189: 6121:Popular culture 6052: 5988:Cultural values 5909: 5858: 5844: 5839: 5771: 5704: 5655: 5653:Further reading 5650: 5649: 5644: 5640: 5605: 5601: 5561: 5555: 5551: 5512: 5508: 5445: 5441: 5380: 5376: 5345: 5341: 5280: 5276: 5239: 5235: 5182: 5175: 5124: 5120: 5081:(6): 20160144. 5075:Biology Letters 5067: 5060: 5028: 5024: 4984: 4980: 4961: 4957: 4947: 4945: 4899: 4895: 4885: 4883: 4873: 4869: 4806: 4802: 4741: 4737: 4690: 4686: 4643: 4642: 4638: 4583: 4579: 4526: 4522: 4516: 4494: 4490: 4481: 4479: 4475: 4428: 4422: 4415: 4376: 4372: 4365: 4351: 4344: 4337: 4311: 4307: 4260: 4256: 4246: 4244: 4232: 4228: 4221: 4207: 4203: 4196: 4182: 4178: 4131: 4127: 4084: 4080: 4025: 4021: 4014: 4000: 3996: 3983:. p. 287. 3969: 3965: 3958: 3936: 3932: 3889:Aquatic Biology 3881: 3874: 3864: 3862: 3860:10.1139/z98-089 3838: 3834: 3795: 3786: 3779: 3765: 3761: 3714: 3710: 3658: 3652: 3648: 3603:Current Biology 3595: 3591: 3581: 3579: 3577: 3559: 3555: 3545: 3543: 3533: 3529: 3476: 3472: 3462: 3460: 3414: 3407: 3376: 3372: 3362: 3360: 3333: 3327: 3323: 3314: 3303: 3293: 3291: 3280: 3276: 3267: 3263: 3258:(4th ed.). 3252: 3248: 3238: 3236: 3201: 3195: 3186: 3176: 3174: 3147: 3141: 3137: 3127: 3125: 3088: 3084: 3033: 3029: 3019: 3017: 3000: 2996: 2986: 2984: 2941: 2937: 2890: 2883: 2851: 2845: 2841: 2832: 2828: 2797: 2793: 2784: 2780: 2740: 2736: 2729: 2707: 2703: 2693: 2691: 2678: 2671: 2622: 2615: 2579: 2570: 2563: 2546: 2539: 2520: 2516: 2509: 2494: 2485: 2481: 2434: 2421: 2414: 2400: 2396: 2367: 2363: 2340: 2333: 2302: 2298: 2275: 2268: 2235: 2228: 2218: 2216: 2192:(10): 542–547. 2181: 2175: 2166: 2156: 2154: 2146: 2145: 2141: 2079: 2075: 2014: 2010: 1949: 1945: 1899: 1892: 1837: 1833: 1826: 1812: 1797: 1750: 1743: 1736: 1722: 1718: 1679: 1664: 1654: 1652: 1632: 1621: 1582: 1578: 1568: 1566: 1528: 1524: 1519: 1514: 1480:Animal training 1475:Animal-made art 1465:Animal language 1444: 1437: 1434: 1417: 1367: 1356: 1350: 1347: 1336: 1324: 1313: 1275: 1245: 1198:Social networks 1195: 1183: 1168: 1147:, July 22, 2021 1136: 1121:External videos 1098: 1048: 1040: 997: 968: 953:External videos 948: 939:cross-fostering 935:trial and error 908:social learning 896: 866: 861: 799: 740: 685: 663:social grooming 603: 601:Primate culture 579: 546: 509: 477: 471: 469:Social learning 462: 389: 352: 321: 301: 280: 267: 251:Richard Dawkins 248: 232: 218: 210: 126: 119: 112: 101: 95: 92: 49: 47: 37: 25: 12: 11: 5: 7644: 7634: 7633: 7628: 7611: 7610: 7608: 7607: 7596: 7593: 7592: 7590: 7589: 7582: 7575: 7572:Animal Welfare 7568: 7561: 7553: 7551: 7547: 7546: 7544: 7543: 7538: 7532: 7530: 7526: 7525: 7523: 7522: 7517: 7512: 7507: 7502: 7497: 7492: 7487: 7485:Desmond Morris 7482: 7477: 7472: 7467: 7462: 7457: 7452: 7447: 7442: 7440:Marian Dawkins 7437: 7435:Charles Darwin 7432: 7427: 7422: 7417: 7411: 7409: 7405: 7404: 7402: 7401: 7396: 7391: 7386: 7381: 7380: 7379: 7374: 7369: 7364: 7354: 7349: 7344: 7339: 7334: 7329: 7324: 7319: 7317:Human ethology 7314: 7309: 7304: 7299: 7294: 7289: 7284: 7283: 7282: 7272: 7267: 7262: 7257: 7252: 7247: 7242: 7237: 7232: 7227: 7225:Animal culture 7222: 7217: 7212: 7206: 7204: 7200: 7199: 7192: 7191: 7184: 7177: 7169: 7160: 7159: 7157: 7156: 7149: 7137: 7125: 7112: 7109: 7108: 7106: 7105: 7100: 7095: 7090: 7085: 7080: 7075: 7070: 7065: 7060: 7055: 7050: 7045: 7040: 7035: 7030: 7025: 7020: 7015: 7010: 7005: 7004: 7003: 6993: 6988: 6983: 6978: 6973: 6968: 6963: 6958: 6953: 6948: 6943: 6938: 6933: 6928: 6923: 6918: 6917: 6916: 6906: 6901: 6896: 6894:Cultural probe 6891: 6886: 6881: 6876: 6871: 6866: 6861: 6856: 6851: 6846: 6841: 6836: 6831: 6826: 6821: 6819:Cross-cultural 6816: 6814:Coffee culture 6811: 6806: 6801: 6796: 6791: 6786: 6784:Animal culture 6780: 6778: 6774: 6773: 6771: 6770: 6765: 6760: 6759: 6758: 6748: 6743: 6742: 6741: 6736: 6731: 6726: 6721: 6716: 6706: 6700: 6698: 6694: 6693: 6691: 6690: 6688:Transculturism 6685: 6680: 6675: 6670: 6669: 6668: 6658: 6653: 6648: 6643: 6638: 6633: 6628: 6623: 6621:Culture change 6618: 6613: 6608: 6603: 6598: 6593: 6588: 6583: 6578: 6573: 6568: 6563: 6558: 6553: 6548: 6543: 6538: 6533: 6528: 6523: 6518: 6513: 6508: 6503: 6498: 6493: 6488: 6483: 6478: 6473: 6468: 6463: 6457: 6455: 6451: 6450: 6448: 6447: 6445:Visual culture 6442: 6437: 6432: 6427: 6425:Safety culture 6422: 6417: 6412: 6407: 6402: 6397: 6392: 6387: 6382: 6377: 6372: 6367: 6362: 6357: 6352: 6347: 6342: 6337: 6332: 6327: 6322: 6317: 6312: 6307: 6302: 6297: 6292: 6291: 6290: 6280: 6275: 6270: 6265: 6260: 6255: 6250: 6249: 6248: 6246:Cross-cultural 6238: 6233: 6228: 6223: 6218: 6213: 6208: 6203: 6197: 6195: 6191: 6190: 6188: 6187: 6182: 6177: 6172: 6171: 6170: 6165: 6160: 6155: 6150: 6140: 6135: 6130: 6129: 6128: 6118: 6117: 6116: 6106: 6101: 6096: 6091: 6086: 6081: 6076: 6071: 6066: 6060: 6058: 6054: 6053: 6051: 6050: 6045: 6040: 6035: 6030: 6025: 6020: 6015: 6010: 6005: 6000: 5995: 5990: 5985: 5980: 5975: 5970: 5965: 5960: 5955: 5950: 5949: 5948: 5943: 5938: 5933: 5923: 5917: 5915: 5911: 5910: 5908: 5907: 5905:Culture theory 5902: 5897: 5892: 5887: 5882: 5877: 5872: 5866: 5864: 5860: 5859: 5849: 5846: 5845: 5838: 5837: 5830: 5823: 5815: 5809: 5808: 5805:Define Culture 5802: 5797: 5792: 5787: 5785:Animal Culture 5782: 5777: 5770: 5769:External links 5767: 5766: 5765: 5716: 5702: 5689: 5654: 5651: 5648: 5647: 5638: 5619:(2): 379–384. 5599: 5549: 5522:(3): 561–570. 5506: 5439: 5374: 5355:(2): 139–147. 5339: 5274: 5253:(1): 247–255. 5233: 5173: 5118: 5058: 5022: 4995:(4): 301–308. 4978: 4955: 4893: 4867: 4800: 4735: 4684: 4636: 4597:(15): 8951–6. 4577: 4520: 4514: 4488: 4413: 4370: 4363: 4342: 4335: 4305: 4254: 4226: 4219: 4201: 4194: 4176: 4125: 4078: 4019: 4012: 3994: 3963: 3956: 3930: 3872: 3832: 3784: 3777: 3759: 3708: 3646: 3589: 3575: 3553: 3527: 3490:(5): 677–695. 3470: 3405: 3386:(6): 410–414. 3370: 3321: 3301: 3274: 3261: 3246: 3212:(2): 309–324. 3184: 3158:(9): 486–493. 3135: 3104:(6): 581–592. 3082: 3027: 2994: 2935: 2906:10.1086/417553 2900:(2): 151–174. 2881: 2862:(2): 255–264. 2839: 2826: 2791: 2778: 2734: 2727: 2701: 2669: 2613: 2592:(4): 274-285. 2568: 2561: 2549:Pagel, Mark D. 2537: 2514: 2507: 2479: 2419: 2412: 2394: 2381:(3): 845–855. 2361: 2331: 2296: 2266: 2245:(2): 204–223. 2226: 2164: 2139: 2073: 2008: 1943: 1890: 1831: 1824: 1795: 1782:10.1086/504162 1766:10.1086/504162 1760:(4): 641–656. 1741: 1734: 1716: 1689:(1): 449–472. 1662: 1619: 1576: 1521: 1520: 1518: 1515: 1513: 1512: 1507: 1502: 1497: 1495:Culture theory 1492: 1487: 1482: 1477: 1472: 1467: 1462: 1457: 1451: 1450: 1449: 1446:Animals portal 1433: 1430: 1416: 1413: 1369: 1368: 1327: 1325: 1318: 1312: 1309: 1274: 1271: 1244: 1241: 1194: 1191: 1182: 1179: 1167: 1164: 1149: 1148: 1132: 1131: 1123: 1122: 1097: 1094: 1047: 1044: 1039: 1036: 981: 980: 964: 963: 955: 954: 947: 944: 904:Norwegian rats 895: 892: 881:mitochondrial 865: 862: 798: 795: 767:humpback whale 739: 736: 684: 681: 676:evolutionarily 620:Kinji Imanishi 602: 599: 595:lexigram board 578: 575: 545: 542: 525:tandem running 508: 505: 473:Main article: 470: 467: 461: 458: 388: 385: 384: 383: 380: 374: 358:Cavalli-Sforza 351: 348: 320: 314: 300: 297: 279: 276: 266: 263: 259:Daniel Dennett 247: 244: 231: 228: 217: 214: 209: 206: 171:vocal learning 164:primatologists 160:Charles Darwin 129:Animal culture 117: 114: 113: 28: 26: 19: 9: 6: 4: 3: 2: 7643: 7632: 7629: 7627: 7624: 7623: 7621: 7606: 7598: 7597: 7594: 7588: 7587: 7583: 7581: 7580: 7576: 7574: 7573: 7569: 7567: 7566: 7562: 7560: 7559: 7555: 7554: 7552: 7548: 7542: 7539: 7537: 7534: 7533: 7531: 7527: 7521: 7518: 7516: 7513: 7511: 7508: 7506: 7503: 7501: 7498: 7496: 7493: 7491: 7490:Thomas Sebeok 7488: 7486: 7483: 7481: 7480:Konrad Lorenz 7478: 7476: 7475:Julian Huxley 7473: 7471: 7470:Heini Hediger 7468: 7466: 7463: 7461: 7458: 7456: 7453: 7451: 7448: 7446: 7443: 7441: 7438: 7436: 7433: 7431: 7428: 7426: 7423: 7421: 7418: 7416: 7413: 7412: 7410: 7406: 7400: 7399:Zoomusicology 7397: 7395: 7392: 7390: 7387: 7385: 7382: 7378: 7375: 7373: 7370: 7368: 7365: 7363: 7360: 7359: 7358: 7355: 7353: 7350: 7348: 7345: 7343: 7340: 7338: 7335: 7333: 7332:Neuroethology 7330: 7328: 7325: 7323: 7320: 7318: 7315: 7313: 7310: 7308: 7305: 7303: 7300: 7298: 7295: 7293: 7290: 7288: 7285: 7281: 7278: 7277: 7276: 7273: 7271: 7268: 7266: 7263: 7261: 7258: 7256: 7253: 7251: 7248: 7246: 7243: 7241: 7240:Anthrozoology 7238: 7236: 7233: 7231: 7228: 7226: 7223: 7221: 7218: 7216: 7213: 7211: 7208: 7207: 7205: 7201: 7197: 7190: 7185: 7183: 7178: 7176: 7171: 7170: 7167: 7155: 7154: 7150: 7148: 7147: 7138: 7136: 7135: 7126: 7124: 7123: 7114: 7113: 7110: 7104: 7103:Youth culture 7101: 7099: 7096: 7094: 7091: 7089: 7088:Urban culture 7086: 7084: 7081: 7079: 7076: 7074: 7073:Remix culture 7071: 7069: 7066: 7064: 7061: 7059: 7056: 7054: 7051: 7049: 7048:Media culture 7046: 7044: 7041: 7039: 7038:Languaculture 7036: 7034: 7031: 7029: 7026: 7024: 7021: 7019: 7016: 7014: 7011: 7009: 7006: 7002: 6999: 6998: 6997: 6994: 6992: 6989: 6987: 6984: 6982: 6979: 6977: 6974: 6972: 6969: 6967: 6966:Culture shock 6964: 6962: 6959: 6957: 6954: 6952: 6949: 6947: 6944: 6942: 6939: 6937: 6934: 6932: 6929: 6927: 6926:Cultural turn 6924: 6922: 6919: 6915: 6912: 6911: 6910: 6907: 6905: 6902: 6900: 6897: 6895: 6892: 6890: 6887: 6885: 6882: 6880: 6877: 6875: 6872: 6870: 6867: 6865: 6862: 6860: 6857: 6855: 6852: 6850: 6847: 6845: 6842: 6840: 6837: 6835: 6832: 6830: 6827: 6825: 6822: 6820: 6817: 6815: 6812: 6810: 6807: 6805: 6802: 6800: 6797: 6795: 6794:Bennett scale 6792: 6790: 6787: 6785: 6782: 6781: 6779: 6775: 6769: 6766: 6764: 6761: 6757: 6754: 6753: 6752: 6749: 6747: 6744: 6740: 6737: 6735: 6732: 6730: 6727: 6725: 6724:Protestantism 6722: 6720: 6717: 6715: 6712: 6711: 6710: 6707: 6705: 6702: 6701: 6699: 6695: 6689: 6686: 6684: 6681: 6679: 6676: 6674: 6671: 6667: 6666:Biculturalism 6664: 6663: 6662: 6659: 6657: 6654: 6652: 6649: 6647: 6644: 6642: 6639: 6637: 6634: 6632: 6629: 6627: 6624: 6622: 6619: 6617: 6614: 6612: 6609: 6607: 6604: 6602: 6599: 6597: 6594: 6592: 6589: 6587: 6584: 6582: 6579: 6577: 6574: 6572: 6569: 6567: 6564: 6562: 6559: 6557: 6554: 6552: 6549: 6547: 6544: 6542: 6539: 6537: 6534: 6532: 6529: 6527: 6524: 6522: 6519: 6517: 6514: 6512: 6509: 6507: 6504: 6502: 6499: 6497: 6494: 6492: 6489: 6487: 6484: 6482: 6479: 6477: 6474: 6472: 6469: 6467: 6464: 6462: 6459: 6458: 6456: 6452: 6446: 6443: 6441: 6438: 6436: 6433: 6431: 6430:Technoculture 6428: 6426: 6423: 6421: 6418: 6416: 6413: 6411: 6410:Print culture 6408: 6406: 6403: 6401: 6398: 6396: 6393: 6391: 6388: 6386: 6383: 6381: 6378: 6376: 6375:Enculturation 6373: 6371: 6368: 6366: 6363: 6361: 6358: 6356: 6353: 6351: 6348: 6346: 6343: 6341: 6338: 6336: 6333: 6331: 6328: 6326: 6323: 6321: 6318: 6316: 6313: 6311: 6308: 6306: 6303: 6301: 6298: 6296: 6295:Cultural icon 6293: 6289: 6286: 6285: 6284: 6281: 6279: 6276: 6274: 6271: 6269: 6266: 6264: 6261: 6259: 6256: 6254: 6251: 6247: 6244: 6243: 6242: 6239: 6237: 6236:Cultural bias 6234: 6232: 6229: 6227: 6224: 6222: 6219: 6217: 6216:Cultural area 6214: 6212: 6209: 6207: 6204: 6202: 6201:Acculturation 6199: 6198: 6196: 6192: 6186: 6183: 6181: 6178: 6176: 6175:Super culture 6173: 6169: 6166: 6164: 6161: 6159: 6156: 6154: 6151: 6149: 6146: 6145: 6144: 6141: 6139: 6136: 6134: 6131: 6127: 6124: 6123: 6122: 6119: 6115: 6112: 6111: 6110: 6107: 6105: 6102: 6100: 6097: 6095: 6092: 6090: 6089:Legal culture 6087: 6085: 6082: 6080: 6077: 6075: 6072: 6070: 6067: 6065: 6062: 6061: 6059: 6055: 6049: 6046: 6044: 6041: 6039: 6038:Sound culture 6036: 6034: 6031: 6029: 6026: 6024: 6021: 6019: 6016: 6014: 6011: 6009: 6006: 6004: 6001: 5999: 5996: 5994: 5991: 5989: 5986: 5984: 5981: 5979: 5976: 5974: 5971: 5969: 5966: 5964: 5961: 5959: 5956: 5954: 5951: 5947: 5944: 5942: 5939: 5937: 5934: 5932: 5929: 5928: 5927: 5924: 5922: 5919: 5918: 5916: 5912: 5906: 5903: 5901: 5898: 5896: 5893: 5891: 5888: 5886: 5883: 5881: 5878: 5876: 5873: 5871: 5868: 5867: 5865: 5861: 5857: 5847: 5843: 5836: 5831: 5829: 5824: 5822: 5817: 5816: 5813: 5806: 5803: 5801: 5798: 5796: 5793: 5791: 5788: 5786: 5783: 5781: 5778: 5776: 5773: 5772: 5762: 5758: 5754: 5750: 5746: 5742: 5738: 5734: 5730: 5726: 5722: 5717: 5713: 5709: 5705: 5699: 5695: 5690: 5686: 5682: 5678: 5674: 5670: 5666: 5662: 5657: 5656: 5642: 5634: 5630: 5626: 5622: 5618: 5614: 5610: 5603: 5595: 5591: 5587: 5583: 5579: 5575: 5571: 5567: 5560: 5553: 5545: 5541: 5537: 5533: 5529: 5525: 5521: 5517: 5510: 5502: 5498: 5493: 5488: 5484: 5480: 5475: 5470: 5466: 5462: 5458: 5454: 5450: 5443: 5435: 5431: 5426: 5421: 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4623: 4618: 4613: 4608: 4604: 4600: 4596: 4592: 4588: 4581: 4573: 4569: 4564: 4559: 4555: 4551: 4547: 4543: 4539: 4535: 4531: 4524: 4517: 4511: 4507: 4503: 4499: 4492: 4478:on 2017-01-07 4474: 4470: 4466: 4462: 4458: 4454: 4450: 4446: 4442: 4438: 4434: 4427: 4420: 4418: 4409: 4405: 4401: 4397: 4393: 4389: 4385: 4381: 4374: 4366: 4360: 4356: 4349: 4347: 4338: 4332: 4328: 4324: 4320: 4316: 4309: 4301: 4297: 4292: 4287: 4283: 4279: 4275: 4271: 4270: 4265: 4258: 4243: 4242: 4237: 4230: 4222: 4216: 4212: 4205: 4197: 4191: 4187: 4180: 4172: 4168: 4163: 4158: 4153: 4148: 4144: 4140: 4136: 4129: 4121: 4117: 4113: 4109: 4105: 4101: 4098:(5): 706–14. 4097: 4093: 4089: 4082: 4074: 4070: 4065: 4060: 4055: 4050: 4046: 4042: 4038: 4034: 4030: 4023: 4015: 4009: 4005: 3998: 3990: 3986: 3982: 3978: 3974: 3967: 3959: 3953: 3949: 3945: 3941: 3934: 3926: 3922: 3917: 3912: 3907: 3902: 3898: 3894: 3890: 3886: 3879: 3877: 3861: 3857: 3853: 3849: 3848: 3843: 3836: 3828: 3824: 3820: 3816: 3812: 3808: 3804: 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1365: 1362: 1354: 1351:February 2022 1344: 1340: 1334: 1333: 1328:This section 1326: 1322: 1317: 1316: 1308: 1306: 1302: 1297: 1295: 1291: 1290:French grunts 1283: 1279: 1270: 1268: 1267: 1261: 1257: 1253: 1249: 1240: 1238: 1237: 1231: 1228: 1223: 1220: 1216: 1210: 1208: 1204: 1199: 1190: 1187: 1178: 1176: 1175: 1163: 1160: 1156: 1146: 1142: 1133: 1129: 1124: 1119: 1116: 1114: 1113: 1107: 1105: 1104: 1093: 1090: 1089:Zebra finches 1085: 1083: 1078: 1073: 1071: 1067: 1066: 1061: 1057: 1053: 1043: 1035: 1033: 1029: 1025: 1020: 1018: 1014: 1010: 992: 989:The songs of 987: 978: 974: 965: 961: 956: 951: 946:Avian culture 943: 942:transmitted. 940: 936: 931: 927: 926:Rattus rattus 923: 918: 916: 911: 909: 905: 901: 891: 887: 884: 878: 875: 871: 860: 855: 853: 848: 845: 841: 835: 833: 829: 824: 820: 816: 815:killer whales 812: 808: 804: 794: 790: 788: 784: 783:Hal Whitehead 780: 776: 772: 768: 764: 759: 757: 753: 749: 745: 735: 732: 728: 724: 721: 716: 712: 710: 705: 703: 699: 695: 690: 680: 677: 672: 669: 668:Frans de Waal 664: 656: 651: 647: 644: 640: 636: 631: 627: 625: 621: 612: 607: 598: 596: 592: 588: 583: 574: 571: 567: 562: 555: 550: 541: 537: 535: 532:teaching and 530: 526: 522: 518: 513: 504: 502: 498: 494: 490: 486: 482: 476: 466: 457: 455: 454: 453:enculturation 449: 444: 443: 442:intra-species 437: 435: 431: 427: 423: 421: 418: 414: 410: 405: 401: 397: 393: 381: 378: 375: 371: 368: 367: 366: 363: 359: 356: 347: 345: 341: 337: 333: 329: 325: 324:Andrew Whiten 319: 313: 310: 306: 296: 294: 289: 285: 275: 273: 262: 260: 256: 252: 243: 241: 237: 227: 224: 213: 205: 203: 198: 196: 192: 188: 184: 180: 176: 172: 167: 165: 161: 157: 153: 149: 144: 142: 138: 134: 130: 124: 110: 107: 99: 88: 85: 81: 78: 74: 71: 67: 64: 60: 57: – 56: 52: 51:Find sources: 45: 41: 35: 34: 29:This article 27: 23: 18: 17: 7584: 7577: 7570: 7563: 7556: 7515:E. O. Wilson 7465:Jane Goodall 7425:Donald Broom 7394:Zoosemiotics 7347:Sociobiology 7224: 7151: 7144: 7132: 7120: 7068:Rape culture 7013:Drug culture 7001:Deaf culture 6986:Cyberculture 6956:Culture hero 6869:Cultural lag 6809:Civilization 6783: 6709:Christianity 6415:Protoculture 6099:Microculture 6079:High culture 6074:Folk culture 6023:Postcritique 5728: 5724: 5693: 5668: 5664: 5660: 5641: 5616: 5612: 5602: 5569: 5565: 5552: 5519: 5515: 5509: 5456: 5452: 5442: 5391: 5387: 5377: 5352: 5348: 5342: 5291: 5287: 5277: 5250: 5246: 5236: 5193: 5189: 5135: 5131: 5121: 5078: 5074: 5036: 5032: 5025: 4992: 4988: 4981: 4964: 4958: 4946:. Retrieved 4910: 4906: 4896: 4884:. Retrieved 4880: 4870: 4817: 4813: 4803: 4752: 4748: 4738: 4700:(1): 43–53. 4697: 4693: 4687: 4652: 4648: 4639: 4594: 4590: 4580: 4537: 4533: 4523: 4497: 4491: 4480:. Retrieved 4473:the original 4436: 4432: 4383: 4379: 4373: 4354: 4318: 4314: 4308: 4273: 4267: 4257: 4245:. 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Retrieved 1539: 1535: 1525: 1510:Sociobiology 1485:Anthropology 1425: 1418: 1409: 1396: 1384: 1372: 1357: 1348: 1337:Please help 1332:verification 1329: 1304: 1298: 1293: 1287: 1273:Fish culture 1264: 1262: 1258: 1254: 1250: 1246: 1236:Corvus corax 1234: 1232: 1224: 1211: 1196: 1184: 1172: 1169: 1152: 1111: 1108: 1102: 1099: 1086: 1074: 1063: 1049: 1041: 1021: 1013:Peter Marler 1007: 925: 919: 912: 897: 888: 879: 873: 867: 849: 844:pulsed calls 836: 811:sperm whales 807:pilot whales 800: 791: 785:, a leading 771:killer whale 760: 741: 733: 729: 725: 717: 713: 706: 686: 673: 660: 638: 632: 628: 616: 580: 559: 538: 514: 510: 501:conspecifics 496: 478: 463: 451: 440: 438: 424: 395: 391: 390: 353: 335: 322: 317: 302: 281: 268: 249: 233: 223:organization 219: 211: 199: 168: 145: 128: 127: 102: 93: 83: 76: 69: 62: 50: 38:Please help 33:verification 30: 7455:Dian Fossey 7420:Marc Bekoff 7408:Ethologists 7146:WikiProject 7078:Tea culture 6981:Culturalism 6951:Culture gap 6914:Pop-culture 6714:Catholicism 6636:Culture war 6094:Low culture 5993:Culturomics 5900:Culturology 5712:j.ctt2jc8mh 5677:2268/162713 5671:: 123–134. 4948:24 February 4886:24 February 4386:: 179–187. 4321:: 119–154. 4247:7 September 3865:20 February 3582:24 February 3546:24 February 3463:24 February 3363:24 February 3344:(1): 1–30. 3239:24 February 3177:24 February 2607:2445/103323 2350:: 187–210. 2285:: 263–287. 2219:23 February 2157:24 February 2117:10023/17427 1963:(1): 2405. 1868:10871/39976 1655:22 February 1569:23 February 1404:coevolution 1391:correlation 1082:loudspeaker 894:Rat culture 805:cetaceans, 803:matrilineal 775:sperm whale 683:Chimpanzees 671:groomers." 643:Masao Kawai 332:chimpanzees 7620:Categories 7357:Structures 7352:Stereotypy 6143:Subculture 5921:Bioculture 4973:1051994025 4482:2019-03-23 3979:(Thesis). 3899:: e12085. 3669:(5): 1–5. 3428:: 95–100. 2950:BioScience 2306:Philosophy 1851:(1): 1–9. 1517:References 1174:Otis tarda 900:black rats 823:nucleotide 787:cetologist 698:tool usage 689:chimpanzee 655:chimpanzee 240:psychology 208:Background 66:newspapers 7586:Behaviour 7529:Societies 7367:Honeycomb 6971:Culturgen 6739:Mormonism 6697:Religions 6370:Cultureme 6288:Destroyed 5914:Subfields 5761:232484226 5745:0036-8075 5685:0002-4619 5586:0003-3472 5572:: 25–31. 5536:0003-3472 5483:0027-8424 5416:1476-4687 5369:0394-9370 5316:2054-5703 5269:0004-8038 5210:0962-8452 5168:206519140 5095:1744-9561 5053:0036-8075 5009:0908-8857 4943:236179560 4844:1932-6203 4777:0028-0836 4714:0376-6357 4679:0028-0836 4400:0003-3472 3625:0960-9822 3522:237280652 3506:1610-7365 3442:0376-6357 3400:145786685 2914:0033-5770 2326:170445206 2247:CiteSeerX 2050:2045-2322 1985:2041-1723 1885:190642523 1877:2055-1045 1774:0011-3204 1711:228894898 1703:2333-9683 1614:196647672 1606:1543-592X 1564:232484226 1166:Migration 1159:cockatoos 1046:Bird song 1028:starlings 1017:songbirds 991:starlings 922:black rat 852:migration 828:sympatric 756:porpoises 561:Imitation 544:Imitation 529:pinnipeds 481:imitation 434:cetaceans 420:pathogens 404:socialize 316:Whiten's 309:imitation 272:languages 189:and some 152:Aristotle 137:behaviors 7631:Ethology 7605:Category 7550:Journals 7377:Instinct 7327:Learning 7322:Instinct 7297:Ethogram 7280:Grooming 7203:Branches 7196:Ethology 7122:Category 6704:Buddhism 6454:Politics 5863:Sciences 5753:33795431 5633:53196374 5594:53188789 5544:53151810 5501:28739943 5434:25470065 5334:27493780 5228:25673683 5160:19498167 5113:27247439 5017:73664186 4935:34437121 4881:ABC News 4862:26488493 4814:PLOS ONE 4795:25470065 4730:26052031 4722:24897247 4572:30333629 4469:15547675 4461:14208581 4408:53198878 4300:30190405 4171:27701411 4120:21321750 4112:20582623 4073:15947077 3925:34532160 3746:16113685 3703:20502702 3663:PLOS ONE 3633:17555968 3514:34427809 3484:Primates 3458:29632261 3450:24780845 3358:12524391 3338:Primates 3234:24052064 3226:11530544 3172:18657877 3122:Archived 3118:26305267 3077:27635227 3014:Archived 2981:Archived 2977:85669317 2930:40567375 2876:53159135 2821:44375991 2773:21208827 2765:21215677 2664:34034521 2636:(1951). 2551:(2002). 2490:(2012). 2466:10385119 2214:32053428 2206:16806574 2134:72336117 2126:30808816 2068:26548756 2003:31160560 1938:33878919 1790:35152891 1556:33795431 1542:(6537). 1432:See also 1380:genetics 1209:(NBDA). 1203:analyzed 1145:ABC News 1077:sparrows 1056:dialects 1032:cowbirds 1024:sparrows 874:Tursiops 864:Dolphins 832:ecotypes 779:dialects 752:dolphins 694:grooming 639:kaluchua 624:macaques 582:Language 577:Language 517:teaching 507:Teaching 493:meerkats 489:language 485:teaching 430:primates 288:learning 183:dolphins 96:May 2023 7307:Feeding 7153:Changes 7134:Commons 6777:Related 6768:Sikhism 6763:Judaism 6194:Aspects 5856:Outline 5842:Culture 5725:Science 5492:5544276 5461:Bibcode 5425:4344839 5396:Bibcode 5325:4968472 5296:Bibcode 5247:The Auk 5219:4345451 5140:Bibcode 5132:Science 5104:4938043 5033:Science 4915:Bibcode 4907:Science 4853:4619304 4822:Bibcode 4786:4344839 4757:Bibcode 4657:Bibcode 4631:9671785 4599:Bibcode 4563:6219627 4542:Bibcode 4441:Bibcode 4433:Science 4278:Bibcode 4269:Science 4162:5049772 4064:1157020 4041:Bibcode 3916:8404572 3827:9831562 3807:Bibcode 3799:Science 3754:4408848 3726:Bibcode 3694:2873264 3671:Bibcode 3641:1236151 3294:May 18, 3128:May 18, 3068:5007745 2987:May 18, 2961:: 489. 2922:1635977 2694:May 18, 2655:8150029 2474:4385871 2446:Bibcode 2096:Bibcode 2088:Science 2059:4637828 2030:Bibcode 1994:6546714 1965:Bibcode 1929:8059593 1536:Science 1490:Culture 1422:guppies 1376:ecology 1103:Paridae 1060:sparrow 920:In the 744:Cetacea 709:ecology 635:Koshima 426:Culture 417:endemic 409:climate 362:Feldman 204:(CMS). 148:culture 133:animals 80:scholar 6153:Fandom 5759: 5751: 5743: 5710: 5700: 5683: 5665:Alauda 5631: 5592: 5584: 5542: 5534: 5499: 5489: 5481: 5432: 5422: 5414: 5388:Nature 5367: 5332: 5322: 5314: 5267: 5226: 5216: 5208: 5166: 5158: 5111: 5101: 5093: 5051: 5015: 5007: 4971: 4941: 4933: 4860: 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