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experimentally demonstrated in guppies. After a group of guppies was trained to swim a more energetically costly route to food, naive guppies were added to the group. These new members were more likely to follow the larger group even if a shorter, more effective route was provided; however, if the naive guppies were not introduced to the trained individuals, they preferentially used the more efficient route to the food source. Norway rats have been shown to abandon previously individually learned habits due to the actions of conspecifics. Rats modified established aversions to certain foods if they observed conspecifics eating those same foods. If this previous aversion was formed for adaptive reasons (i.e. the food was nutrient-poor), reversion to consumption of this food source could reduce the fitness of the individual. More theoretically, social learning can become maladaptive after a certain point in animal populations. If there are more social learners than asocial learners in a particular group, the information transferred between individuals is less likely to be reliable. This could result in maladaptive information transfer to social learners, decreasing the fitness for social learners in comparison to that of asocial learners. Therefore, social learning is only an adaptive strategy if the number of individual learners is equal to or greater than the number of social learners.
115:). One pilot study determined that black rats living in the forests of Palestine preferentially fed on pine cones instead of other fresh fruits and vegetation nearby. These rats also methodically stripped pine cones rather than gnawing at them randomly. To determine how these food preferences developed, researchers provided naive adult black rats with fresh pine cones in captivity and observed their behavior. After three months of experimentation, they found that none of the rats had successfully opened the pine cones and had instead haphazardly and inefficiently attempted to feed on the cones. In further experiments, the rats were allowed to observe experienced individuals opening the pine cones but were still unable to pick up the skill of pinecone stripping. Eventually, the researchers determined that naive adult rats could learn to strip pine cones efficiently if presented with an already partially stripped cone. This is consistent with opportunity providing because experienced individuals inadvertently provide naive rats with partially stripped cones that facilitate their learning without altering the experienced rats' behavior.
897:) have been shown to teach pups essential prey-handling skills. Meerkats often consume dangerous prey (such as venomous scorpions) that inexperienced pups appear unable to safely subdue and consume without assistance from older individuals. Adults only display teaching behavior in response to pup begging calls, and adults modify their specific teaching behaviors based upon the age of the pup begging (providing more assistance to younger, presumably less experienced pups). In some instances, providing prey to pups appears costly to adults. Pups are initially unable to find and consume any of their own prey and more rapidly gain predatory abilities through learning experiences from a “teacher”, suggesting that adult teaching facilitates both the speed and efficiency of skill acquisition.
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achieve a goal such as reaching otherwise inaccessible food after observing another to do such a task but using the tool in a different way than the model. The term 'emulation' encompasses a scope of distinct social learning processes, including object movement re-enactment, end-state emulation, and affordance learning. Object movement re-enactment is the extraction and copying of certain steps of a process of a model moving an object. End-state emulation is the copying of the results of a model's actions using the observer's own unique means. Affordance learning deals with the idea that an observer can gain information about physical properties of the environment and objects within it and how those may interact and then use such information to complete a task.
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actively demonstrates a behavior pattern or draws attention to a location with the specific function of transmitting information to the pupil. An individual must meet three criteria to qualify as a “teacher”: it modifies its behavior only in the presence of a naive observer, it incurs some cost (or at least, no benefit) to itself in doing so, and the naive observer acquires knowledge or skill more rapidly or efficiently than it might otherwise. All of these criteria are rarely met in animals, and only recently have convincing examples been adequately described. Most identified examples have still not been conclusively proven to meet all criteria and primarily serve to suggest that teaching may occur while acknowledging that further research is needed.
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in this task was established in the demonstrator pigeon, other learner pigeons were placed in a video-monitored observation chamber. After every second observed trial, these learner pigeons were then individually placed in the demonstrator pigeon's box and presented the same test. The learner pigeons displayed competent performance on the task, and thus it was concluded that the learner pigeons had formed a response-outcome association while observing. However, the researchers noted that an alternative interpretation of these results could be that the learner pigeons had instead acquired outcome-response associations that guided their behavior and that further testing was needed to establish if this was a valid alternative.
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observer learns the relationship between a stimulus and a result but does not directly copy the behavior of the experienced individual. In imitation, the observer directly copies the behavior of the animal in order to complete a novel task. In emulation, the observer learns a goal from the observed animal’s behavior and seeks to achieve the same results while not following all of the same steps. All of these mechanisms are possible through inadvertent social learning, without active facilitation from the experienced individual. When an individual more actively influences another's behavior through any one of these mechanisms, the individual becomes a teacher.
866:” to their young. Stranding in these circumstances is a behavior in which a whale temporarily beaches itself to reach prey on land near to the shoreline. Adult whales have been observed helping juveniles with many aspects of this behavior by pushing them up and down the beach, guiding them towards prey, and intervening when the juveniles find themselves in danger. Adults only engage in this behavior with apparently naive juveniles. They are more successful when hunting alone than when teaching juveniles, but case studies suggest that juveniles taught the stranding technique by adults are able to master it more than a year earlier than their non-taught peers.
1038:) also utilizes social learning behaviors to find food sources. Mature rats leave sensory trails to and from food sources that are preferentially followed by naive pups. Animals can also learn what to eat from social learning with conspecifics. Experimentally discerning harmful foods from edible foods can be dangerous for a naive individual; however, inexperienced individuals can avoid this cost through observing older individuals that already have acquired this knowledge. For example, several species of bird will avoid food if they see another individual eat it and become ill. Observation can also educate the observer about how to eat particular foods.
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208:) where young sparrows were exposed to song models with controlled trill rates. Higher trill rates are more difficult to perform and thus are likely more desired in birdsong performance. When exposed to low-performance models, it was found that the learner sparrows sacrificed imitative accuracy for higher performance, while when exposed to high-performance models, imitation was very accurate. The researchers suggest that this study may provide insight into how behaviors learned through imitation can still be selected for due to level of performance.
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the individual to obtain. Therefore, individuals that are able to capitalize on other individuals' self-acquired information may experience a fitness benefit. However, because social learning relies on the actions of others rather than direct contact, it can be unreliable. This is especially true in variable environments, where appropriate behaviors may change frequently. Consequently, social learning is most beneficial in stable environments, in which predators, food, and other stimuli are not likely to change rapidly.
160:), young monkeys that observed their parents fearfully responding to model snakes also developed a fear of snakes without direct contact. After three months, the observer rhesus macaques still showed strong fearful reactions toward snakes, suggesting that they had formed a strong connection from just observing their parents’ behavior. Another example of this is how blackbirds learn to identify predators; they observe other birds mobbing unfamiliar objects they haven't seen before.
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95:) demonstrated how local enhancement influenced foraging behavior. Untrained adult female guppies (observers) were given five days of experience swimming with demonstrator fish trained to take one of two equivalent routes to food. When these observers were later isolated, they preferentially used the route their demonstrators had used. These results indicate that guppies learn about their local environments through the social learning mechanism of local enhancement.
1089:. Even when the originally trained “founder” guppies were removed from the group and only initially naive individuals (trained by the founders) were present, the tradition persisted and new guppies learned the costly path. Socially learning the more costly route also resulted in slower learning of the more efficient route when it was subsequently presented, suggesting that even maladaptive strategies can be socially learned and incorporated into local traditions.
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observes another (human or monkey) performing a similar action. Ferrari, Bonini, and
Fogassi worked to explain how the mirror neuron framework could account for imitation of a multiple phenomena with ranging complexities and cognitive demands; they proposed a ‘direct mirror pathway’ for earlier, more automatic imitation and an ‘indirect mirror pathway’ that seems important for more complex and efficient imitative behaviors.
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taps on the leader's body, showing that the leader (teacher) modifies its behavior in the presence of the naive follower. Tandem running appears to impose a significant cost on the leader, slowing its speed to ¼ of what it would be if running alone. But the benefit is clear: evidence suggests that followers find food much more quickly through tandem running than from searching alone.
215:). When given to foster parents, it was shown that fledgling crossbills will imitate the particular variations in their foster parents' calls. It was thus hypothesized that such idiosyncrasies in call could aid in creating familial cohesion and that when such call variants are passed down generations, those variants are direct signals of the crossbill's genetic and familial history.
981:). Blackbird groups were more likely to mob an object if a member of the group had previously been conditioned to recognize it as a predator. This behavior has also been recorded in guppies, where naive guppies from environments with less natural predators significantly improved their anti-predator behavior when placed in a group with guppies from a high predation environment.
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62:. Mechanisms of inadvertent social learning relate primarily to psychological processes in the observer, whereas teaching processes relate specifically to activities of the demonstrator. Studying the mechanisms of information transmission allows researchers to better understand how animals make decisions by observing others' behaviors and obtaining information.
188:'over-imitation.' Meanwhile chimpanzee responses depended more on context such as causal relevance of actions. Fidelity of behavior copying and transfer reportedly plays a role in cultural transmission, so understanding copying fidelity in non-human animals may be important for understanding their capacity for cultural transmission and cumulative culture.
251:) that explored the importance of neural motor circuitry on birdsong learning. If the premotor nucleus was disrupted while a juvenile finch was learning a song from an older finch, the song was not copied. Images of a finch undergoing various neural manipulations showed that premotor circuits aid in encoding information about songs.
140:) found that individuals that had previously observed a human opening a box preferentially spent more time investigating the box and attempting to open it via trial and error. These geese were also more successful at eventually opening the box in comparison to control geese that had not previously observed a human opening the box.
1021:) were subsequently more likely to mob cuckoos but not harmless controls. These results indicate that social learning provides a mechanism by which hosts can rapidly increase their nest defense against brood parasites, enabling the hosts to track fine-scale spatiotemporal variation in the local risk of brood parasitism.
239:) and compared to adult humans. The macaques showed weak evidence of imitative learning compared to the adult humans. It was hypothesized that because the macaques were adults, they were less likely to imitate than juvenile monkeys, as accurate imitation may be an adaptation that is more useful to juveniles.
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Social learning is a beneficial means to gather information if asocial learning is particularly costly and would increase risk of predation, parasitism, or of expending unnecessary energy. In these types of scenarios, social learners may observe the behaviors of an experienced individual. This allows
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Behavioral mechanisms have also been studied. Cecilia Heyes at the
University of Oxford argues that the mechanisms underlying social learning in both humans and nonhumans are analogous to those of non-social learning. Observational learning, then, only becomes social when perceptual, attentional, and
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Mirror neurons have been implicated as the link in primate brains between visual observation and motor representation. These special neurons, originally discovered in area F5 of the ventral premotor cortex of monkeys, are activated when an individual performs a certain action and when that individual
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Though the exact definition of imitation is a topic of debate within scientific literature, broadly, in imitation a learner observes a unique action performed by the demonstrator and learns to reproduce the behavior with detectable behavior matching. (This differs from "copying" in which the learner
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If copying errors are common, or if each observer adds individually learned modifications to a new behavior pattern, stable traditions are unlikely to develop and persist over time. However, even when no longer adaptive, traditions can be passed down if individuals learn primarily through observing
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Social learning does not necessarily mean that the transmitted behavior is the most efficient response to a stimulus. If a socially learned behavior expends unnecessary energy, and there is a more efficient strategy that is not being utilized, employing social learning is maladaptive. This has been
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Pigeons are able to learn behaviors that lead to the delivery of a reward by watching a demonstrator pigeon. A demonstrator pigeon was trained to peck a panel in response to one stimulus (e.g. a red light) and hop on the panel in response to a second stimulus (e.g. a green light). After proficiency
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MacDougall-Shackleton summarized research that suggests developmental stressors affect bird song learning, and that such discrepancies in call can be identified and selected against in some species of birds. He suggests that while many studies have shown that several species of songbirds prefer the
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Social learning is fundamentally different from individual learning, or asocial learning, which involves learning the appropriate responses to an environment through experience and trial and error. Though asocial learning may result in the acquisition of reliable information, it is often costly for
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provides evidence that teaching can occur even without a large brain with complex cognitive abilities. This behavior is shown by an ant who has located a food source in order to guide a naive ant to the desired location. The leading ant only continues the tandem run if the following ant frequently
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Many of the mechanisms involved in inadvertent social learning are also employed during teaching; the distinction is drawn based upon the role of the demonstrator. From the perspective of the pupil, teaching would be identical to its inadvertent social learning equivalent, but in teaching, a tutor
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Adolescence and adulthood: When rats forage on their own, their food choices are influenced by social interactions that may take place far away from foraging sites. They smell foods on the fur, whiskers and especially the breath of other rats and strongly prefer the foods those rats had previously
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Social learning also provides individuals with information about food sources in an environment. This can include information on where to find food, what to eat, and how to eat it. There are several examples in the animal kingdom in which animals utilize social learning to find food. For example,
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Much of the research that has been conducted on imitation and emulation in animals has centered around primates due to their advanced cognitive capacities and evolutionary proximity to humans. Examples of studies that have explored these capacities and tendencies in primates are listed in a table
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Emulation is similar to imitation in that after observing a demonstrator interacting with objects in its environment, an observer is more likely to act to bring about a similar effect on those objects, but not necessarily through the same method. For example, emulation may include using a tool to
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Observational conditioning is a phenomenon similar to stimulus enhancement. In observational conditioning, the behavior of the demonstrator exposes the learner to a new relationship between stimuli that it had not previously known, and causes the learner to form an association between them. In an
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Social learning occurs when one individual influences the learning of another through various processes. In local enhancement and opportunity providing, the attention of an individual is drawn to a specific location or situation. In stimulus enhancement, emulation, observational conditioning, the
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through direct experience. Thus, predator learning is very costly and increases the predation risk for each individual. In group learning scenarios, a few members can experience the danger of predation and transmit this acquired predator recognition throughout the group. Consequently, in future
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Beyond the studies listed, in a naturalistic environment, imitative learning is seen in many animal species. Many species of songbirds learn their songs through imitation, and it has been hypothesized that chimpanzees' understanding of intentionality of action in other members of a social group
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by chimpanzees, in which a mother chimpanzee may contribute to the development of her offspring's nut cracking technique by leaving "hammer", either hard wood or rocks, and nuts in the nest. The infant is thus given the chance to use the hammer in the “proper” context. The mother chimpanzee may
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can occur. These learned behavior complexes shared by individuals appear in the population generation after generation and persist in the behavioral repertoire of individual organisms even following removal from the immediate learning situation. One of the most commonly recognized examples of
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One key distinction between imitation and emulation is copying fidelity. High-fidelity is associated with imitation. In studies comparing behaviors of chimpanzees and human children, the human children were typically shown to perform high-fidelity imitation, what may even be considered
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Opportunity providing is a social learning mechanism in which the experienced individual puts the observer in a situation that facilitates the acquisition of knowledge or a new skill. A well known example of unintentional opportunity providing is the transmission of feeding behavior in
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song dialect of their local area, current data is lacking in explaining why this is so. It has been argued that genetic factors may play into this preference as well as social learning. In this study, three separate groups of laboratory-raised house finches (
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tradition in animals is found in songbirds, in which the same song pattern is transmitted from generation to generation by vocal imitation. Even "alien" syllable types not produced by their biological parents can be learned by
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animals, much of the research has centered around the presence or absence of these abilities in primates. The table below provides an overview of the field of research related to possible imitation and emulation in primates.
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While asocial learners' fitness remains relatively unchanged regardless of frequency, social learners' fitness is higher when they are relatively rare in the population. Social learners' fitness decreases as their frequency
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In stimulus enhancement, a demonstrator exposes an observer to a particular stimulus, leading to the observer learning the relationship between a stimulus and its result. A study investigating stimulus enhancement in
223:) were raised hearing the local song dialect, a foreign song dialect, or no song. When adults, all finches showed sexual preference for the local dialect, suggesting an inherited component to song preference.
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Before birth: In utero, fetal rats detect odor-bearing particles that come from their mother's diet and cross the placental barrier. Shortly after birth, newborn rats respond positively to these foods.
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Weaning: When young rats are weaning and eating solid foods for the first time, they use adult rats as guides. They forage where the adults are foraging or where adults have previously scent-marked.
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Galef, Bennett G. (1986). "Social interaction modifies learned aversions, sodium appetite, and both palatability and handling-time induced dietary preference in rats (Rattus norvegicus)".
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Hernandez, Alexandra M.; MacDougall-Shackleton, Scott A. (2004). "Effects of early song experience on song preferences and song control and auditory brain regions in female house finches (
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Rats use social learning in a wide range of situations, but perhaps especially so in acquiring food preferences. Learning about suitable foods can be divided into four life stages.
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Kelley, Jennifer L.; Evans, Jonathan P.; Ramnarine, Indar W.; Magurran, Anne E. (2003). "Back to school: Can antipredator behavior in guppies be enhanced through social learning?".
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The mental life of the monkeys by Edward L. Thorndike: This
Monograph is also Vol. IX, No. 1, of Columbia University Contributions to Philosophy, Psychology and Education
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refers to learning that is facilitated by observation of, or interaction with, another animal or its products. Social learning has been observed in a variety of animal
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Ferrari, Maud C. O.; Messier, François; Chivers, Douglas P. (June 2007). "First
Documentation of Cultural Transmission of Predator Recognition by Larval Amphibians".
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experienced individuals rather than through asocial learning techniques. This was observed in guppies in the laboratory, in the group foraging example explained
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Learning about brood parasites through direct experience can also be costly and error-prone. Birds can actively defend themselves against brood parasitism from
873:, adult hens encourage safe food choices by responding with increased pecking and scratching at palatable food when chicks consume apparently unpalatable food.
947:, adults emit "food-offering" vocalizations only in the presence of infants that appear to indicate the presence of food or discovery of a hidden prey item.
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to guide naive workers to an identified food source. This dance provides information about the direction, distance and quality of the food source. In
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Einzelausgabe aus der
Anthropoidenstaton auf Teneriffa. Ziele und Aufgaben der Station sowie rste Beobachtungen an den auf ihr gehaltenen Schimpansen
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Fritz, Johannes; Bisenberger, Agnes; Kotrschal, Kurt (2000). "Stimulus enhancement in greylag geese: Socially mediated learning of an operant task".
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Terkel, Joseph (1996). "Cultural
Transmission of Feeding Behavior in the Black Rat (Rattus rattus)". In Heyes, Cecilia M.; Galef, Bennett G. (eds.).
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Custance, Deborah; Prato-Previde, Emanuela; Spiezio, Caterina; Rigamonti, Marco M.; Poli, Marco (2006). "Social learning in pig-tailed macaques (
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Kendal, Rachel L.; Coolen, Isabelle; Van Bergen, Yfke; Laland, Kevin N. (2005). "Trade-Offs in the
Adaptive Use of Social and Asocial Learning".
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encounters, the entire group can recognize the threat of predation and respond accordingly. This social learning method has been shown in group
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Whiten, Andrew; van de Waal, Erica (November 2017). "Social learning, culture and the 'socio-cultural brain' of human and non-human primates".
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During nursing: Nursing rats receive information about their mother's diet through her milk. They prefer the foods she ate during lactation.
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Mechanisms that support imitative learning have been studied on the neurological level. Roberts et al. performed research on zebra finches (
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Kameda, Tatsuya; Nakanishi, Daisuke (2002). "Cost–benefit analysis of social/cultural learning in a nonstationary uncertain environment".
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Galef, Bennett G.; Giraldeau, Luc-Alain (2001). "Social influences on foraging in vertebrates: Causal mechanisms and adaptive functions".
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As a sexually selected trait, variation in learning of songbird calls is often studied. Lahti et al. performed a study on swamp sparrows (
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reproduces that same action but this is performed with a different part of the body e.g. the left paw is used instead of the right paw.)
1958:"Copying results and copying actions in the process of social learning: chimpanzees (Pan troglodytes) and human children (Homo sapiens)"
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Reader, Simon; Kendal, Jeremy; Laland, Kevin (2003). "Social learning of foraging sites and escape routes in wild
Trinidadian guppies".
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Trigger light or sound by using head, foot, or sitting (greater imitation when observed was hands free and making unconstrained action)
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The lead worker (on the left) has returned to the nest and is leading the remaining workers back to the food source via tandem running.
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Nicol, Christine J.; Pope, Stuart J. (1996). "The maternal feeding display of domestic hens is sensitive to perceived chick error".
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Hirata, Satoshi; Celli, Maura L. (2003). "Role of mothers in the acquisition of tool-use behaviours by captive infant chimpanzees".
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Mineka, Susan; Davidson, Mark; Cook, Michael; Keir, Richard (1984). "Observational conditioning of snake fear in rhesus monkeys".
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Sewall explored the variation in learned bird songs in relation to social and genetic intermixing of families of red crossbills (
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The performance of opening an artificial fruit after watching a demonstrator was tested in groups of adult pig-tailed macaques (
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Lahti, David C.; Moseley, Dana L.; Podos, Jeffrey (2011). "A Tradeoff
Between Performance and Accuracy in Bird Song Learning".
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Other species appear to similarly teach their young through the provisioning of weakened or otherwise subdued prey. In both
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Alem, Sylvain; Perry, Clint J.; Zhu, Xingfu; Loukola, Olli J.; Ingraham, Thomas; Søvik, Eirik; Chittka, Lars (2016-10-04).
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Call, Josep (2009). "Contrasting the Social
Cognition of Humans and Nonhuman Apes: The Shared Intentionality Hypothesis".
908:, adults catch live prey animals and transport them back to cubs, allowing the cubs to learn and practice hunting skills.
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Hoppitt, W; Brown, G; Kendal, R; Rendell, L; Thornton, A; Webster, M; Laland, K (2008). "Lessons from animal teaching".
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In local enhancement, a demonstrator attracts an observer's attention to a particular location. One seminal study with
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1046:) have been shown to more successfully open hickory nuts after watching more experienced squirrels open the nuts.
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4302:"Processes of social learning in the tool use of chimpanzees (Pan troglodytes) and human children (Homo sapiens)"
2016:"Causal knowledge and imitation/emulation switching in chimpanzees (Pan troglodytes) and children (Homo sapiens)"
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Curio, E.; Ernst, U.; Vieth, W. (1978). "Cultural Transmission of Enemy Recognition: One Function of Mobbing".
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When many individuals residing within the same area employ social learning, local traditions can be formed and
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Carpenter, Clarence Ray (1937). "An Observational Study of Two Captive Mountain Gorillas (Gorilla Beringei)".
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via a variety of behaviors, including mobbing, and these too can be socially learned. For example, a study on
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Accessing food reward from puzzle box (in clear condition when causal vs irrelevant steps could be observed)
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Door opening, using keys in locks, regulating water supply with lever, scrubbing floor, sweeping with broom
4935:"Rogers' Paradox Recast and Resolved: Population Structure and the Evolution of Social Learning Strategies"
1254:"Associative Mechanisms Allow for Social Learning and Cultural Transmission of String Pulling in an Insect"
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Aisner, R.; Terkel, J. (1992). "Ontogeny of pine cone opening behaviour in the black rat, Rattus rattus".
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Local enhancement has also been observed to transmit foraging information in among birds, rats, and pigs.
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4122:"Social influences on the acquisition of tool-using behaviors in tufted capuchin monkeys (Cebus apella)"
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Roberts, Todd F.; Gobes, Sharon M. H.; Murugan, Malavika; Ölveczky, Bence P.; Mooney, Richard (2012).
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MacDougall-Shackleton, Scott A. (2009). "The importance of development: What songbirds can teach us".
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My friend Toto: The adventures of a chimpanzee and the story of his journey from the Congo to London
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3863:"Spontaneous tool use and sensorimotor intelligence in Cebus compared with other monkeys and apes"
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2133:"Early learning of discrete call variants in red crossbills: implications for reliable signaling"
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the observer to gain information about the environment without individually being put at risk.
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When social learning is actively facilitated by an experienced individual, it is classified as
2674:"Orang-utan tool-use at Tanjung Puting Reserve, Central Indonesian Borneo (Kalimantan Tengah)"
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2445:"From monkey mirror neurons to primate behaviours: possible 'direct' and 'indirect' pathways"
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Other evidence of teaching through local enhancement can be seen in a variety of species. In
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2818:"Identification : A process of enculturation in the subhuman society ofMacaca fuscata"
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Red squirrels are more successful at opening nuts after watching an experienced individual.
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Whiten, Andrew; McGuigan, Nicola; Marshall-Pescini, Sarah; Hopper, Lydia M. (2009-08-27).
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Mundinger, Paul C. (1980). "Animal cultures and a general theory of cultural evolution".
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1898:"Emulation, imitation, over-imitation and the scope of culture for child and chimpanzee"
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motivational factors are focused on other organisms by genetic or developmental forces.
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Galef, Bennett G. (1982). "Studies of social learning in norway rats: A brief review".
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Adams-Curtis, L.E. (1987). "Social context of manipulative behaviour in Cebus apella".
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4380:
4286:
4200:
4106:
3958:
3844:
3624:
Buttelmann, David; Carpenter, Malinda; Call, Josep; Tomasello, Michael (July 2007).
3610:
3536:
3415:
2990:
2849:
2390:"Neurophysiological mechanisms underlying the understanding and imitation of action"
1997:
1882:
1793:
1682:
1647:
1446:
1238:
526:
Solving stick and tunnel problems, solving stick and string problems, throwing ball
5037:
4995:
4987:
4946:
4907:
4878:
4834:
4795:
4758:
4711:
4664:
4641:
4619:
4561:
4518:
4480:
4429:
4360:
4313:
4266:
4227:
4180:
4133:
4086:
4075:"Tool use in capuchin monkeys: Distinguishing between performing and understanding"
4028:
3989:
3938:
3874:
3816:
3750:
3684:
3637:
3590:
3516:
3469:
3395:
3333:
3286:
3110:
3048:
3017:
2970:
2923:
2880:
2876:
2829:
2771:
2744:
2724:
2685:
2654:
2615:
2519:
2472:
2456:
2429:
2401:
2360:
2352:
2313:
2270:
2228:
2189:
2152:
2144:
2113:
2078:
2055:
2027:
1969:
1925:
1909:
1862:
1816:
1773:
1730:
1670:
1635:
1600:
1560:
1523:
1450:
1442:
1407:
1370:
1354:
1320:
1283:
1265:
1218:
1178:
1030:
birds are more likely to forage in areas where they already see birds feeding. The
974:
4215:
1750:
4595:
4016:
3909:
3777:
3430:
3218:
3156:
1866:
1480:
1270:
153:
4911:
4317:
4137:
4032:
3367:
van Lawick-Goodall, Jane (1973). "Cultural elements in a chimpanzee community".
2317:
1347:
Philosophical Transactions of the Royal Society of London B: Biological Sciences
118:
Opportunity providing has also been found to be important in the acquisition of
3052:
2274:
1820:
1604:
1338:
1098:
944:
924:
657:
Putting things in mouth, bell ringing, object examination, bucket interactions
4763:
4746:
4484:
3878:
3578:
3550:
Gardner, R. Allen; Gardner, Beatrix T.; Cantfort, Thomas E. Van (1989-01-01).
3504:
3383:
3189:
The Ape and the Child: A Study of Environmental Influence Upon Early Behaviour
3098:
3006:"Development of tool-use to obtain food in a captive group of Macaca tonkeana"
2817:
2341:"Motor circuits are required to encode a sensory model for imitative learning"
2148:
2031:
1973:
1734:
5054:
4883:
4866:
4395:
4372:
4325:
4278:
4239:
4192:
4145:
4098:
4040:
4001:
3950:
3886:
3828:
3696:
3649:
3602:
3579:"A longitudinal investigation of gestural communication in young chimpanzees"
3528:
3481:
3407:
3345:
3298:
3251:
3122:
3060:
2982:
2935:
2888:
2841:
2736:
2627:
2580:
2468:
2413:
2039:
1981:
1921:
1464:
1419:
1366:
1279:
1039:
1014:
855:
808:
303:
133:
4715:
4565:
2928:
10.1002/1097-4679(195904)15:2<155::AID-JCLP2270150211>3.0.CO;2-P
2388:
Rizzolatti, Giacomo; Fogassi, Leonardo; Gallese, Vittorio (September 2001).
192:
within the ‘Research on Imitation and Emulation in Primates’ section below.
5009:
4991:
4960:
4848:
4772:
4723:
4668:
4633:
4573:
4522:
3836:
3820:
3762:
3704:
3673:"New evidence on imitation in an enculturated chimpanzee (Pan troglodytes)"
3657:
3353:
3306:
2943:
2896:
2531:
2486:
2460:
2421:
2374:
2325:
2282:
2240:
2201:
2166:
2090:
2047:
1989:
1939:
1913:
1874:
1785:
1777:
1742:
1639:
1612:
1472:
1384:
1358:
1297:
1230:
1222:
940:
123:
provide this opportunity unknowing or actively depending on the situation.
4919:
4807:
4799:
4449:
4333:
4153:
3068:
3035:
Cook, Michael; Mineka, Susan; Wolkenstein, Bonnie; Laitsch, Karen (1985).
2704:
2643:"Imitative Learning of a Flaked Stone Technology-The Case of an Orangutan"
2389:
1828:
1500:
Proceedings of the 32nd Annual Conference of the Cognitive Science Society
1311:
Brown, Culum; Kevin Laland (2003). "Social Learning in Fishes: A review".
766:
436:
4348:
4254:
4168:
4074:
3926:
2958:
4839:
4822:
4441:
4420:
Caro, TM; Hauser, MD (1992). "Is there teaching in nonhuman animals?".
4364:
4270:
4184:
4090:
3973:
3942:
3594:
3520:
3399:
3384:"Tool-using and -making behavior in wild chimpanzees at Bossou, Guinea"
3259:
3235:
2974:
2833:
2783:
2759:
1455:
881:
770:
668:
Banging objects together, putting tub in moat, putting cloth on branch
550:
Cracking nuts with stones, techniques for reaching lower tree branches
440:
108:
40:
4980:
Philosophical Transactions of the Royal Society B: Biological Sciences
3993:
3503:
Sumita, Kyoko; Kitahara-Frisch, Jean; Norikoshi, Kohshi (April 1985).
3037:"Observational conditioning of snake fear in unrelated rhesus monkeys"
2588:
2564:
2449:
Philosophical Transactions of the Royal Society B: Biological Sciences
2232:
1902:
Philosophical Transactions of the Royal Society B: Biological Sciences
1167:"Social learning in animals: Empirical studies and theoretical models"
126:
4017:"The manufacture and use of tools by capuchin monkeys (Cebus apella)"
3688:
3337:
3290:
3114:
2728:
2523:
2405:
2295:
2193:
1031:
510:
Brushing hair, applying lipstick, brushing teeth, sharpening pencils
307:
172:
4624:
4599:
4253:
Tomasello, M.; Davis-Dasilva, M.; Camak, L.; Bard, K. (April 1987).
3577:
Tomasello, Michael; Gust, Deborah; Frost, G. Thomas (January 1989).
2775:
2356:
1895:
4433:
3473:
1343:"Social learning in birds and its role in shaping a foraging niche"
1142:
Social Learning: An Introduction to Mechanisms, Methods, and Models
1002:
965:
901:
890:
870:
36:
4471:
Bauer, Gordon B.; Harley, Heidi E. (2001). "The mimetic dolphin".
262:
143:
4692:"Social Transmission of a Host Defense Against Cuckoo Parasitism"
4300:
Nagell, Katherine; Olguin, Raquel S.; Tomasello, Michael (1993).
3458:"Chimpanzees, Tools, and Termites: Another Example From Tanzania"
1956:
Call, Josep; Carpenter, Malinda; Tomasello, Michael (July 2005).
1078:
998:
614:
79:
48:
44:
24:
4976:"Potential disadvantages of using socially acquired information"
4544:
Thornton, A.; McAuliffe, K (2006). "Teaching in Wild Meerkats".
4349:"The social learning of tool use by orangutans (Pongo pygmaeus)"
3623:
869:
Coaching appears to occur in other species as well. In domestic
4252:
2500:
Heyes, Cecilia (2012). "What's social about social learning?".
1013:) showed that individuals that observed their neighbors mob of
3974:"Acquisition of Nut-Cracking Behaviour by 2 Capuchin Monkeys (
3502:
2910:
Miller, Robert E.; Murphy, John V.; Mirsky, I. Arthur (1959).
2565:"Observations on the Mentality of Chimpanzees and Orang-Utans"
2214:
4867:"Social transmission of maladaptive information in the guppy"
3925:
Antinucci, Francesco; Visalberghi, Elisabetta (August 1986).
2263:
Journal of Experimental Psychology: Animal Behavior Processes
268:
88:
32:
4654:
3034:
2760:"Mental Life of Two Macacus rhesus Monkeys in Captivity. II"
1517:
1086:
3671:
Carrasco, Lara; Posada, Sandra; Colell, Montserrat (2009).
2338:
1184:
10.1641/0006-3568(2005)055[0489:SLIAES]2.0.CO;2
936:
928:
28:
20:
4974:
Giraldeau, L.-A.; Valone, T. J.; Templeton, J. J. (2002).
4823:"Social learning about predators: A review and prospectus"
4073:
Visalberghi, Elisabetta; Trinca, Loredana (October 1989).
3505:"The acquisition of stone-tool use in captive chimpanzees"
3236:"The Intellectual Development of a Home-Raised Chimpanzee"
1049:
4933:
Rendell, Luke; Fogarty, Laurel; Laland, Kevin N. (2010).
4255:"Observational learning of tool-use by young chimpanzees"
3099:"Some observations on the intelligence of the chimpanzee"
2254:
Saggerson, A. L.; George, David N.; Honey, R. C. (2005).
1763:
1590:
984:
905:
4973:
2387:
1024:
462:
Digging with spade, screwing screw, scrubbing, sweeping
2797:
Carner, R (1955). "Botanical Collecting with Monkeys".
470:
Wiping nose with handkerchief, hammering nails, sewing
4120:
Fragaszy, Dorothy M.; Visalberghi, Elisabetta (1989).
4015:
Westergaard, Gregory C.; Fragaszy, Dorothy M. (1987).
3549:
2443:
Ferrari, P. F.; Bonini, L.; Fogassi, L. (2009-08-27).
1955:
1806:
388:
Potato washing, rice throwing, consumption of caramel
4299:
2179:
1397:
4932:
3924:
3860:
3670:
3136:
Sheak, W.H. (1923). "Anthropoid apes I have known".
2442:
2253:
242:
4464:
4119:
4014:
3456:Nishida, Toshisada; Uehara, Shigeo (October 1980).
3326:
Journal of Comparative and Physiological Psychology
3279:
Journal of Comparative and Physiological Psychology
1498:. In Ohlsson, Stellan; Catrambone, Richard (eds.).
1251:
127:
Stimulus enhancement and observational conditioning
3576:
3382:Sugiyama, Yukimaru; Koman, Jeremy (October 1979).
3366:
2909:
1625:
534:Patting head, clapping hands, sticking out tongue
4543:
4072:
3240:Proceedings of the American Philosophical Society
2865:"Observational Learning of an Avoidance Response"
2863:Presley, W. J.; Riopelle, A. J. (December 1959).
2569:Proceedings of the American Philosophical Society
2103:
2009:
2007:
1432:
1310:
992:
478:Using paintbrush, stacking boxes to reach banana
5052:
4594:
3861:Chevalier-Skolnikoff, Suzanne (September 1989).
3429:Waal, Frans de; Waal, Frans B. M. (2000-04-10).
3322:"Picture perception in a home-raised chimpanzee"
2604:"A STUDY OF ORANG-UTAN BEHAVIOUR IN THE 1959-60"
1698:Social Learning In Animals: The Roots of Culture
1496:"Spatial Factors of Social and Asocial Learning"
4744:
4689:
2862:
2547:Narrative of a Journey in the Interior of China
1951:
1949:
1844:
1550:
950:
730:Nut cracking, moving reward in tube with stick
263:Research on Imitation and Emulation in Primates
4864:
3802:
3186:
2014:Horner, Victoria; Whiten, Andrew (July 2005).
2004:
1337:
1208:
380:Botanical collection (assisting experimenter)
267:In considering imitation and emulation in non-
226:
4779:
4738:
4346:
3626:"Enculturated chimpanzees imitate rationally"
3455:
3432:Chimpanzee Politics: Power and Sex Among Apes
3381:
3081:
1139:
1135:
1133:
396:Jumping over barrier to avoid electric shock
147:One adult female and one young rhesus macaque
65:
4393:
4053:
3739:"Observational Learning in Monkeys and Apes"
2013:
1946:
1660:
1513:
1511:
1509:
1165:Galef, Bennett G.; Laland, Kevin N. (2005).
1131:
1129:
1127:
1125:
1123:
1121:
1119:
1117:
1115:
1113:
4470:
3971:
2182:Canadian Journal of Experimental Psychology
1720:
1494:Metz, Alexander; Shultz, Thomas R. (2010).
1164:
1140:Hoppitt, William; Laland, Kevin N. (2013).
959:
822:
4865:Laland, Kevin N.; Williams, Kerry (1998).
4504:
4502:
4419:
3320:Hayes, Keith J.; Hayes, Catherine (1953).
3319:
3273:Hayes, Keith J.; Hayes, Catherine (1952).
3272:
3234:Hayes, Keith J.; Hayes, Catherine (1951).
3233:
1586:
1584:
1582:
1580:
1578:
1576:
1574:
1493:
1204:
1202:
1200:
1198:
1196:
1194:
911:
876:
372:Plug pulling, lever pressing, box opening
163:
5027:
5023:
5021:
5019:
4999:
4950:
4882:
4838:
4762:
4623:
4508:
3907:
3717:
2702:
2601:
2513:
2476:
2364:
2156:
1929:
1506:
1454:
1374:
1287:
1269:
1182:
1110:
4860:
4858:
4690:Davies, N. B.; Welbergen, J. A. (2009).
4347:Call, J.; Tomasello, M. (October 1994).
3428:
3096:
3003:
2959:"Tool use by captive pigtailed macaques"
2815:
2757:
2671:
2549:. Longman, Hurst, Rees, Orme, and Brown.
1840:
1838:
1435:Neuroscience & Biobehavioral Reviews
1053:
983:
915:
880:
199:
167:
142:
101:
78:
4820:
4499:
4169:"Tool use in captive hamadryas baboons"
3927:"Tool use inCebus apella: A case study"
3803:Hamilton, W. J.; Tilson, R. L. (1985).
3275:"Imitation in a home-raised chimpanzee"
3171:
2562:
1571:
1191:
1050:Maladaptive examples of social learning
964:An animal generally learns its natural
502:Spitting, imitating facial expressions
5053:
5016:
3805:"Fishing baboons at desert waterholes"
3775:
3201:
3154:
2796:
2672:Galdikas, Biruté M.F. (January 1982).
2640:
2130:
1695:
1081:raised by foster canaries in the lab.
196:influences their imitative behaviors.
4897:
4855:
4785:
3856:
3854:
3553:Teaching Sign Language to Chimpanzees
3216:
3135:
2558:
2556:
2499:
2304:) on a two-action artificial fruit".
1835:
1025:Learning about foraging opportunities
4745:Welbergen, J.A; Davies, N.B (2009).
4213:
4167:Beck, Benjamin B. (September 1972).
4166:
3931:International Journal of Primatology
3736:
3086:. Berlin: Abh Preuss. pp. 1–20.
3004:Anderson, James R. (November 1985).
2956:
2912:"Non-verbal communication of affect"
2544:
2068:
862:) teach the predatory technique of “
74:
4214:Beck, Benjamin B. (November 1974).
3206:. New Haven: Yale University Press.
2137:Behavioral Ecology and Sociobiology
1067:
13:
3851:
3755:10.1111/j.2044-8295.1963.tb00877.x
2764:The American Journal of Psychology
2659:10.1111/j.1835-9310.1972.tb00451.x
2620:10.1111/j.1748-1090.1962.tb03396.x
2553:
939:, knowledgeable workers perform a
619:Matching moods and play behaviors
590:Play initiation by throwing chips
412:Throwing tool at unreachable food
14:
5077:
4900:Journal of Comparative Psychology
4600:"Teaching in tandem-running ants"
4396:"How do rats choose what to eat?"
4306:Journal of Comparative Psychology
4216:"Baboons, chimpanzees, and tools"
4126:Journal of Comparative Psychology
4021:Journal of Comparative Psychology
3677:Journal of Comparative Psychology
3187:Kellogg, WN; Kellogg, LA (1933).
2869:The Journal of Genetic Psychology
2502:Journal of Comparative Psychology
2306:Journal of Comparative Psychology
1593:Trends in Ecology & Evolution
1522:. Vol. 35. pp. 333–79.
1520:Advances in the Study of Behavior
693:Antinucci and Visalberghi (1986)
494:Brushing hair, opening cupboards
243:Mechanisms for imitative learning
4952:10.1111/j.1558-5646.2009.00817.x
3972:Visalberghi, Elisabetta (1987).
3642:10.1111/j.1467-7687.2007.00630.x
3204:Chimpanzees: a laboratory colony
2118:10.1111/j.1439-0310.2011.01930.x
2083:10.1111/j.1756-8765.2009.01025.x
1412:10.1111/j.1439-0310.2007.01362.x
1341:; Wiebe, Karen L. (2011-04-12).
1325:10.1046/j.1467-2979.2003.00122.x
733:Fragaszy and Visalberghi (1989)
709:Westergaard and Fragaszy (1987)
4967:
4926:
4891:
4814:
4683:
4648:
4588:
4537:
4422:The Quarterly Review of Biology
4413:
4387:
4340:
4293:
4246:
4207:
4160:
4113:
4066:
4056:American Journal of Primatology
4047:
4008:
3965:
3918:
3901:
3809:American Journal of Primatology
3796:
3769:
3730:
3726:: 175–200 – via ProQuest.
3711:
3664:
3617:
3570:
3543:
3496:
3449:
3422:
3375:
3360:
3313:
3266:
3227:
3210:
3195:
3180:
3165:
3148:
3129:
3090:
3075:
3028:
2997:
2957:Beck, Benjamin B. (July 1976).
2950:
2903:
2856:
2809:
2790:
2751:
2696:
2665:
2641:Wright, R. V. S. (2009-02-10).
2634:
2595:
2538:
2493:
2436:
2381:
2332:
2289:
2247:
2208:
2173:
2124:
2097:
2062:
1889:
1800:
1757:
1714:
1689:
1654:
1619:
1544:
1447:10.1016/j.neubiorev.2016.12.018
885:Adult meerkats search for prey.
337:Building nests, feeding habits
4827:Animal Learning & Behavior
3776:Marais, Eugène Nielen (1990).
3737:Hall, K. R. L. (August 1963).
3082:Rothman, M; Teuber, E (1915).
3041:Journal of Abnormal Psychology
2916:Journal of Clinical Psychology
2881:10.1080/00221325.1959.10534265
2816:Imanishi, Kinji (March 1957).
2758:Kinnaman, A. J. (April 1902).
2703:Carpenter, Alfred (May 1887).
1809:Journal of Abnormal Psychology
1487:
1426:
1391:
1331:
1304:
1245:
1158:
1144:. Princeton University Press.
993:Learning about brood parasites
725:Visalberghi and Trinca (1987)
420:Reaching food with metal rods
1:
4473:Behavioral and Brain Sciences
3908:Thorndike, Edward L. (1901).
3867:Behavioral and Brain Sciences
3743:British Journal of Psychology
3022:10.1016/S0047-2484(85)80072-5
2690:10.1016/S0047-2484(82)80028-6
1565:10.1016/S1090-5138(02)00101-0
1528:10.1016/S0065-3454(05)35008-X
1104:
630:Digging in certain locations
574:Cracking walnuts with stones
283:Positive or negative support
5042:10.1016/0162-3095(80)90008-4
4232:10.1016/0047-2484(74)90011-6
3369:Precultural Primate Behavior
2563:Furness, William H. (1916).
1867:10.1126/science.202.4370.899
1700:. Elsevier. pp. 17–47.
1675:10.1016/0003-3472(92)90038-B
1553:Evolution and Human Behavior
1271:10.1371/journal.pbio.1002564
951:Relevance of social learning
671:Chevalier-Skolnikoff (1989)
660:Chevalier-Skolnikoff (1989)
399:Presley and Riopelle (1959)
364:Opening oysters with stones
7:
4912:10.1037/0735-7036.100.4.432
4788:Developmental Psychobiology
4318:10.1037/0735-7036.107.2.174
4138:10.1037/0735-7036.103.2.159
4033:10.1037/0735-7036.101.2.159
2602:Harrisson, Barbara (1962).
2394:Nature Reviews Neuroscience
2318:10.1037/0735-7036.120.3.303
2071:Topics in Cognitive Science
1092:
845:
649:Hamilton and Tilson (1985)
638:Fruit cracking with stones
569:Nishida and Hiraiwa (1982)
497:Kellogg and Kellogg (1933)
329:Lifting lid of sewage tank
227:Visual behavioral imitation
10:
5082:
4598:; Richardson, Tom (2006).
4220:Journal of Human Evolution
3103:Journal of Animal Behavior
3053:10.1037/0021-843X.94.4.591
3010:Journal of Human Evolution
2678:Journal of Human Evolution
2608:International Zoo Yearbook
2275:10.1037/0097-7403.31.3.289
2131:Sewall, Kendra B. (2011).
1821:10.1037/0021-843X.93.4.355
1605:10.1016/j.tree.2008.05.008
816:Call and Tomasello (1994)
553:Sugiyama and Koman (1979)
545:van Lawick-Goodall (1973)
449:Rothman and Teuber (1915)
435:
299:
296:
280:Likely learning mechanism
66:Social learning mechanisms
5030:Ethology and Sociobiology
4764:10.1016/j.cub.2008.12.041
4485:10.1017/S0140525X01243969
3879:10.1017/S0140525X00057678
3155:Köhler, Wolfgang (1925).
2705:"Monkeys opening Oysters"
2149:10.1007/s00265-010-1022-0
2032:10.1007/s10071-004-0239-6
1974:10.1007/s10071-004-0237-8
1735:10.1007/s10071-003-0187-6
803:Horner and Whiten (2005)
765:
762:
759:
752:Reaching pan using stick
678:
675:
626:
601:Buttelmann et al. (2007)
360:
332:Yerkes and Yerkes (1927)
302:
4400:Rat behavior and biology
3172:Kearton, Cherry (1925).
3097:Shepherd, W. T. (1915).
960:Learning about predators
823:At different life stages
813:Reaching food with tool
784:Reaching food with tool
779:Tomasello et al. (1987)
776:Reaching food with tool
741:Reaching food with tool
593:Tomasello et al. (1989)
4821:Griffin, A. S. (2004).
4716:10.1126/science.1172227
4566:10.1126/science.1128727
2221:Journal of Neurobiology
1011:Acrocephalus scirpaceus
912:Using local enhancement
877:Providing opportunities
654:Ateles (Spider Monkey)
609:Carrasco et al. (2009)
537:Hayes and Hayes (1953)
529:Hayes and Hayes (1952)
521:Hayes and Hayes (1951)
164:Imitation and emulation
4992:10.1098/rstb.2002.1065
4884:10.1093/beheco/9.5.493
4669:10.1006/anbe.2003.2076
4523:10.1006/anbe.1996.0221
3821:10.1002/ajp.1350080308
3202:Yerkes, R. M. (1943).
2461:10.1098/rstb.2009.0062
1914:10.1098/rstb.2009.0069
1778:10.1006/anbe.2000.1424
1640:10.1006/anbe.2003.2252
1359:10.1098/rstb.2010.0343
1223:10.1006/anbe.2000.1557
1060:
1007:Eurasian reed warblers
989:
921:
886:
722:Moving reward in tube
566:Ant species selection
176:
148:
84:
4800:10.1002/dev.420150402
4394:Hanson, Anne (2012).
3630:Developmental Science
3217:Hayes, Cathy (1951).
3176:. London: Arrowsmith.
3158:The Mentality of Apes
2545:Abel, Clarke (1818).
1074:cultural transmission
1057:
987:
919:
884:
787:Nagell et al. (1993)
577:Sumita et al. (1985)
518:Imitation on command
407:Miller et al. (1959)
200:Imitation of birdsong
171:
146:
102:Opportunity providing
82:
3462:Current Anthropology
3220:The Ape in Our House
2300:) and adult humans (
2217:Carpodacus mexicanus
945:callitrichid monkeys
717:Adams-Curtis (1987)
345:Flaking stone tools
221:Carpodacus mexicanus
4708:2009Sci...324.1318D
4702:(5932): 1318–1320.
4616:2006Natur.439..153F
4558:2006Sci...313..227T
3982:Folia Primatologica
3779:The Soul of the Ape
3191:. Whittlesey House.
2721:1887Natur..36...53C
2455:(1528): 2311–2323.
2345:Nature Neuroscience
1908:(1528): 2417–2428.
1859:1978Sci...202..899C
1502:. pp. 1685–90.
795:Call et al. (2005)
701:Visalberghi (1987)
585:Fouts et al (1989)
431:Cook et al. (1985)
321:Digging with spade
276:
249:Taeniopygia guttata
206:Melospiza georgiana
93:Poecilia reticulata
4871:Behavioral Ecology
4840:10.3758/BF03196014
4365:10.1007/bf02435516
4271:10.1007/bf02436405
4185:10.1007/bf01730574
4091:10.1007/bf02380877
3943:10.1007/bf02693700
3595:10.1007/BF02381209
3521:10.1007/BF02382016
3400:10.1007/BF02373433
3161:. Harcourt, Brace.
2975:10.1007/bf02382787
2834:10.1007/BF01667196
1313:Fish and Fisheries
1061:
990:
922:
895:Suricata suricatta
887:
706:Probing for syrup
275:
177:
149:
85:
4986:(1427): 1559–66.
3994:10.1159/000156320
3789:978-0-14-012848-2
3782:. Penguin Books.
3563:978-0-88706-965-9
3442:978-0-8018-6336-3
2799:Proc. Roy. Instil
2351:(10): 1454–1459.
2298:Macaca nemestrina
2233:10.1002/neu.10312
1853:(4370): 899–901.
1707:978-0-08-054131-0
1537:978-0-12-004535-8
1353:(1567): 969–977.
1151:978-1-4008-4650-4
1036:Rattus norvegicus
820:
819:
685:Thorndike (1901)
665:Cebus (Capuchin)
622:Carpenter (1937)
606:Action imitation
367:Carpenter (1887)
361:Macaca (Macaque)
237:Macaca nemestrina
213:Loxia curvirostra
75:Local enhancement
5073:
5066:Animal cognition
5046:
5045:
5025:
5014:
5013:
5003:
4971:
4965:
4964:
4954:
4930:
4924:
4923:
4895:
4889:
4888:
4886:
4862:
4853:
4852:
4842:
4818:
4812:
4811:
4783:
4777:
4776:
4766:
4742:
4736:
4735:
4687:
4681:
4680:
4657:Animal Behaviour
4652:
4646:
4645:
4627:
4596:Franks, Nigel R.
4592:
4586:
4585:
4541:
4535:
4534:
4511:Animal Behaviour
4506:
4497:
4496:
4468:
4462:
4461:
4417:
4411:
4410:
4408:
4406:
4391:
4385:
4384:
4344:
4338:
4337:
4297:
4291:
4290:
4250:
4244:
4243:
4211:
4205:
4204:
4164:
4158:
4157:
4117:
4111:
4110:
4070:
4064:
4063:
4051:
4045:
4044:
4012:
4006:
4005:
3988:(3–4): 168–181.
3969:
3963:
3962:
3922:
3916:
3915:
3905:
3899:
3898:
3858:
3849:
3848:
3800:
3794:
3793:
3773:
3767:
3766:
3734:
3728:
3727:
3715:
3709:
3708:
3689:10.1037/a0016275
3668:
3662:
3661:
3621:
3615:
3614:
3574:
3568:
3567:
3547:
3541:
3540:
3500:
3494:
3493:
3453:
3447:
3446:
3426:
3420:
3419:
3379:
3373:
3372:
3364:
3358:
3357:
3338:10.1037/h0053704
3317:
3311:
3310:
3291:10.1037/h0053609
3270:
3264:
3263:
3231:
3225:
3224:
3214:
3208:
3207:
3199:
3193:
3192:
3184:
3178:
3177:
3169:
3163:
3162:
3152:
3146:
3145:
3133:
3127:
3126:
3115:10.1037/h0070497
3094:
3088:
3087:
3079:
3073:
3072:
3032:
3026:
3025:
3001:
2995:
2994:
2954:
2948:
2947:
2907:
2901:
2900:
2860:
2854:
2853:
2813:
2807:
2806:
2794:
2788:
2787:
2755:
2749:
2748:
2729:10.1038/036053d0
2700:
2694:
2693:
2669:
2663:
2662:
2638:
2632:
2631:
2599:
2593:
2592:
2560:
2551:
2550:
2542:
2536:
2535:
2524:10.1037/a0025180
2517:
2497:
2491:
2490:
2480:
2440:
2434:
2433:
2406:10.1038/35090060
2385:
2379:
2378:
2368:
2336:
2330:
2329:
2293:
2287:
2286:
2260:
2251:
2245:
2244:
2212:
2206:
2205:
2194:10.1037/a0015414
2177:
2171:
2170:
2160:
2128:
2122:
2121:
2101:
2095:
2094:
2066:
2060:
2059:
2020:Animal Cognition
2011:
2002:
2001:
1962:Animal Cognition
1953:
1944:
1943:
1933:
1893:
1887:
1886:
1842:
1833:
1832:
1804:
1798:
1797:
1772:(6): 1119–1125.
1766:Animal Behaviour
1761:
1755:
1754:
1723:Animal Cognition
1718:
1712:
1711:
1693:
1687:
1686:
1663:Animal Behaviour
1658:
1652:
1651:
1628:Animal Behaviour
1623:
1617:
1616:
1588:
1569:
1568:
1548:
1542:
1541:
1515:
1504:
1503:
1491:
1485:
1484:
1458:
1430:
1424:
1423:
1395:
1389:
1388:
1378:
1335:
1329:
1328:
1308:
1302:
1301:
1291:
1273:
1264:(10): e1002564.
1249:
1243:
1242:
1211:Animal Behaviour
1206:
1189:
1188:
1186:
1162:
1156:
1155:
1137:
1068:Local traditions
1044:Sciurus vulgaris
975:common blackbird
971:mobbing behavior
542:Termite Fishing
486:Washing clothes
457:Shepherd (1915)
423:Anderson (1985)
391:Imanishi (1957)
375:Kinnaman (1902)
356:Galdikas (1982)
340:Harrison (1960)
277:
274:
175:eating a coconut
152:experiment with
5081:
5080:
5076:
5075:
5074:
5072:
5071:
5070:
5051:
5050:
5049:
5026:
5017:
4972:
4968:
4931:
4927:
4896:
4892:
4863:
4856:
4819:
4815:
4784:
4780:
4751:Current Biology
4743:
4739:
4688:
4684:
4653:
4649:
4625:10.1038/439153a
4593:
4589:
4552:(5784): 227–9.
4542:
4538:
4507:
4500:
4469:
4465:
4418:
4414:
4404:
4402:
4392:
4388:
4353:Human Evolution
4345:
4341:
4298:
4294:
4259:Human Evolution
4251:
4247:
4212:
4208:
4165:
4161:
4118:
4114:
4071:
4067:
4052:
4048:
4013:
4009:
3970:
3966:
3923:
3919:
3906:
3902:
3859:
3852:
3801:
3797:
3790:
3774:
3770:
3735:
3731:
3716:
3712:
3669:
3665:
3622:
3618:
3575:
3571:
3564:
3548:
3544:
3501:
3497:
3454:
3450:
3443:
3427:
3423:
3380:
3376:
3365:
3361:
3318:
3314:
3271:
3267:
3232:
3228:
3215:
3211:
3200:
3196:
3185:
3181:
3170:
3166:
3153:
3149:
3138:Natural History
3134:
3130:
3095:
3091:
3080:
3076:
3033:
3029:
3002:
2998:
2955:
2951:
2908:
2904:
2861:
2857:
2814:
2810:
2795:
2791:
2776:10.2307/1412738
2756:
2752:
2701:
2697:
2670:
2666:
2639:
2635:
2600:
2596:
2561:
2554:
2543:
2539:
2515:10.1.1.401.6757
2498:
2494:
2441:
2437:
2386:
2382:
2357:10.1038/nn.3206
2337:
2333:
2294:
2290:
2258:
2252:
2248:
2213:
2209:
2178:
2174:
2129:
2125:
2102:
2098:
2067:
2063:
2012:
2005:
1954:
1947:
1894:
1890:
1843:
1836:
1805:
1801:
1762:
1758:
1719:
1715:
1708:
1694:
1690:
1659:
1655:
1624:
1620:
1589:
1572:
1549:
1545:
1538:
1516:
1507:
1492:
1488:
1431:
1427:
1396:
1392:
1339:Slagsvold, Tore
1336:
1332:
1309:
1305:
1250:
1246:
1207:
1192:
1163:
1159:
1152:
1138:
1111:
1107:
1095:
1070:
1052:
1027:
1019:Cuculus canorus
995:
962:
953:
914:
879:
848:
825:
714:Puzzle solving
627:Papio (Baboon)
561:De Waal (1982)
489:Kearton (1925)
465:Furness (1916)
428:Fear of snakes
324:Furness (1916)
313:Kiss imitation
265:
245:
229:
202:
166:
154:rhesus macaques
129:
104:
77:
68:
17:Social learning
12:
11:
5:
5079:
5069:
5068:
5063:
5048:
5047:
5036:(3): 183–233.
5015:
4966:
4925:
4890:
4854:
4813:
4778:
4757:(3): 235–240.
4737:
4682:
4647:
4587:
4536:
4498:
4463:
4434:10.1086/417553
4412:
4386:
4359:(4): 297–313.
4339:
4312:(2): 174–186.
4292:
4265:(2): 175–183.
4245:
4226:(6): 509–516.
4206:
4179:(3): 277–295.
4159:
4132:(2): 159–170.
4112:
4085:(4): 511–521.
4065:
4046:
4027:(2): 159–168.
4007:
3964:
3937:(4): 351–363.
3917:
3900:
3873:(3): 561–588.
3850:
3815:(3): 255–257.
3795:
3788:
3768:
3749:(3): 201–226.
3729:
3710:
3683:(4): 385–390.
3663:
3636:(4): F31–F38.
3616:
3569:
3562:
3556:. SUNY Press.
3542:
3515:(2): 168–181.
3495:
3474:10.1086/202545
3468:(5): 671–672.
3448:
3441:
3421:
3394:(4): 513–524.
3374:
3359:
3332:(6): 470–474.
3312:
3285:(5): 450–459.
3265:
3246:(2): 105–109.
3226:
3209:
3194:
3179:
3164:
3147:
3128:
3109:(5): 391–396.
3089:
3074:
3047:(4): 591–610.
3027:
3016:(7): 637–645.
2996:
2969:(3): 301–310.
2949:
2922:(2): 155–158.
2902:
2875:(2): 251–254.
2855:
2808:
2789:
2770:(2): 173–218.
2750:
2695:
2664:
2653:(4): 296–306.
2633:
2594:
2575:(3): 281–290.
2552:
2537:
2508:(2): 193–202.
2492:
2435:
2400:(9): 661–670.
2380:
2331:
2312:(3): 303–313.
2288:
2269:(3): 289–300.
2246:
2227:(2): 247–258.
2207:
2172:
2143:(2): 157–166.
2123:
2112:(9): 802–811.
2096:
2077:(2): 368–379.
2061:
2026:(3): 164–181.
2003:
1968:(3): 151–163.
1945:
1888:
1834:
1799:
1756:
1713:
1706:
1688:
1653:
1634:(4): 729–739.
1618:
1570:
1543:
1536:
1505:
1486:
1425:
1406:(6): 621–627.
1390:
1330:
1319:(3): 280–288.
1303:
1244:
1190:
1157:
1150:
1108:
1106:
1103:
1102:
1101:
1099:Animal culture
1094:
1091:
1069:
1066:
1051:
1048:
1026:
1023:
994:
991:
961:
958:
952:
949:
925:Tandem running
913:
910:
878:
875:
847:
844:
843:
842:
838:
835:
832:
824:
821:
818:
817:
814:
811:
805:
804:
801:
797:
796:
793:
789:
788:
785:
781:
780:
777:
774:
769:(Specifically
764:
761:
757:
756:
753:
750:
746:
745:
742:
739:
735:
734:
731:
727:
726:
723:
719:
718:
715:
711:
710:
707:
703:
702:
699:
698:Cracking nuts
695:
694:
691:
690:Cracking nuts
687:
686:
683:
680:
677:
673:
672:
669:
666:
662:
661:
658:
655:
651:
650:
647:
643:
642:
641:Marais (1969)
639:
635:
634:
631:
628:
624:
623:
620:
617:
611:
610:
607:
603:
602:
599:
595:
594:
591:
587:
586:
583:
582:Sign language
579:
578:
575:
571:
570:
567:
563:
562:
559:
555:
554:
551:
547:
546:
543:
539:
538:
535:
531:
530:
527:
523:
522:
519:
515:
514:
511:
507:
506:
505:Yerkes (1943)
503:
499:
498:
495:
491:
490:
487:
483:
482:
481:Kohler (1925)
479:
475:
474:
471:
467:
466:
463:
459:
458:
455:
454:Opening watch
451:
450:
447:
444:
439:(Specifically
433:
432:
429:
425:
424:
421:
417:
416:
413:
409:
408:
405:
404:Fear response
401:
400:
397:
393:
392:
389:
385:
384:
383:Carner (1955)
381:
377:
376:
373:
369:
368:
365:
362:
358:
357:
354:
350:
349:
348:Wright (1972)
346:
342:
341:
338:
334:
333:
330:
326:
325:
322:
318:
317:
314:
311:
301:
298:
294:
293:
290:
287:
284:
281:
264:
261:
244:
241:
228:
225:
201:
198:
165:
162:
158:Macaca mulatta
128:
125:
103:
100:
76:
73:
67:
64:
9:
6:
4:
3:
2:
5078:
5067:
5064:
5062:
5059:
5058:
5056:
5043:
5039:
5035:
5031:
5024:
5022:
5020:
5011:
5007:
5002:
4997:
4993:
4989:
4985:
4981:
4977:
4970:
4962:
4958:
4953:
4948:
4945:(2): 534–48.
4944:
4940:
4936:
4929:
4921:
4917:
4913:
4909:
4905:
4901:
4894:
4885:
4880:
4876:
4872:
4868:
4861:
4859:
4850:
4846:
4841:
4836:
4833:(1): 131–40.
4832:
4828:
4824:
4817:
4809:
4805:
4801:
4797:
4794:(4): 279–95.
4793:
4789:
4782:
4774:
4770:
4765:
4760:
4756:
4752:
4748:
4741:
4733:
4729:
4725:
4721:
4717:
4713:
4709:
4705:
4701:
4697:
4693:
4686:
4678:
4674:
4670:
4666:
4663:(4): 655–62.
4662:
4658:
4651:
4643:
4639:
4635:
4631:
4626:
4621:
4617:
4613:
4610:(7073): 153.
4609:
4605:
4601:
4597:
4591:
4583:
4579:
4575:
4571:
4567:
4563:
4559:
4555:
4551:
4547:
4540:
4532:
4528:
4524:
4520:
4516:
4512:
4505:
4503:
4494:
4490:
4486:
4482:
4478:
4474:
4467:
4459:
4455:
4451:
4447:
4443:
4439:
4435:
4431:
4428:(2): 151–74.
4427:
4423:
4416:
4401:
4397:
4390:
4382:
4378:
4374:
4370:
4366:
4362:
4358:
4354:
4350:
4343:
4335:
4331:
4327:
4323:
4319:
4315:
4311:
4307:
4303:
4296:
4288:
4284:
4280:
4276:
4272:
4268:
4264:
4260:
4256:
4249:
4241:
4237:
4233:
4229:
4225:
4221:
4217:
4210:
4202:
4198:
4194:
4190:
4186:
4182:
4178:
4174:
4170:
4163:
4155:
4151:
4147:
4143:
4139:
4135:
4131:
4127:
4123:
4116:
4108:
4104:
4100:
4096:
4092:
4088:
4084:
4080:
4076:
4069:
4061:
4057:
4050:
4042:
4038:
4034:
4030:
4026:
4022:
4018:
4011:
4003:
3999:
3995:
3991:
3987:
3983:
3979:
3977:
3968:
3960:
3956:
3952:
3948:
3944:
3940:
3936:
3932:
3928:
3921:
3913:
3912:
3904:
3896:
3892:
3888:
3884:
3880:
3876:
3872:
3868:
3864:
3857:
3855:
3846:
3842:
3838:
3834:
3830:
3826:
3822:
3818:
3814:
3810:
3806:
3799:
3791:
3785:
3781:
3780:
3772:
3764:
3760:
3756:
3752:
3748:
3744:
3740:
3733:
3725:
3721:
3720:Human Biology
3714:
3706:
3702:
3698:
3694:
3690:
3686:
3682:
3678:
3674:
3667:
3659:
3655:
3651:
3647:
3643:
3639:
3635:
3631:
3627:
3620:
3612:
3608:
3604:
3600:
3596:
3592:
3588:
3584:
3580:
3573:
3565:
3559:
3555:
3554:
3546:
3538:
3534:
3530:
3526:
3522:
3518:
3514:
3510:
3506:
3499:
3491:
3487:
3483:
3479:
3475:
3471:
3467:
3463:
3459:
3452:
3444:
3438:
3435:. JHU Press.
3434:
3433:
3425:
3417:
3413:
3409:
3405:
3401:
3397:
3393:
3389:
3385:
3378:
3370:
3363:
3355:
3351:
3347:
3343:
3339:
3335:
3331:
3327:
3323:
3316:
3308:
3304:
3300:
3296:
3292:
3288:
3284:
3280:
3276:
3269:
3261:
3257:
3253:
3249:
3245:
3241:
3237:
3230:
3222:
3221:
3213:
3205:
3198:
3190:
3183:
3175:
3168:
3160:
3159:
3151:
3143:
3139:
3132:
3124:
3120:
3116:
3112:
3108:
3104:
3100:
3093:
3085:
3078:
3070:
3066:
3062:
3058:
3054:
3050:
3046:
3042:
3038:
3031:
3023:
3019:
3015:
3011:
3007:
3000:
2992:
2988:
2984:
2980:
2976:
2972:
2968:
2964:
2960:
2953:
2945:
2941:
2937:
2933:
2929:
2925:
2921:
2917:
2913:
2906:
2898:
2894:
2890:
2886:
2882:
2878:
2874:
2870:
2866:
2859:
2851:
2847:
2843:
2839:
2835:
2831:
2827:
2823:
2819:
2812:
2804:
2800:
2793:
2785:
2781:
2777:
2773:
2769:
2765:
2761:
2754:
2746:
2742:
2738:
2734:
2730:
2726:
2722:
2718:
2714:
2710:
2706:
2699:
2691:
2687:
2683:
2679:
2675:
2668:
2660:
2656:
2652:
2648:
2644:
2637:
2629:
2625:
2621:
2617:
2613:
2609:
2605:
2598:
2590:
2586:
2582:
2578:
2574:
2570:
2566:
2559:
2557:
2548:
2541:
2533:
2529:
2525:
2521:
2516:
2511:
2507:
2503:
2496:
2488:
2484:
2479:
2474:
2470:
2466:
2462:
2458:
2454:
2450:
2446:
2439:
2431:
2427:
2423:
2419:
2415:
2411:
2407:
2403:
2399:
2395:
2391:
2384:
2376:
2372:
2367:
2362:
2358:
2354:
2350:
2346:
2342:
2335:
2327:
2323:
2319:
2315:
2311:
2307:
2303:
2299:
2292:
2284:
2280:
2276:
2272:
2268:
2264:
2257:
2250:
2242:
2238:
2234:
2230:
2226:
2222:
2218:
2211:
2203:
2199:
2195:
2191:
2187:
2183:
2176:
2168:
2164:
2159:
2154:
2150:
2146:
2142:
2138:
2134:
2127:
2119:
2115:
2111:
2107:
2100:
2092:
2088:
2084:
2080:
2076:
2072:
2065:
2057:
2053:
2049:
2045:
2041:
2037:
2033:
2029:
2025:
2021:
2017:
2010:
2008:
1999:
1995:
1991:
1987:
1983:
1979:
1975:
1971:
1967:
1963:
1959:
1952:
1950:
1941:
1937:
1932:
1927:
1923:
1919:
1915:
1911:
1907:
1903:
1899:
1892:
1884:
1880:
1876:
1872:
1868:
1864:
1860:
1856:
1852:
1848:
1841:
1839:
1830:
1826:
1822:
1818:
1815:(4): 355–72.
1814:
1810:
1803:
1795:
1791:
1787:
1783:
1779:
1775:
1771:
1767:
1760:
1752:
1748:
1744:
1740:
1736:
1732:
1729:(4): 235–44.
1728:
1724:
1717:
1709:
1703:
1699:
1692:
1684:
1680:
1676:
1672:
1668:
1664:
1657:
1649:
1645:
1641:
1637:
1633:
1629:
1622:
1614:
1610:
1606:
1602:
1599:(9): 486–93.
1598:
1594:
1587:
1585:
1583:
1581:
1579:
1577:
1575:
1566:
1562:
1559:(5): 373–93.
1558:
1554:
1547:
1539:
1533:
1529:
1525:
1521:
1514:
1512:
1510:
1501:
1497:
1490:
1482:
1478:
1474:
1470:
1466:
1462:
1457:
1452:
1448:
1444:
1440:
1436:
1429:
1421:
1417:
1413:
1409:
1405:
1401:
1394:
1386:
1382:
1377:
1372:
1368:
1364:
1360:
1356:
1352:
1348:
1344:
1340:
1334:
1326:
1322:
1318:
1314:
1307:
1299:
1295:
1290:
1285:
1281:
1277:
1272:
1267:
1263:
1259:
1255:
1248:
1240:
1236:
1232:
1228:
1224:
1220:
1216:
1212:
1205:
1203:
1201:
1199:
1197:
1195:
1185:
1180:
1176:
1172:
1168:
1161:
1153:
1147:
1143:
1136:
1134:
1132:
1130:
1128:
1126:
1124:
1122:
1120:
1118:
1116:
1114:
1109:
1100:
1097:
1096:
1090:
1088:
1082:
1080:
1075:
1065:
1056:
1047:
1045:
1041:
1040:Red squirrels
1037:
1033:
1022:
1020:
1016:
1015:common cuckoo
1012:
1008:
1004:
1000:
986:
982:
980:
979:Turdus merula
976:
972:
967:
957:
948:
946:
942:
938:
933:
930:
926:
918:
909:
907:
906:domestic cats
903:
898:
896:
892:
883:
874:
872:
867:
865:
861:
857:
856:killer whales
852:
839:
836:
833:
830:
829:
828:
815:
812:
810:
807:
806:
802:
799:
798:
794:
792:Tube opening
791:
790:
786:
783:
782:
778:
775:
772:
768:
758:
754:
751:
748:
747:
743:
740:
737:
736:
732:
729:
728:
724:
721:
720:
716:
713:
712:
708:
705:
704:
700:
697:
696:
692:
689:
688:
684:
681:
674:
670:
667:
664:
663:
659:
656:
653:
652:
648:
645:
644:
640:
637:
636:
632:
629:
625:
621:
618:
616:
613:
612:
608:
605:
604:
600:
597:
596:
592:
589:
588:
584:
581:
580:
576:
573:
572:
568:
565:
564:
560:
558:Limping gait
557:
556:
552:
549:
548:
544:
541:
540:
536:
533:
532:
528:
525:
524:
520:
517:
516:
513:Hayes (1951)
512:
509:
508:
504:
501:
500:
496:
493:
492:
488:
485:
484:
480:
477:
476:
473:Sheak (1923)
472:
469:
468:
464:
461:
460:
456:
453:
452:
448:
445:
442:
438:
434:
430:
427:
426:
422:
419:
418:
414:
411:
410:
406:
403:
402:
398:
395:
394:
390:
387:
386:
382:
379:
378:
374:
371:
370:
366:
363:
359:
355:
352:
351:
347:
344:
343:
339:
336:
335:
331:
328:
327:
323:
320:
319:
316:Abel (1818)
315:
312:
309:
305:
295:
291:
288:
285:
282:
279:
278:
273:
270:
260:
256:
252:
250:
240:
238:
233:
224:
222:
216:
214:
209:
207:
197:
193:
189:
185:
181:
174:
170:
161:
159:
155:
145:
141:
139:
135:
134:greylag geese
124:
121:
116:
114:
113:Rattus rattus
110:
99:
96:
94:
90:
81:
72:
63:
61:
56:
52:
50:
46:
42:
38:
34:
30:
26:
22:
18:
5033:
5029:
4983:
4979:
4969:
4942:
4938:
4928:
4906:(4): 432–9.
4903:
4899:
4893:
4877:(5): 493–9.
4874:
4870:
4830:
4826:
4816:
4791:
4787:
4781:
4754:
4750:
4740:
4699:
4695:
4685:
4660:
4656:
4650:
4607:
4603:
4590:
4549:
4545:
4539:
4514:
4510:
4479:(2): 326–7.
4476:
4472:
4466:
4425:
4421:
4415:
4403:. Retrieved
4399:
4389:
4356:
4352:
4342:
4309:
4305:
4295:
4262:
4258:
4248:
4223:
4219:
4209:
4176:
4172:
4162:
4129:
4125:
4115:
4082:
4078:
4068:
4059:
4055:
4049:
4024:
4020:
4010:
3985:
3981:
3976:Cebus apella
3975:
3967:
3934:
3930:
3920:
3914:. Macmillan.
3910:
3903:
3870:
3866:
3812:
3808:
3798:
3778:
3771:
3746:
3742:
3732:
3723:
3719:
3713:
3680:
3676:
3666:
3633:
3629:
3619:
3589:(1): 35–50.
3586:
3582:
3572:
3552:
3545:
3512:
3508:
3498:
3465:
3461:
3451:
3431:
3424:
3391:
3387:
3377:
3368:
3362:
3329:
3325:
3315:
3282:
3278:
3268:
3243:
3239:
3229:
3219:
3212:
3203:
3197:
3188:
3182:
3173:
3167:
3157:
3150:
3141:
3137:
3131:
3106:
3102:
3092:
3083:
3077:
3044:
3040:
3030:
3013:
3009:
2999:
2966:
2962:
2952:
2919:
2915:
2905:
2872:
2868:
2858:
2825:
2821:
2811:
2802:
2798:
2792:
2767:
2763:
2753:
2712:
2708:
2698:
2684:(1): 19–33.
2681:
2677:
2667:
2650:
2646:
2636:
2614:(1): 57–68.
2611:
2607:
2597:
2572:
2568:
2546:
2540:
2505:
2501:
2495:
2452:
2448:
2438:
2397:
2393:
2383:
2348:
2344:
2334:
2309:
2305:
2302:Homo sapiens
2301:
2297:
2291:
2266:
2262:
2249:
2224:
2220:
2216:
2210:
2188:(1): 74–79.
2185:
2181:
2175:
2140:
2136:
2126:
2109:
2105:
2099:
2074:
2070:
2064:
2023:
2019:
1965:
1961:
1905:
1901:
1891:
1850:
1846:
1812:
1808:
1802:
1769:
1765:
1759:
1726:
1722:
1716:
1697:
1691:
1666:
1662:
1656:
1631:
1627:
1621:
1596:
1592:
1556:
1552:
1546:
1519:
1499:
1489:
1438:
1434:
1428:
1403:
1399:
1393:
1350:
1346:
1333:
1316:
1312:
1306:
1261:
1258:PLOS Biology
1257:
1247:
1214:
1210:
1174:
1170:
1160:
1141:
1083:
1071:
1062:
1043:
1035:
1028:
1018:
1010:
996:
978:
963:
954:
941:waggle dance
934:
923:
899:
894:
888:
868:
860:Orcinus orca
859:
853:
849:
826:
755:Beck (1974)
744:Beck (1972)
682:Box opening
633:Hall (1963)
415:Beck (1976)
289:Behavior(s)
266:
257:
253:
248:
246:
236:
234:
230:
220:
217:
212:
210:
205:
203:
194:
190:
186:
182:
178:
157:
150:
137:
130:
117:
112:
105:
97:
92:
86:
69:
57:
53:
16:
15:
2828:(1): 1–29.
2715:(916): 53.
1456:10023/12387
1217:(1): 3–15.
771:Chimpanzees
441:Chimpanzees
138:Anser anser
47:(including
5055:Categories
4517:(4): 767.
3371:: 144–184.
1669:: 327–36.
1177:(6): 489.
1171:BioScience
1105:References
1059:increases.
1032:Norway rat
760:Emulation
297:Imitation
109:black rats
83:Male guppy
41:amphibians
23:, such as
4939:Evolution
4732:206519140
4493:144947666
4405:24 August
4373:0393-9375
4326:1939-2087
4279:0393-9375
4240:0047-2484
4193:0032-8332
4146:1939-2087
4099:0032-8332
4041:0735-7036
4002:1421-9980
3951:0164-0291
3895:145467809
3887:0140-525X
3829:0275-2565
3697:1939-2087
3650:1363-755X
3603:0032-8332
3529:0032-8332
3490:145349296
3482:0011-3204
3408:0032-8332
3346:0021-9940
3299:0021-9940
3252:0003-049X
3223:. Harper.
3123:0095-9928
3061:1939-1846
2983:0032-8332
2936:1097-4679
2889:0022-1325
2842:0032-8332
2737:0028-0836
2628:1748-1090
2581:0003-049X
2510:CiteSeerX
2469:0962-8436
2414:1471-003X
2040:1435-9448
1982:1435-9448
1922:0962-8436
1465:0149-7634
1441:: 58–75.
1420:0179-1613
1367:0962-8436
1280:1545-7885
966:predators
864:stranding
763:Positive
676:Negative
353:Tool use
308:Orangutan
300:Positive
292:Citation
173:Orangutan
5061:Ethology
5010:12495513
4961:19674093
4849:15161148
4773:19185495
4724:19498167
4677:53186108
4634:16407943
4582:11490465
4574:16840701
4531:53162892
4458:40567375
4381:84534346
4287:84211164
4201:42897869
4173:Primates
4107:19285572
4079:Primates
3959:23901545
3845:84278221
3837:31986808
3763:14051442
3705:19929107
3658:17552931
3611:21156104
3583:Primates
3537:25897929
3509:Primates
3416:22621124
3388:Primates
3354:13109075
3307:13000013
3144:: 45–55.
2991:35080987
2963:Primates
2944:13631102
2897:14434758
2850:30040660
2822:Primates
2532:21895355
2487:19620103
2422:11533734
2375:22983208
2326:16893268
2283:16045384
2241:15085541
2202:19271818
2167:30337770
2106:Ethology
2091:25164939
2048:15549502
1998:17599865
1990:15490290
1940:19620112
1883:33299917
1875:17752463
1794:20465133
1786:10877890
1743:13680401
1683:53148456
1648:53169300
1613:18657877
1473:28034660
1400:Ethology
1385:21357219
1298:27701411
1239:38321280
1231:11170692
1093:See also
1003:cowbirds
902:cheetahs
891:meerkats
871:chickens
846:Teaching
646:Fishing
120:tool use
60:teaching
49:primates
37:reptiles
5001:1693065
4920:3802787
4808:7049794
4704:Bibcode
4696:Science
4642:4416276
4612:Bibcode
4554:Bibcode
4546:Science
4450:1635977
4442:2831436
4334:8370271
4154:2736909
3260:3143327
3069:4078162
2784:1412738
2745:4112014
2717:Bibcode
2647:Mankind
2478:2865083
2430:6792943
2366:3458123
2158:6191197
2056:1949770
1931:2865074
1855:Bibcode
1847:Science
1829:6542574
1376:3049099
1289:5049772
1079:finches
999:cuckoos
973:in the
749:Macaca
615:Gorilla
89:guppies
45:mammals
25:insects
5008:
4998:
4959:
4918:
4847:
4806:
4771:
4730:
4722:
4675:
4640:
4632:
4604:Nature
4580:
4572:
4529:
4491:
4456:
4448:
4440:
4379:
4371:
4332:
4324:
4285:
4277:
4238:
4199:
4191:
4152:
4144:
4105:
4097:
4062:: 325.
4039:
4000:
3957:
3949:
3893:
3885:
3843:
3835:
3827:
3786:
3761:
3703:
3695:
3656:
3648:
3609:
3601:
3560:
3535:
3527:
3488:
3480:
3439:
3414:
3406:
3352:
3344:
3305:
3297:
3258:
3250:
3121:
3067:
3059:
2989:
2981:
2942:
2934:
2895:
2887:
2848:
2840:
2782:
2743:
2735:
2709:Nature
2626:
2589:984118
2587:
2579:
2530:
2512:
2485:
2475:
2467:
2428:
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