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was occupied elsewhere. This was supported by the fact that the infant picked up the stone once the mother dropped it and allowed her to groom him while he played with it. This behavior was seen in a few other members of the colony, but not seen throughout the species. In another instance, a female chimpanzee at the Delta
Regional Primate Research Center created a "toothbrush" by stripping a twig of its leaves, and used this toothbrush to groom her infant over several instances. However, both examples concern tool use in primates, which is already widely studied and scientifically backed. The wide working memory capacities and causal understanding capabilities of primates permit them to fashion and utilize tools far more extensively than other non-human animals. Apart from physical and mental constraints, perhaps a reason allogrooming animals do not use tools is because a major purpose of social grooming is social bonding and involves emotional exchanges, much of which is conveyed by touch.
928:, an opioid antagonist that blocks the opioid receptor and inhibits the effects of endogenous opioids. In comparison to the control females, who were given saline solutions, the naloxone females groomed their infants and other members of their group less. The naloxone females were also observed to be less protective of their young, which is uncharacteristic of new mothers. This decline in social interactions upon naloxone injection suggests that opioid antagonists interfere with maternal involvement in social actions—here, social grooming. it could therefore be hypothesized that higher levels of opioids in new rhesus mothers cause increased levels of social involvement and maternal behavior, aiding the development and learning of the newborn.
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mates; they chose grooming mates based on who would reciprocate rather than who would not. More importantly, if the delay between two chimpanzees grooming each other is very little, then the chimpanzees tend to "time match": i.e., the second groomer grooms the first for the same amount of time that he/she was groomed. This "episodic memory" requires a demanding amount of cognitive function and emotional recognition, and has been tested experimentally with respect to food preferences, where apes chose between tasty perishable and non-tasty non-perishable food at shorter and longer delays respectively after trying the food. Hence, apes can distinguish between different events that occurred at different times.
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groomed by members of its group. However, it has instead been seen that when group size increases, members ensure that they spend an appropriate amount of time grooming everyone. Hence, the fact that animals, particularly primates, groom each other more frequently than necessary from a hygienic standpoint suggests that the social aspect of allogrooming plays an equally, if not more, important role. Another point of evidence for the importance of the social aspect is that in comparison to how and how much a primate grooms itself (autogrooming), allogrooming involves longer periods of time and different techniques, some of which have connotations of being affectionate gestures.
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940:, and a narrow range of species within primates themselves. As a result, the literature does not provide a well-rounded idea of what the cognitive or behavioral basis for social grooming is, nor does it completely outline all of its effects, positive or negative. Even in well-studied species, it may be that not all the data relevant to social grooming has been collected. Secondly, data for most species is derived based on the members of a single group. In primates, whose behavior is highly flexible depending on the socio-environmental conditions, this poses a particular challenge. Thirdly, most studies are short-term and
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259:, it was seen that more dominant group members were "stroked" more than they were "picked at" when being groomed, compared to lower-ranking group members. From a utilitarian standpoint, stroking is a less effective technique for grooming than picking, but it is construed as being a more affectionate gesture. Hence, grooming a higher ranking individual could be done in order to placate a potential aggressor and reduce tension. Moreover, individuals closer in rank tend to groom each other more reciprocally than individuals further apart in rank.
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pressure levels than rats who did not receive any touch. This was found to be the result of an increased vagal nerve tone, meaning they had had a higher parasympathetic nervous response and a lower sympathetic nervous response to stimuli, resulting in a lower stress response. Social grooming is a form of innocuous sensory activation. Innocuous sensory activation, characterized by non-aggressive contact, stimulates an entirely separate neural pathway from nocuous aggressive sensory activation. Innocuous sensations are transmitted through the
499:, infant females mimic their mothers' actions by grooming their mothers more often than their male counterparts do and by grooming the same group members that their mothers groom. This mimicry is suggested to indicate identification-based observational learning in infant stump-tailed macaques, and the daughters' penchants for maternal mimicry and kin-biased grooming versus the sons' penchants for rank-biased grooming falls in line with their social roles in groups, where adult males require alliances in order to gain and maintain rank.
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indicative of higher stress; however, females with reliable and well-established grooming partners have less of a fGC rise than those with weaker grooming networks. Hence, the social support received from a "friendship" aids baboons in stress management. Similarly, fGC levels are also seen to rise in females when a close "friend" dies; however, these rising fGC levels are seen to decrease in females that form new grooming partners, replacing their deceased friends.
419:, this rule governs the idea that kin selection causes genes to increase in frequency when the genetic relatedness (r) of a recipient to an actor multiplied by the benefit to the recipient (B) is greater than the reproductive cost to the actor (C). Thus, it is advantageous for an individual to partake in altruistic behaviors, such as social grooming, so long as the individual receiving the benefits of the behavior is related to the one providing the behavior.
280:, infants are seen as a valuable commodity that can be exchanged for favours; mothers allow non-mothers to handle their infants for short durations in exchange for being groomed. Tibetan macaques measure and perceive the value of the infants by noting the relative ratio of infants in the group; as the number of infants increase, their "value" decreases, and so does the amount of grooming performed by non-mothers for mothers in exchange for infant-handling.
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682:, for example, engages in social grooming by cleaning body parts that cannot be reached by the receiving bee. The receiving bee extends its wings perpendicular to its body while its wings, mouth parts, and antennae are cleaned in order to remove dust and pollen. This removal of dust and pollen allows for the sharpening of olfactory senses, thus contributing to the overall well-being of the group.
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include relaxation, healing, and digestion stimulation. Reproductive benefits have also been found: studies in rats have shown that the release of oxytocin can increase male reproductive success. Oxytocin plays an important role in maternal pair bonding, and is hypothesized to promote similar bonding in social groups as a result of positive feedback loops from social interactions.
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shown that increased social stress results in glucocorticoid resistance, further inhibiting immune function. Macaques who participated in social grooming showed decreased levels of viral load, which points toward decreased levels of social stress resulting in increased immune function and glucocorticoid sensitivity. Additionally, a 1997 study concluded that an increase in
227:, the females initiate grooming with the males just before flight at dusk. The male and his close-knit female harem release bodily secretions onto each other, which may allow them to recognize the female's reproductive status. A 2016 study by Kumar et al. chemically analyzed these secretions, concluding that they may be required in chemosensory mediated communication and
208:(associated with social and emotional processing/analysis) when the monkeys were shown pictures of their friends' faces, compared to when they were shown less familiar faces. Hence, primates recognize familiar and well-liked individuals ("friends") and spend more time grooming them than less favoured partners. In species with a more tolerant social style, such as
162:. Social affiliation during a mild stressor was shown to correlate with lower levels of mammary tumor development and longer lifespan in rats, while lack of this affiliation was demonstrated to be a major risk factor. On the other hand, it could be argued that the hygienic aspect of allogrooming does not play as important a role as the social aspect of it.
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constraints and predation pressure strongly affect group sizes and thereby have an indirect effect on primate grooming time". By analyzing past data and studies done on this topic, the authors found that a primate group greater than 40 will face greater ecological problems and, thus, time spent during social grooming is affected.
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receiving significant attention appear to be the regions where the baboons themselves cannot reach. Grooming activity in these regions is used to remove parasites, dirt, dead skin, and tangled fur in order to help keep the animal's health in good condition despite the individual's inability to reach and clean certain areas.
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231:. Similarly, in the less aggressive herb-field mouse, males are observed to groom females for longer durations and even allow females to not reciprocate. Since the mating demands of males are greater than those offered by females, the females use social grooming to choose mates and males use it to incite mating.
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position. It has been found that in times of higher conflict and competition, this is less likely to occur. Researchers have suggested that primates may see a need to balance the uses of grooming, swapping between its use as a means to increase social standing and its use as a means to keep oneself clean.
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is not as important a factor as friendship or mate preference when choosing a grooming mate as previously thought. In a 2018 study of captive chimpanzees, Phelps et al. found that the animals remembered interactions that were "successful" or "unsuccessful" and used these as a basis to choose grooming
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One of the most critical functions of social grooming is to establish social networks and relationships. In many species, individuals form close social connections dubbed "friendships" due to long periods of time spent together doing activities. In primates especially, grooming is known to have major
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It is often argued as to whether the overarching importance of social grooming is to boost an organism's health and hygiene or whether the social side of social grooming plays an equally or more important role. Traditionally, it is thought that the primary function of social grooming is the upkeep of
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Grooming stimulates the release of beta-endorphins, which is one physiological reason for the relaxing effect of grooming. Beta-endorphins are found in neurons in the hypothalamus and the pituitary gland. Beta-endorphins are found to be opioid agonists. Opioids are molecules that act on receptors to
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and pre-existing coalitions, and for reconciliation after conflicts. Primates groom socially in moments of boredom as well, and the act has been shown to reduce tension and stress. This reduction in stress is often associated with observed periods of relaxed behavior, and primates have been known to
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social grooming results in the formation of crucial relationships among partners. These social relationships serve to aid cooperation and facilitate protection against combative groups composed of other males, which can oftentimes cause physical harm. Furthermore, social relationships have also been
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It has been suggested that in male bonobos, grooming is exchanged in favour of some emotional component because grooming familiar individuals involves larger time differences (i.e., the duration for which each individual grooms the other is not equal) and reduced reciprocity (i.e., the likelihood of
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Recent studies regarding chimpanzees have determined the direct correlation of the release of oxytocin to consolatory behavior. This behavior, as well as release, has been noted in primates such as the Vervet monkey, a primate species that actively engages in social grooming from early childhood to
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Grooming in primates is not only utilized for alliance formation and maintenance, but to exchange resources such as communal food, sex, and hygiene. Wild baboons have been found to utilize social grooming as an activity to remove ticks and other insects from others; in this grooming, the body areas
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at
Bucknell University, a mother macaque was seen to choose a stone after observing several stones on the ground, and then use this stone to groom her infant. It was hypothesized that the stone was used as a distractor for the infant so that the mother could adequately clean him while his attention
373:' reciprocal altruism theory under the title "tit-for-tat". In conjunction with altruism, kin selection bears an emphasis on favoring the reproductive success of an organism's relatives, even at a cost to the organism's own survival and reproduction. Because of this, kin selection is an instance of
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Social bonds established by grooming may provide an adaptive advantage in the form of conflict resolution and protection from aggression. In wild savannah baboons, social affiliations were shown to augment fitness by increasing tolerance from more dominant group members and increasing the chance of
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groom each other in the form of stroking, scratching, and massaging. This activity often serves to remove foreign material from the body to promote the communal success of these socially active animals. There exists a wide array of socially grooming species throughout the animal kingdom, including
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performed on 44 different primate species suggest that the number of times a species allogrooms, on average, correlates with its group size rather than with its body size. If allogrooming was required from a purely hygienic standpoint, then the larger an animal, the more and more often it would be
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In most cases, allogrooming is an action that is learned from an individual's mother. Infants are groomed by their mothers and mimic these actions on each other and the mothers as juveniles. This action is reciprocated on other group members (non-mother or of a different rank) more often once the
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Faecal glucocorticoid (fGCs) is a hormone metabolite associated with stress that is seen to be present in lower levels in female baboons with stronger, well-established grooming networks. When potentially infanticidal male baboons immigrate into a group, the females' fGC levels are seen to rise,
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are steroid hormones that are synthesized in the adrenal cortex and are a part of the group of corticosteroids. Glucocorticoids are involved in immune function, and are a part of the feedback system that reduces inflammation. They are also involved in glucose metabolism. Studies in macaques have
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have shown that the intranasal injection of oxytocin increases the amount of allogrooming done by female bats. The release of oxytocin, found to be stimulated by positive touches (such as allogrooming), smells, and sounds, can provide physiological benefits for the individual; these benefits can
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has on infant survival. A positive relationship is established between infant survival to one year and a composite sociality index, a measure of sociality based on proximity and social grooming. Evidence has also been provided for the effect of sociality on adult survival in wild baboons. Direct
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Allogrooming is a behavior commonly seen in many types of cattle, including dairy and beef breeds. The act of social licking can be seen specifically in heifers to initiate social dominance, emphasize companionship and improve hygiene of oneself or others. This behavior seen in cows may provide
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Recent studies have determined that vampire bats engage in social grooming much more than other types of bats to promote the well-being of the group. Facing higher levels of parasitic infection, vampire bats engage in cleaning one another as well as sharing food via regurgitation. This activity
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which promote physiological responses in stress reduction. These responses can occur from the production of hormones and endorphins, or through the growth or reduction in nerve structures. For example, in studies of suckling rats, rats who received warmth and touch when feeding had lower blood
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There are benefits to initiating grooming. The one that starts the grooming will in return be groomed themselves, getting the benefit of being cleaned. Research has found that primates lower on the social ladder may initiate grooming with a higher-ranking primate in order to increase their own
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Primates provide perhaps one of the best examples of mutual grooming, due to the intensive research performed regarding their varying lifestyles and the direct variation in the means of social grooming across different species. Among primates, social grooming plays a significant role in animal
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blockades, which decrease the level of beta-endorphins, monkeys were observed to respond with an increased desire to be groomed; conversely, the monkeys' desire to be groomed reduced significantly when they were given morphine. However, beta-endorphin levels are difficult to measure in animal
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Grooming is often offered by an individual in exchange for a certain behavioral response or action. Social grooming is critical for vampire bats especially, since it is necessary for them to maintain food-sharing relationships in order to sustain their food regurgitation sharing behavior. In
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The time primates spend grooming increases with group size, but too-large group sizes can lead to decreased group cohesion because time spent grooming is usually impacted by other factors, which include ecological, phylogenetic, and life history. For example, the article states, "Cognitive
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are chemical molecules produced in the brains of organisms that serve to create feelings of relaxation, happiness, and pain relief. In primates, laughter and social grooming trigger opioid release in the brain, which is thought to form and maintain social bonds. In a study performed on
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primates, insects, birds, and bats. While thorough research has yet to be conducted, much has been learned about social grooming in non-human animals via the study of primates. The driving force behind mammalian social grooming is primarily believed to be rooted in adaptation to
212:, it is seen that females choose their grooming mates based on whom they know better rather than on social rank. In addition to primates, animals such as deer, cows, horses, voles, mice, meerkats, coatis, lions, birds, and bats also form social bonds through grooming behavior.
798:. In 2010, researchers determined the existence of a form of social grooming as a consolation behavior within ravens via a form of bystander contact, whereby observer ravens would act to console a distressed victim via contact sitting, preening, and beak-to-beak touching.
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have been proven to contribute to greater social cohesion and stability. Groups of gibbons with more stable social networks formed grooming networks that were significantly more complex, while groups with low stability networks formed far fewer grooming pairs.
300:) has been studied extensively, with numerous studies showing an increase in fitness as a result of social bonds formed through social grooming behavior. One such study, which collected 16 years of behavioral data on wild baboons, highlighted the effects that
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Guan ZH, Huang B, Ning WH, Ni QY, Sun GZ, Jiang XL (December 2013). "Significance of grooming behavior in two polygynous groups of western black crested gibbons: Implications for understanding social relationships among immigrant and resident group members".
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siblings often have conflicts over grooming allocation by their mother, yet grooming remains an activity that mediates tension and is low cost for alliance formation and maintenance. This grooming occurs both between siblings and between mother and child.
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438:. The grooming was evenly balanced across multiple rather than single bouts, suggesting that females are not constrained to complete exchanges with single transactions and use social grooming to solidify long-term relationships with those in their
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Social grooming is considered a behavior of facultative altruism—the behavior itself is a temporary loss of direct fitness (with potential for indirect fitness gain), followed by personal reproduction. This tradeoff has been compared to the
255:. Researchers have observed that in this system, dominant males receive more grooming while grooming others less, thereby indicating that less dominant males groom more dominant individuals to maintain relationships. In a study conducted on
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have shown that males will groom females in order to procure sex. One study found that a female has a greater likelihood of engaging in sexual activity with a male if he has recently groomed her, compared to males who have not.
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on corticosteroid secretion and prevent the undesirable side effects of an abnormal physiologic stress response. Social grooming can change the number of glucocorticoid receptors, which can result in increased immune function.
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promote feelings of relaxation and reduce pain. A study in monkeys found that the changes in opiate expression in the body, mirroring changes in beta-endorphin levels, influences desire for social grooming. When injected with
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advantages including reduced parasite loads, social tension, and competition at the feed bunk. It is understood that social licking can provide long-term benefits such as promoting positive emotions and a relaxed environment.
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Martel, Frances L.; Nevison, Claire M.; Rayment, F. David; Simpson, Michael J. A.; Keverne, Eric B. (1 January 1993). "Opioid receptor blockade reduces maternal affect and social grooming in rhesus monkeys".
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Liu D, Diorio J, Tannenbaum B, Caldji C, Francis D, Freedman A, et al. (September 1997). "Maternal care, hippocampal glucocorticoid receptors, and hypothalamic-pituitary-adrenal responses to stress".
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Social grooming behavior has been shown to elicit an array of health benefits in a variety of species. For example, group member connection has the potential to mitigate the potentially harmful effects of
810:" where parasites and other contaminants on the surface of the body are actively removed. This removal of foreign material is primarily performed on hard-to-reach areas, such as the neck, via nibbling.
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during instances of within-group contest interactions. In the yellow baboon, adult females form relationships with their kin, who offer support during times of violent conflict within social groups. In
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Manninen, Sandra; Tuominen, Lauri; Dunbar, Robin I.; Karjalainen, Tomi; Hirvonen, Jussi; Arponen, Eveliina; Hari, Riitta; Jääskeläinen, Iiro P.; Sams, Mikko; Nummenmaa, Lauri (21 June 2017).
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In nearly all instances of social grooming, individuals use their own body parts, such as hands, teeth, or tongue, to groom a group member or infant. It is very rare to observe instances of
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Mondragón-Ceballos, Ricardo; Chiappa, Pilar; Mayagoitia, Lilian; Lee, Phyllis (1 July 2010). "Sex differences in learning the allocation of social grooming in infant stumptailed macaques".
2444:"Origin of Species : Chapter VIII. Instinct : Objections to the theory of natural selection as applied to instincts: neuter and sterile insects by Charles Darwin @ Classic Reader"
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Val-Laillet D, Guesdon V, Keyserlingk MA, Passillé AM, Rushen J (2009). "Allogrooming in cattle: Relationships between social preferences, feeding displacements and social dominance".
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Oxytocin is a peptide hormone known to help express social emotions such as altruism, which in turn provide a positive feedback mechanism for social behaviors. For example, studies of
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Social grooming has been shown to be correlated with changes in endocrine levels within individuals. Specifically, there is a large correlation between the brain's release of
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fall asleep while receiving grooming. Conflict among primates has been observed by researchers as increasing stress among the group, making mutual grooming very advantageous.
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and social grooming. Oxytocin is hypothesized to promote prosocial behaviors due to its positive emotional response when released. Further, social grooming also releases
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of the females in their group. Though the behavior of social grooming itself was not beneficial to the one providing the service, the opportunity to mate and subsequent
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grooming the other is unpredictable). Hence, the presence of some sort of social bond between individuals results in greater "generosity" and tolerance between them.
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species—in contrast to oxytocin, which can be measured by sampling cerebrospinal fluid—and therefore they have not been linked as strongly with social behaviors.
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correlations between measures of social connectedness (which focuses on social grooming) and median survival time for both female and male baboons were modeled.
93:, indicates social grooming between members of the same species. Grooming is a major social activity and a means by which animals who live in close proximity may
146:, and that the amount of time spent allogrooming regions did not vary significantly even if the body part had a more important social or communicatory function.
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Laister S, Stockinger B, Regner AM, Zenger K, Knierim U, Winckler C (2011). "Social licking in dairy cattle—Effects on heart rate in performers and receivers".
349:, which requires the conscious intention to help another. As a behavior, altruism is not evaluated in moral terms, but rather as a consequence of an action for
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Keverne EB, Martensz ND, Tuite B (1 January 1989). "Beta-endorphin concentrations in cerebrospinal fluid of monkeys are influenced by grooming relationships".
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In general, social grooming is an activity that is directed up-hierarchy—i.e., a lower ranking individual grooms a higher ranking individual in the group. In
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It has been questioned whether some animals are instead using altruistic behaviors as a market strategy to trade for something desirable. In olive baboons,
944:, so the direct link between social grooming and fitness or mate choice outcomes cannot be studied directly as in long-term direct or captive studies.
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Keverne EB, Martensz ND, Tuite B (1989). "Beta-endorphin concentrations in cerebrospinal fluid of monkeys are influenced by grooming relationships".
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Keverne EB, Martensz ND, Tuite B (1989). "Beta-endorphin concentrations in cerebrospinal fluid of monkeys are influenced by grooming relationships".
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it has been found that individuals perform altruistic behaviors as a form of trade in which a behavior is provided in exchange for benefits, such as
377:, which combines the number of offspring produced with the number an individual can ensure the production of by supporting others, such as siblings.
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an animal's hygiene. Evidence to support this statement involves the fact that all grooming concentrates on body parts that are inaccessible by
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on target tissue in the neonatal rat. In the study on neonatal rats, it was found that the receptor number was altered due to a change in both
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Moore D, Angel JE, Cheeseman IM, Robinson GE, Fahrbach SE (1995). "A highly specialized social grooming honey bee (Hymenoptera: Apidae)".
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Berghänel A, Ostner J, Schröder U, Schülke O (2011). "Social bonds predict future cooperation in male
Barbary macaques, Macaca sylvanus".
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Moore D, Angel JE, Cheeseman IM, Robinson GE, Fahrbach SE (1995). "A highly specialized social grooming honey bee(Hymenoptera: Apidae)".
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in social grooming in non-human animals; however, a few such instances have been observed in primates. In a 1981 observational study of
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Ray JC, Sapolsky RM (1 January 1992). "Styles of male social behavior and their endocrine correlates among high-ranking wild baboons".
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Kutsukake N, Clutton-Brock TH (1 February 2010). "Grooming and the value of social relationships in cooperatively breeding meerkats".
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of another individual while decreasing the fitness of the one performing the behavior. This differs from the philosophical concept of
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Social grooming relationships have been proven to provide direct fitness benefits to a variety of species. In particular, grooming in
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Schino G, Scucchi S, Maestripieri D, Turillazzi PG (1988). "Allogrooming as a tension-reduction mechanism: a behavioral approach".
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Schino G, Scucchi S, Maestripieri D, Turillazzi PG (1988). "Allogrooming as a tension-reduction mechanism: a behavioral approach".
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Barelli C, Reichard UH, Mundry R (1 October 2011). "Is grooming used as a commodity in wild white-handed gibbons, Hylobates lar?".
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Dunbar RI (February 2010). "The social role of touch in humans and primates: behavioral function and neurobiological mechanisms".
3877:"Social stress results in altered glucocorticoid regulation and shorter survival in simian acquired immune deficiency syndrome"
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of those participating in the behavior. This study also found that social grooming performance cycled with that of the females
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3836:"How do glucocorticoids influence stress responses? Integrating permissive, suppressive, stimulatory, and preparative actions"
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Silk JB, Alberts SC, Altmann J (1 March 2004). "Patterns of coalition formation by adult female baboons in
Amboseli, Kenya".
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Studies have also shown that male baboons who participate more in social grooming show lower basal cortisol concentrations.
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Aureli F, Van Schaik CP, Van Hooff JA (1989). "Functional aspects of reconciliation among captive long-tailed macaques (
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2553:"Impatient traders or contingent reciprocators? Evidence for the extended time-course of grooming exchanges in baboons"
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Kumar, Rathina; Cantor, Mauricio; Senthilkumar, K.; Panneerselvam, Vimal; Kaliraj, P.; Marimuthu, G. (1 January 2017).
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Aureli F, van Schaik C, van Hooff J (1989). "Functional aspects of reconciliation among captive long-tailed macaques (
3365:"Allopreening in birds is associated with parental cooperation over offspring care and stable pair bonds across years"
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97:, reinforce social structures and family links, and build companionship. Social grooming is also used as a means of
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Physiology and
Pharmacology for Anesthesia: Foundations and Clinical Application: Expert Consult – Online and Print
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Feh, C., De
Mazieres, J. Grooming at a preferred site reduces heart rate in horses. Anim. Behav. 1993;46:1191–1194.
2042:"Social affiliation matters: both same-sex and opposite-sex relationships predict survival in wild female baboons"
1248:"A functional analysis of social grooming patterns through direct comparison with self-grooming in rhesus monkeys"
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Uvnäs-Moberg K (November 1998). "Oxytocin may mediate the benefits of positive social interaction and emotions".
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and stress levels, maintenance of social structures, and direct improvement of fitness as a measure of survival.
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1163:"Reciprocal affiliation among adolescent rats during a mild group stressor predicts mammary tumors and lifespan"
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Above all, the main criticism regarding studies concerning social grooming is that almost all of them focus on
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There are a variety of proposed mechanisms by which social grooming behavior has been hypothesized to increase
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social significance and function in the formation and maintenance of these friendships. Studies performed on
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Kimura R (1998). "Mutual grooming and preferred associate relationships in a band of free-ranging horses".
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Schino G (1 August 2001). "Grooming, competition and social rank among female primates: a meta-analysis".
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3270:"Trade-offs in primate grooming reciprocation: testing behavioral flexibility and correlated evolution"
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In insects, grooming often performs the important role of removing foreign material from the body. The
353:. It is often questioned why the behavior persists if it is costly to the one performing it; however,
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Social grooming may also serve to establish and recognize mates or amorous partners. For example, in
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typically describes the act of grooming between two individuals, often as a part of social grooming,
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van Lawick-Goodall J (1968). "The behavior of free living chimpanzees in the Gombe Stream
Reserve".
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van Lawick-Goodall J (1968). "The behavior of free living chimpanzees in the Gombe Stream
Reserve".
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2797:"Tool Use by Non-Human Primates | Center for Academic Research and Training in Anthropogeny (CARTA)"
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Spruijt BM, van Hooff JA, Gispen WH (July 1992). "Ethology and neurobiology of grooming behavior".
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Spruijt BM, van Hooff JA, Gispen WH (July 1992). "Ethology and neurobiology of grooming behavior".
1986:, Alberts SC, Altmann J (November 2003). "Social bonds of female baboons enhance infant survival".
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1889:"Interchange between grooming and infant handling in female Tibetan macaques (Macaca thibetana)"
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1388:"Whom to Groom and for What? Patterns of Grooming in Female Barbary Macaques (Macaca sylvanus)"
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129:. These evolutionary advantages may come in the form of health benefits including reduction in
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251:, social grooming has been shown to carry the role of maintaining relationships that increase
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495:
Male and female members of a species may differ in learning how, when, and whom to groom. In
387:
366:
184:
4632:"Genetic identification of C fibres that detect massage-like stroking of hairy skin in vivo"
1938:"Social preferences influence the short-term exchange of social grooming among male bonobos"
4643:
3888:
3627:
3425:
3085:
2762:
2409:
2230:
1995:
1841:
1646:
1458:
1399:
496:
462:
350:
341:
in the biological sense refers to a behavior performed by an individual that increases the
263:
4682:
2933:
1386:
Roubová, Veronika; Konečná, Martina; Šmilauer, Petr; Wallner, Bernard (10 February 2015).
8:
4280:
3353:
de Waal, F. (1989). Peacemaking among primates. Cambridge, MA: Harvard
University Press.
3023:
941:
772:
743:
508:
163:
98:
4647:
3892:
3631:
3618:
Uvnäs-Moberg K (January 1997). "Physiological and endocrine effects of social contact".
3429:
3089:
2766:
2413:
2325:
2234:
1999:
1845:
1650:
1462:
1403:
1298:
742:
behavior, whereby the primates engage in establishing and maintaining alliances through
312:
The relationship between median survival time and social connectedness in female baboons
4719:
4664:
4631:
4539:
4502:
4462:
4412:
4362:
4325:
4266:
4215:
4158:
4125:
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3998:
3816:
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3639:
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3554:
3448:
3413:
3389:
3364:
3336:
3164:
3116:
3073:
2970:
2866:
2778:
2731:
2698:
Weinberg, Susan M.; Candland, Douglas K. (1981). ""Stone-grooming" in Macaca fuscata".
2612:
2497:
2382:
2251:
2218:
2199:
2164:
2129:
2066:
2041:
2019:
1965:
1913:
1904:
1888:
1864:
1829:
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1187:
1162:
1116:
1077:
1025:
978:
963:
958:
731:
338:
94:
4715:
4437:
4395:
Lee PC (1987). "Sibships: Cooperation and
Competition Among Immature Vervet Monkeys".
4387:
3588:
3004:
2966:
4779:
4774:
4669:
4618:
4577:
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3990:
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2256:
2121:
2071:
2011:
1957:
1937:
1918:
1869:
1814:
1802:
1731:
1713:
1674:
1635:"Precise time-matching in chimpanzee allogrooming does not occur after a short delay"
1598:
1579:
1561:
1510:
1486:
1427:
1302:
1247:
1211:
1192:
1141:
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1069:
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concentrations. An increase in the number of receptors might influence the amount of
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252:
224:
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126:
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4366:
4270:
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3168:
2974:
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2501:
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2133:
2023:
1969:
1770:
1341:
1271:
1120:
1081:
1029:
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4711:
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4523:
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4404:
4383:
4346:
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4250:
4191:
4153:
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4137:
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3982:
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3800:
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3681:
3635:
3584:
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3520:
3516:
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3433:
3384:
3376:
3328:
3281:
3190:
3148:
3111:
3093:
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3000:
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2901:
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2782:
2770:
2707:
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2417:
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2339:
2331:
2246:
2238:
2191:
2156:
2113:
2105:
2061:
2053:
2003:
1949:
1908:
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1859:
1849:
1794:
1758:
1721:
1705:
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1610:
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1522:
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1417:
1407:
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1223:
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1009:
719:
512:
4604:
4490:
4321:
3986:
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3332:
2897:
2608:
2195:
2168:
2160:
1762:
1692:
Martin-Ordas, Gema; Haun, Daniel; Colmenares, Fernando; Call, Josep (March 2010).
4552:
4474:
3970:
3438:
3098:
1887:
Jiang, Qi; Xia, Dong-Po; Wang, Xi; Zhang, Dao; Sun, Binghua; Li, Jin-Hua (2019).
1854:
1659:
1471:
1412:
1178:
1000:
Henazi SP, Barrett L (January 1999). "The value of grooming to female primates".
863:
466:
358:
323:
277:
209:
1285:
Seyfarth RM, Cheney DL (5 July 2011). "The evolutionary origins of friendship".
3881:
Proceedings of the National Academy of Sciences of the United States of America
3537:
Kemp AH, Guastella AJ (8 August 2011). "The Role of Oxytocin in Human Affect".
921:
875:
660:
642:
565:
487:
individual is a fully developed adult and can follow normal grooming patterns.
416:
370:
354:
256:
193:
4067:
Platt, Michael L.; Seyfarth, Robert M.; Cheney, Dorothy L. (5 February 2016).
1953:
1709:
1615:
1526:
832:
663:
purposes in the exchange of resources such as food, sex, and communal hygiene.
4768:
4203:
4149:
4092:
4037:
3777:
3550:
3295:
3107:
3050:
2719:
2674:
2571:
2534:
2493:
2378:
2293:
2125:
1717:
1633:
Phelps, Steve; Ng, Wing; Musolesi, Mirco; Russell, Yvan (11 September 2018).
1565:
880:
764:
651:
458:
454:
293:
235:
215:
205:
82:
3380:
2443:
2421:
2007:
968:
28:
4673:
4535:
4527:
4262:
4254:
4167:
4110:
4084:
4045:
4029:
3901:
3861:
3852:
3835:
3693:
3457:
3398:
3160:
3152:
3125:
2727:
2711:
2335:
2242:
2219:"Infanticide risk and the evolution of male-female association in primates"
2075:
2057:
2015:
1961:
1922:
1873:
1806:
1735:
1678:
1583:
1490:
1431:
1333:
1306:
1196:
1073:
1065:
1021:
953:
924:, lactating females with 4- to 10-week-old infants were given low doses of
791:
690:
439:
429:
318:
110:
33:
4622:
4498:
4358:
4211:
3994:
3956:
3920:
3812:
3736:
3647:
3596:
3194:
2915:
2429:
2260:
2109:
170:
1097:"Grooming site preferences in primates and their functional implications"
849:
739:
656:
228:
89:, clean or maintain one another's bodies or appearances. A related term,
4739:
4655:
1694:"Keeping track of time: evidence for episodic-like memory in great apes"
1557:
4458:
4408:
2862:
1263:
1112:
1013:
589:
446:
435:
159:
4073:
Philosophical Transactions of the Royal Society B: Biological Sciences
3570:
3568:
2660:
1798:
1542:"Evolutionary ecology of chemosensation and its role in sensory drive"
1227:
4613:
3479:
2906:
2774:
2683:
2400:
Axelrod R, Hamilton WD (March 1981). "The evolution of cooperation".
2117:
1983:
888:
807:
795:
679:
666:
481:
301:
4630:
Vrontou S, Wong AM, Rau KK, Koerber HR, Anderson DJ (January 2013).
2469:
2313:(Fall 2013 ed.). Metaphysics Research Lab, Stanford University.
1508:
4513:
3565:
3138:
2485:
2370:
2357:
Trivers RL (1 March 1971). "The Evolution of Reciprocal Altruism".
2091:"The evolution of female social relationships in nonhuman primates"
925:
828:
346:
151:
114:
54:
3219:"Primate Sociality and Social Systems | Learn Science at Scitable"
2040:
Archie EA, Tung J, Clark M, Altmann J, Alberts SC (October 2014).
4123:
937:
549:
248:
155:
4572:
Smuts B, Cheney D, Seyfarth R, Wrangham R, Struhsaker T (1987).
1161:
Yee JR, Cavigelli SA, Delgado B, McClintock MK (November 2008).
4753:
2656:
2654:
1357:"Primate brains react differently to faces of friends and VIPs"
916:
698:
prevents ongoing infection while also promoting group success.
19:
This article is about the social activity. For other uses, see
4126:"Social Laughter Triggers Endogenous Opioid Release in Humans"
2181:
1691:
308:
200:
scans of the monkeys' brains lit up more significantly at the
3506:
3016:
3014:
1385:
1160:
86:
4477:(September 1997). "The importance of a well-groomed child".
3973:(September 1997). "The importance of a well-groomed child".
3875:
Capitanio JP, Mendoza SP, Lerche NW, Mason WA (April 1998).
3414:"Do ravens show consolation? Responses to distressed others"
2651:
330:
proven to decrease risk of infanticide in several primates.
4571:
4444:
4180:
3874:
3049:
2848:
612:
534:
242:
175:
3011:
361:
as the mechanism behind the clear advantages of altruism.
4236:
3933:
1045:
3074:"Social Grooming in Bats: Are Vampire Bats Exceptional?"
1936:
Surbeck, Martin; Hohmann, Gottfried (18 December 2014).
1830:"Social Grooming in Bats: Are Vampire Bats Exceptional?"
1447:"Social Grooming in Bats: Are Vampire Bats Exceptional?"
1212:"Functional Significance of Social Grooming in Primates"
4069:"Adaptations for social cognition in the primate brain"
3314:"Group size, grooming, and social cohesion in primates"
1632:
806:
Horses engage in mutual grooming via the formation of "
171:
Reinforcing social structure and building relationships
4590:
2986:
2984:
2883:
2039:
1748:
4629:
3268:
Schino, Gabriele; Aureli, Filippo (1 November 2008).
2749:
Mcgrew, William; TUTIN, CAROLINE (16 February 1973).
2594:
390:
219:
Indian short-nosed fruit bats in a tight-knit cluster
4336:
3790:
3714:
2327:
The Descent of Man, and Selection in Relation to Sex
1597:
Stopka, Pavel; Stopková, Romana (1 September 2001).
1140:. University of California Press. pp. 193–224.
490:
453:
males were more attentive to social grooming during
4066:
3833:
2981:
1381:
1379:
1377:
4423:
4279:
3834:Sapolsky RM, Romero LM, Munck AU (February 2000).
3362:
3022:
2990:
2216:
2146:
2089:Sterck EH, Watts DP, Schaik CP (1 November 1997).
1982:
1628:
1626:
1599:"Conditional allogrooming in the herb-field mouse"
482:General learning and reciprocation of allogrooming
405:
3667:
3665:
3072:Carter, Gerald; Leffer, Lauren (7 October 2015).
2088:
1886:
883:grooming resulted in a proportionate increase in
204:(associated with recognition and memory) and the
187:family, with the center pair grooming one another
4766:
3363:Kenny E, Birkhead TR, Green JP (1 August 2017).
2927:
2925:
2697:
1374:
1241:
1239:
1237:
2822:"Primate Cognition | Learn Science at Scitable"
2399:
1783:
1623:
931:
3676:. Touch, Temperature, Pain/Itch and Pleasure.
3662:
3307:
3305:
2035:
2033:
1935:
1596:
1540:Yohe, Laurel R; Brand, Philipp (August 2018).
1511:"Social grooming among short-nosed fruit bats"
1504:
1502:
1500:
1284:
476:
287:
3536:
3411:
3244:"Social Grooming in Primates: Adaptive Value"
2922:
2470:"Coefficients of Inbreeding and Relationship"
1234:
999:
4683:"Social grooming in the common vampire bat,
4303:"Payment for sex in a macaque mating market"
3617:
3574:
3267:
3180:
3178:
3071:
2934:"Social grooming in the common vampire bat,
2844:
2842:
2217:van Schaik CP, Kappeler PM (November 1997).
1827:
1444:
1041:
1039:
870:
158:, social grooming has been proven to reduce
4286:. Cambridge, MA: Harvard University Press.
4015:
3708:
3539:Current Directions in Psychological Science
3302:
3029:. Cambridge, MA: Harvard University Press.
2877:
2748:
2628:
2626:
2307:Okasha S (1 January 2013). Zalta EN (ed.).
2276:Selection : the mechanism of evolution
2030:
1497:
3620:Annals of the New York Academy of Sciences
3132:
1133:
120:
4705:
4680:
4663:
4612:
4157:
4100:
3910:
3900:
3851:
3447:
3437:
3388:
3285:
3274:Biological Journal of the Linnean Society
3175:
3115:
3097:
2956:
2931:
2905:
2839:
2682:
2250:
2065:
1912:
1863:
1853:
1725:
1668:
1658:
1614:
1573:
1539:
1480:
1470:
1421:
1411:
1186:
1036:
270:
4576:. Chicago: University of Chicago Press.
4473:
3969:
3757:
2751:"Chimpanzee Tool Use in Dental Grooming"
2623:
995:
993:
705:
689:
665:
307:
243:Enforcing hierarchy and social structure
214:
179:
176:Creation and maintenance of social bonds
60:
39:
27:
4277:
3311:
3061:. Chicago: University of Chicago Press.
3020:
2634:"Social Grooming in Primates: Ontogeny"
2356:
2310:The Stanford Encyclopedia of Philosophy
369:model, and out of this comparison came
4767:
4373:
4300:
3674:Neuroscience and Biobehavioral Reviews
3671:
2467:
2323:
2306:
2273:
1319:
1245:
1209:
1094:
4550:
3532:
3530:
3184:
2590:
2588:
2550:
2546:
2544:
2514:
1828:Carter G, Leffer L (7 October 2015).
1445:Carter G, Leffer L (7 October 2015).
990:
838:dorsal column-medial lemniscal system
422:
2581:on 4 October 2011 – via Brill.
1354:
1252:International Journal of Primatology
1134:Aureli F, Waal FB (1 January 2000).
1101:International Journal of Primatology
822:
4394:
3312:Lehmann, Johannes (December 2007).
2098:Behavioral Ecology and Sociobiology
1299:10.1146/annurev-psych-120710-100337
794:, which researchers believe builds
786:Preening (bird) § Allopreening
13:
4229:
3640:10.1111/j.1749-6632.1997.tb51917.x
3527:
2585:
2541:
1905:10.24272/j.issn.2095-8137.2018.049
856:
519:
380:
136:
74:A male cat grooms a female kitten.
14:
4801:
4732:
4557:. New York: Aldine Publications.
3412:Fraser ON, Bugnyar T (May 2010).
1246:Boccia, Maria (1 December 1983).
1095:Barton, Robert (1 October 1985).
645:allogrooming while infant suckles
491:Sex-based differences in learning
4752:
4738:
4376:Applied Animal Behaviour Science
3509:Applied Animal Behaviour Science
3482:Applied Animal Behaviour Science
3287:10.1111/j.1095-8312.2008.01067.x
2223:Proceedings. Biological Sciences
2046:Proceedings. Biological Sciences
1210:Dunbar, Robin (1 January 1991).
1002:Primates; Journal of Primatology
635:
620:
597:
572:
557:
541:
526:
4516:American Journal of Primatology
4243:American Journal of Primatology
4174:
4117:
4060:
4018:American Journal of Primatology
4009:
3963:
3927:
3868:
3827:
3784:
3751:
3686:10.1016/j.neubiorev.2008.07.001
3611:
3500:
3473:
3464:
3405:
3356:
3347:
3261:
3236:
3211:
3141:American Journal of Primatology
3065:
3043:
2814:
2789:
2742:
2700:American Journal of Primatology
2691:
2508:
2461:
2436:
2393:
2359:The Quarterly Review of Biology
2350:
2317:
2300:
2267:
2210:
2175:
2140:
2082:
1976:
1929:
1880:
1821:
1787:American Journal of Primatology
1777:
1742:
1685:
1590:
1533:
1438:
1348:
1053:American Journal of Primatology
4142:10.1523/JNEUROSCI.0688-16.2017
3758:Hemmings HC (1 January 2013).
3521:10.1016/j.applanim.2010.12.003
3494:10.1016/j.applanim.2008.08.005
3189:. Taylor & Francis Group.
1313:
1278:
1203:
1154:
1127:
1088:
1:
4743:The dictionary definition of
4716:10.1016/S0003-3472(86)80274-3
4605:10.1152/physrev.1992.72.3.825
4554:Sex and Friendship in Baboons
4491:10.1126/science.277.5332.1620
4438:10.1016/S0066-1856(68)80003-2
4388:10.1016/S0168-1591(97)00129-9
4322:10.1016/j.anbehav.2007.03.009
3987:10.1126/science.277.5332.1620
3949:10.1126/science.277.5332.1659
3589:10.1016/S0306-4530(98)00056-0
3333:10.1016/j.anbehav.2006.10.025
3187:Sex and Friendship in Baboons
3005:10.1016/s0066-1856(68)80003-2
2967:10.1016/S0003-3472(86)80274-3
2898:10.1152/physrev.1992.72.3.825
2609:10.1016/j.anbehav.2011.07.012
2196:10.1016/j.anbehav.2011.02.009
2161:10.1016/j.anbehav.2003.07.001
1763:10.1016/j.anbehav.2009.10.014
984:
726:) grooming each other at the
502:
4351:10.1016/0306-4530(89)90065-6
4196:10.1016/0306-4530(93)90027-I
3805:10.1016/0306-4530(89)90065-6
3729:10.1016/0306-4530(89)90065-6
3439:10.1371/journal.pone.0010605
3099:10.1371/journal.pone.0138430
2344:2027/bc.ark:/13960/t96696g2b
1855:10.1371/journal.pone.0138430
1660:10.1371/journal.pone.0201810
1472:10.1371/journal.pone.0138430
1413:10.1371/journal.pone.0117298
1179:10.1097/psy.0b013e31818425fb
932:Criticism for studies quoted
7:
4426:Animal Behaviour Monographs
4301:Gumert MD (December 2007).
4130:The Journal of Neuroscience
2993:Animal Behaviour Monographs
2519:. Oxford University Press.
2515:Smith JM (1 January 1996).
2278:. Oxford University Press.
2274:Graham B (1 January 2008).
1287:Annual Review of Psychology
1137:Natural Conflict Resolution
947:
893:thyroid-stimulating hormone
843:
701:
477:Ontogeny of social grooming
333:
288:Direct fitness consequences
10:
4808:
4447:Journal of Insect Behavior
4282:Peacemaking among Primates
3025:Peacemaking among primates
2851:Journal of Insect Behavior
910:
783:
673:
18:
16:Behavior in social animals
1954:10.1007/s10071-014-0826-0
1710:10.1007/s10071-009-0282-4
1527:10.1163/1568539X-00003410
871:Glucocorticoids receptors
813:
801:
592:preening the male (video)
101:, maternal behavior, and
4339:Psychoneuroendocrinology
4184:Psychoneuroendocrinology
3793:Psychoneuroendocrinology
3717:Psychoneuroendocrinology
3577:Psychoneuroendocrinology
3551:10.1177/0963721411417547
3053:, Cheney D, Seyfarth R,
2675:10.1163/000579510X505436
2572:10.1163/156853909x406455
2551:Frank R (January 2009).
2468:Wright S (1 July 1922).
885:glucocorticoid receptors
779:
3057:, Struhsaker T (1987).
2474:The American Naturalist
2422:10.1126/science.7466396
2008:10.1126/science.1088580
1616:10.1093/beheco/12.5.584
685:
406:{\displaystyle rB>C}
121:Evolutionary advantages
81:is a behavior in which
4528:10.1002/ajp.1350160106
4255:10.1002/ajp.1350190105
4085:10.1098/rstb.2015.0096
4030:10.1002/ajp.1350280402
3902:10.1073/pnas.95.8.4714
3853:10.1210/edrv.21.1.0389
3153:10.1002/ajp.1350160106
2801:carta.anthropogeny.org
2712:10.1002/ajp.1350010412
2336:10.5962/bhl.title.2092
2243:10.1098/rspb.1997.0234
2058:10.1098/rspb.2014.1261
1334:10.1006/anbe.2001.1750
1167:Psychosomatic Medicine
1066:10.1002/ajp.1350190105
734:
728:Jigokudani Monkey Park
694:
670:
407:
313:
271:Interchange of favours
225:short-nosed fruit bats
220:
188:
75:
58:
37:
21:Groom (disambiguation)
4681:Wilkinson GS (1986).
4593:Physiological Reviews
3381:10.1093/beheco/arx078
3195:10.4324/9781315129204
2932:Wilkinson GS (1986).
2886:Physiological Reviews
2517:Evolutionary genetics
2448:www.classicreader.com
2110:10.1007/s002650050390
744:dominance hierarchies
718:
693:
669:
629:yellow-billed babbler
497:stump-tailed macaques
408:
311:
264:black crested gibbons
262:Grooming networks in
218:
185:North American beaver
183:
164:Observational studies
73:
52:
31:
4761:at Wikimedia Commons
773:crab-eating macaques
657:consolatory behavior
552:grooming one another
463:reproductive fitness
388:
351:reproductive fitness
131:disease transmission
4790:Mutualism (biology)
4656:10.1038/nature11810
4648:2013Natur.493..669V
4239:Macaca fascicularis
3893:1998PNAS...95.4714C
3632:1997NYASA.807..146U
3430:2010PLoSO...510605F
3090:2015PLoSO..1038430C
2767:1973Natur.241..477M
2414:1981Sci...211.1390A
2235:1997RSPSB.264.1687V
2000:2003Sci...302.1231S
1893:Zoological Research
1846:2015PLoSO..1038430C
1651:2018PLoSO..1301810P
1463:2015PLoSO..1038430C
1404:2015PLoSO..1017298R
1216:Folia Primatologica
1048:Macaca fascicularis
564:Social grooming in
533:Mutual grooming in
447:white-handed gibbon
317:obtaining aid from
99:conflict resolution
4459:10.1007/BF02009512
4409:10.1007/BF02382182
4278:de Waal F (1989).
4079:(1687): 20150096.
3369:Behavioral Ecology
3021:de Waal F (1989).
2863:10.1007/BF02009512
2052:(1793): 20141261.
1603:Behavioral Ecology
1264:10.1007/BF02735602
1113:10.1007/BF02735574
1014:10.1007/BF02557701
979:Group cohesiveness
964:Community cohesion
959:Cleaning symbiosis
771:Recent studies of
735:
695:
671:
436:reduced aggression
423:Use as a commodity
403:
367:Prisoner's Dilemma
314:
298:Papio cynocephalus
221:
202:perirhinial cortex
189:
115:precoital activity
76:
59:
38:
36:grooms a juvenile.
4757:Media related to
4685:Desmodus rotundus
4574:Primate Societies
4564:978-0-202-02027-3
4136:(25): 6125–6131.
3943:(5332): 1659–62.
3840:Endocrine Reviews
3204:978-1-315-12920-4
3185:Smuts BB (1985).
3059:Primate Societies
2936:Desmodus rotundus
2761:(5390): 477–478.
2324:Darwin C (1871).
2229:(1388): 1687–94.
1799:10.1002/ajp.22178
1558:10.1093/cz/zoy048
1355:Woodward, Aylin.
1228:10.1159/000156574
974:Personal grooming
897:negative feedback
823:Endocrine effects
720:Japanese macaques
716:
606:
583:
513:Japanese macaques
471:courting behavior
375:inclusive fitness
105:in some species.
71:
57:groom a juvenile.
50:
4797:
4756:
4742:
4727:
4709:
4700:(6): 1880–1889.
4694:Animal Behaviour
4691:
4677:
4667:
4642:(7434): 669–73.
4626:
4616:
4587:
4568:
4551:Smuts B (1985).
4547:
4510:
4485:(5332): 1620–1.
4470:
4441:
4420:
4391:
4370:
4333:
4316:(6): 1655–1667.
4310:Animal Behaviour
4307:
4297:
4285:
4274:
4224:
4223:
4178:
4172:
4171:
4161:
4121:
4115:
4114:
4104:
4064:
4058:
4057:
4013:
4007:
4006:
3981:(5332): 1620–1.
3967:
3961:
3960:
3931:
3925:
3924:
3914:
3904:
3872:
3866:
3865:
3855:
3831:
3825:
3824:
3788:
3782:
3781:
3755:
3749:
3748:
3712:
3706:
3705:
3669:
3660:
3659:
3615:
3609:
3608:
3572:
3563:
3562:
3534:
3525:
3524:
3504:
3498:
3497:
3488:(2–4): 141–149.
3477:
3471:
3468:
3462:
3461:
3451:
3441:
3409:
3403:
3402:
3392:
3375:(4): 1142–1148.
3360:
3354:
3351:
3345:
3344:
3327:(6): 1617–1629.
3321:Animal Behaviour
3318:
3309:
3300:
3299:
3289:
3265:
3259:
3258:
3256:
3254:
3240:
3234:
3233:
3231:
3229:
3215:
3209:
3208:
3182:
3173:
3172:
3136:
3130:
3129:
3119:
3101:
3084:(10): e0138430.
3069:
3063:
3062:
3047:
3041:
3040:
3028:
3018:
3009:
3008:
2988:
2979:
2978:
2960:
2951:(6): 1880–1889.
2945:Animal Behaviour
2942:
2929:
2920:
2919:
2909:
2881:
2875:
2874:
2846:
2837:
2836:
2834:
2832:
2818:
2812:
2811:
2809:
2807:
2793:
2787:
2786:
2775:10.1038/241477a0
2746:
2740:
2739:
2695:
2689:
2688:
2686:
2669:(9): 1073–1099.
2658:
2649:
2648:
2646:
2644:
2630:
2621:
2620:
2597:Animal Behaviour
2592:
2583:
2582:
2580:
2574:. Archived from
2566:(8): 1123–1135.
2557:
2548:
2539:
2538:
2512:
2506:
2505:
2480:(645): 330–338.
2465:
2459:
2458:
2456:
2454:
2440:
2434:
2433:
2408:(4489): 1390–6.
2397:
2391:
2390:
2354:
2348:
2347:
2321:
2315:
2314:
2304:
2298:
2297:
2271:
2265:
2264:
2254:
2214:
2208:
2207:
2190:(6): 1109–1116.
2184:Animal Behaviour
2179:
2173:
2172:
2149:Animal Behaviour
2144:
2138:
2137:
2095:
2086:
2080:
2079:
2069:
2037:
2028:
2027:
1994:(5648): 1231–4.
1980:
1974:
1973:
1942:Animal Cognition
1933:
1927:
1926:
1916:
1884:
1878:
1877:
1867:
1857:
1840:(10): e0138430.
1825:
1819:
1818:
1781:
1775:
1774:
1751:Animal Behaviour
1746:
1740:
1739:
1729:
1698:Animal Cognition
1689:
1683:
1682:
1672:
1662:
1630:
1621:
1620:
1618:
1594:
1588:
1587:
1577:
1537:
1531:
1530:
1506:
1495:
1494:
1484:
1474:
1457:(10): e0138430.
1442:
1436:
1435:
1425:
1415:
1383:
1372:
1371:
1369:
1367:
1352:
1346:
1345:
1322:Animal Behaviour
1317:
1311:
1310:
1282:
1276:
1275:
1243:
1232:
1231:
1207:
1201:
1200:
1190:
1158:
1152:
1151:
1131:
1125:
1124:
1092:
1086:
1085:
1043:
1034:
1033:
997:
790:Birds engage in
717:
639:
627:Allopreening in
624:
615:grooming another
608:
607:
585:
584:
561:
545:
530:
412:
410:
409:
404:
324:Barbary macaques
278:Tibetan macaques
210:Barbary macaques
72:
51:
4807:
4806:
4800:
4799:
4798:
4796:
4795:
4794:
4765:
4764:
4759:Social grooming
4746:social grooming
4735:
4730:
4707:10.1.1.539.5104
4689:
4584:
4565:
4345:(1–2): 155–61.
4305:
4294:
4232:
4230:Further reading
4227:
4179:
4175:
4122:
4118:
4065:
4061:
4014:
4010:
3968:
3964:
3932:
3928:
3873:
3869:
3832:
3828:
3799:(1–2): 155–61.
3789:
3785:
3770:
3756:
3752:
3723:(1–2): 155–61.
3713:
3709:
3670:
3663:
3616:
3612:
3573:
3566:
3535:
3528:
3505:
3501:
3478:
3474:
3469:
3465:
3410:
3406:
3361:
3357:
3352:
3348:
3316:
3310:
3303:
3266:
3262:
3252:
3250:
3242:
3241:
3237:
3227:
3225:
3217:
3216:
3212:
3205:
3183:
3176:
3137:
3133:
3070:
3066:
3048:
3044:
3037:
3019:
3012:
2989:
2982:
2958:10.1.1.539.5104
2940:
2930:
2923:
2882:
2878:
2847:
2840:
2830:
2828:
2820:
2819:
2815:
2805:
2803:
2795:
2794:
2790:
2747:
2743:
2696:
2692:
2659:
2652:
2642:
2640:
2632:
2631:
2624:
2593:
2586:
2578:
2555:
2549:
2542:
2527:
2513:
2509:
2466:
2462:
2452:
2450:
2442:
2441:
2437:
2398:
2394:
2355:
2351:
2330:. John Murray.
2322:
2318:
2305:
2301:
2286:
2272:
2268:
2215:
2211:
2180:
2176:
2145:
2141:
2093:
2087:
2083:
2038:
2031:
1981:
1977:
1934:
1930:
1885:
1881:
1826:
1822:
1793:(12): 1165–73.
1782:
1778:
1747:
1743:
1690:
1686:
1645:(9): e0201810.
1631:
1624:
1595:
1591:
1546:Current Zoology
1538:
1534:
1507:
1498:
1443:
1439:
1398:(2): e0117298.
1384:
1375:
1365:
1363:
1353:
1349:
1318:
1314:
1283:
1279:
1244:
1235:
1208:
1204:
1159:
1155:
1148:
1132:
1128:
1093:
1089:
1044:
1037:
998:
991:
987:
950:
934:
913:
876:Glucocorticoids
873:
864:opiate receptor
859:
857:Beta-endorphins
846:
833:beta-endorphins
825:
816:
804:
788:
782:
706:
704:
688:
676:
646:
640:
631:
625:
616:
609:
598:
593:
586:
573:
568:
566:hyacinth macaws
562:
553:
546:
537:
531:
522:
520:Mutual grooming
505:
493:
484:
479:
469:, similar to a
425:
389:
386:
385:
383:
381:Hamilton's rule
359:group selection
336:
290:
273:
245:
194:rhesus macaques
178:
173:
139:
137:Health benefits
123:
107:Mutual grooming
79:Social grooming
61:
40:
24:
17:
12:
11:
5:
4805:
4804:
4793:
4792:
4787:
4782:
4777:
4763:
4762:
4750:
4734:
4733:External links
4731:
4729:
4728:
4678:
4627:
4588:
4582:
4569:
4563:
4548:
4511:
4471:
4453:(6): 855–861.
4442:
4421:
4392:
4382:(4): 265–276.
4371:
4334:
4298:
4292:
4275:
4233:
4231:
4228:
4226:
4225:
4190:(4): 307–321.
4173:
4116:
4059:
4024:(4): 231–250.
4008:
3962:
3926:
3867:
3826:
3783:
3769:978-1437716795
3768:
3750:
3707:
3661:
3610:
3564:
3545:(4): 222–231.
3526:
3515:(3–4): 81–90.
3499:
3472:
3463:
3404:
3355:
3346:
3301:
3280:(3): 439–446.
3260:
3235:
3223:www.nature.com
3210:
3203:
3174:
3131:
3064:
3042:
3035:
3010:
2980:
2921:
2876:
2857:(6): 855–861.
2838:
2826:www.nature.com
2813:
2788:
2741:
2706:(4): 465–468.
2690:
2650:
2622:
2603:(4): 801–809.
2584:
2540:
2526:978-0198542155
2525:
2507:
2486:10.1086/279872
2460:
2435:
2392:
2371:10.1086/406755
2349:
2316:
2299:
2285:978-0198569725
2284:
2266:
2209:
2174:
2155:(3): 573–582.
2139:
2104:(5): 291–309.
2081:
2029:
1975:
1948:(2): 573–579.
1928:
1899:(2): 139–145.
1879:
1820:
1776:
1757:(2): 271–279.
1741:
1704:(2): 331–340.
1684:
1622:
1609:(5): 584–589.
1589:
1552:(4): 525–533.
1532:
1496:
1437:
1373:
1347:
1328:(2): 265–271.
1312:
1277:
1258:(4): 399–418.
1233:
1222:(3): 121–131.
1202:
1153:
1146:
1126:
1107:(5): 519–532.
1087:
1035:
988:
986:
983:
982:
981:
976:
971:
966:
961:
956:
949:
946:
933:
930:
922:rhesus monkeys
912:
909:
872:
869:
858:
855:
845:
842:
824:
821:
815:
812:
803:
800:
784:Main article:
781:
778:
724:Macaca fuscata
703:
700:
687:
684:
675:
672:
648:
647:
643:Bonnet macaque
641:
634:
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619:
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525:
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492:
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483:
480:
478:
475:
461:increases the
451:Hylobates lar)
424:
421:
402:
399:
396:
393:
382:
379:
371:Robert Trivers
355:Charles Darwin
335:
332:
294:yellow baboons
289:
286:
272:
269:
257:rhesus monkeys
244:
241:
177:
174:
172:
169:
138:
135:
122:
119:
103:reconciliation
83:social animals
15:
9:
6:
4:
3:
2:
4803:
4802:
4791:
4788:
4786:
4783:
4781:
4778:
4776:
4773:
4772:
4770:
4760:
4755:
4751:
4749:at Wiktionary
4748:
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4736:
4725:
4721:
4717:
4713:
4708:
4703:
4699:
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4679:
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4649:
4645:
4641:
4637:
4633:
4628:
4624:
4620:
4615:
4610:
4606:
4602:
4599:(3): 825–52.
4598:
4594:
4589:
4585:
4583:9780226767161
4579:
4575:
4570:
4566:
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3887:(8): 4714–9.
3886:
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3625:
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3068:
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3052:
3046:
3038:
3036:9780674659209
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2946:
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2937:
2928:
2926:
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2899:
2895:
2892:(3): 825–52.
2891:
2887:
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2047:
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2021:
2017:
2013:
2009:
2005:
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1997:
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1963:
1959:
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1932:
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955:
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945:
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942:observational
939:
929:
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765:Vervet monkey
760:
756:
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523:
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514:
510:
500:
498:
488:
474:
472:
468:
467:ovarian cycle
464:
460:
459:fertilization
456:
452:
448:
445:In addition,
443:
441:
440:social groups
437:
433:
431:
420:
418:
417:W.D. Hamilton
415:Developed by
413:
400:
397:
394:
391:
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376:
372:
368:
362:
360:
356:
352:
348:
344:
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310:
306:
303:
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295:
285:
281:
279:
268:
265:
260:
258:
254:
250:
240:
237:
236:kin selection
232:
230:
226:
217:
213:
211:
207:
206:temporal pole
203:
199:
195:
186:
182:
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165:
161:
157:
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147:
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128:
118:
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4522:(1): 43–50.
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4478:
4450:
4446:
4429:
4425:
4403:(1): 47–59.
4400:
4396:
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4249:(1): 39–51.
4246:
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3880:
3870:
3846:(1): 55–89.
3843:
3839:
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3796:
3792:
3786:
3762:. Elsevier.
3759:
3753:
3720:
3716:
3710:
3680:(2): 260–8.
3677:
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3502:
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3417:
3407:
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3320:
3277:
3273:
3263:
3251:. Retrieved
3248:www.reed.edu
3247:
3238:
3226:. Retrieved
3222:
3213:
3186:
3147:(1): 43–50.
3144:
3140:
3134:
3081:
3077:
3067:
3058:
3045:
3024:
2996:
2992:
2948:
2944:
2935:
2889:
2885:
2879:
2854:
2850:
2829:. Retrieved
2825:
2816:
2804:. Retrieved
2800:
2791:
2758:
2754:
2744:
2703:
2699:
2693:
2666:
2662:
2641:. Retrieved
2638:www.reed.edu
2637:
2600:
2596:
2576:the original
2563:
2559:
2516:
2510:
2477:
2473:
2463:
2451:. Retrieved
2447:
2438:
2405:
2401:
2395:
2365:(1): 35–57.
2362:
2358:
2352:
2326:
2319:
2309:
2302:
2275:
2269:
2226:
2222:
2212:
2187:
2183:
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2152:
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2101:
2097:
2084:
2049:
2045:
1991:
1987:
1978:
1945:
1941:
1931:
1896:
1892:
1882:
1837:
1833:
1823:
1790:
1786:
1779:
1754:
1750:
1744:
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1592:
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1545:
1535:
1518:
1514:
1454:
1450:
1440:
1395:
1391:
1364:. Retrieved
1360:
1350:
1325:
1321:
1315:
1290:
1286:
1280:
1255:
1251:
1219:
1215:
1205:
1170:
1166:
1156:
1136:
1129:
1104:
1100:
1090:
1060:(1): 39–51.
1057:
1051:
1047:
1008:(1): 47–59.
1005:
1001:
954:Cleaner fish
935:
914:
905:
902:
874:
860:
850:vampire bats
847:
826:
817:
805:
792:allopreening
789:
770:
761:
757:
753:
749:
736:
723:
696:
677:
649:
506:
494:
485:
450:
444:
430:Papio anubis
428:
426:
414:
384:
363:
337:
326:
319:conspecifics
315:
297:
291:
282:
274:
261:
246:
233:
222:
196:showed that
190:
148:
144:autogrooming
140:
124:
111:pair bonding
106:
91:allogrooming
90:
85:, including
78:
77:
34:olive baboon
25:
4475:Sapolsky RM
4432:: 161–311.
3971:Sapolsky RM
3253:13 November
3228:13 November
2999:: 161–311.
915:Endogenous
763:adulthood.
740:consolation
661:utilitarian
659:as well as
229:mate choice
4769:Categories
3055:Wrangham R
2831:30 January
2806:30 January
2643:30 January
1366:30 January
1293:: 153–77.
985:References
808:pair bonds
796:pair bonds
732:Yamanouchi
590:budgerigar
509:tool usage
503:Tool usage
160:heart rate
55:red wolves
53:Two adult
4702:CiteSeerX
4614:1874/3750
4204:0306-4530
4150:0270-6474
4093:0962-8436
4054:196609839
4038:1098-2345
3778:830351627
3559:145122328
3296:0024-4066
3108:1932-6203
2953:CiteSeerX
2907:1874/3750
2720:1098-2345
2684:1893/2990
2663:Behaviour
2560:Behaviour
2535:954574132
2494:0003-0147
2379:0033-5770
2294:781154368
2126:0340-5443
2118:1874/1183
1815:205330388
1718:1435-9448
1566:1674-5507
1521:: 37–63.
1515:Behaviour
889:serotonin
680:honey bee
357:proposed
302:sociality
234:Finally,
152:stressors
32:An adult
4780:Ethology
4775:Cleaning
4724:11214563
4674:23364746
4544:86566868
4536:31968881
4507:45145411
4467:40236926
4417:21449948
4397:Primates
4367:24350695
4271:86673865
4263:31964021
4220:11262534
4168:28536272
4111:26729935
4046:31941207
4003:45145411
3862:10696570
3821:24350695
3745:24350695
3702:30450770
3694:18662717
3656:46171612
3605:24789741
3458:20485685
3418:PLOS ONE
3399:29622926
3341:14866172
3169:86566868
3161:31968881
3126:26445502
3078:PLOS ONE
2975:11214563
2871:40236926
2736:85138037
2728:31995914
2617:53190848
2502:83865141
2453:23 March
2387:19027999
2204:53144146
2134:25163826
2076:25209936
2024:24970809
2016:14615543
1970:16030429
1962:25519436
1923:29955029
1874:26445502
1834:PLOS ONE
1807:23843246
1771:53203763
1736:19784852
1679:30204753
1639:PLOS ONE
1584:30108633
1491:26445502
1451:PLOS ONE
1432:25668722
1392:PLOS ONE
1342:53145237
1307:21740224
1272:24431777
1197:18842748
1121:37114535
1082:86673865
1074:31964021
1030:34106634
1022:23179531
948:See also
938:primates
926:naloxone
881:maternal
844:Oxytocin
829:oxytocin
702:Primates
550:macaques
347:altruism
339:Altruism
334:Altruism
249:meerkats
156:macaques
4785:Hygiene
4665:3563425
4644:Bibcode
4623:1320764
4499:9312858
4479:Science
4359:2525263
4330:9104008
4212:8391149
4159:6596504
4102:4745018
3995:9312858
3975:Science
3957:9287218
3937:Science
3921:9539804
3889:Bibcode
3813:2525263
3737:2525263
3648:9071348
3628:Bibcode
3597:9924739
3449:2868892
3426:Bibcode
3390:5873249
3117:4596566
3086:Bibcode
3051:Smuts B
2916:1320764
2783:4166467
2763:Bibcode
2430:7466396
2410:Bibcode
2402:Science
2261:9404030
2252:1688726
2231:Bibcode
2067:4173677
1996:Bibcode
1988:Science
1984:Silk JB
1914:6378558
1865:4596566
1842:Bibcode
1727:2822233
1670:6133271
1647:Bibcode
1575:6084603
1482:4596566
1459:Bibcode
1423:4323340
1400:Bibcode
1188:5767077
969:Licking
917:opioids
911:Opioids
674:Insects
652:animals
588:Female
343:fitness
253:fitness
127:fitness
113:, or a
4722:
4704:
4672:
4662:
4636:Nature
4621:
4580:
4561:
4542:
4534:
4505:
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4465:
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4091:
4052:
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3811:
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3735:
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3646:
3603:
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3167:
3159:
3124:
3114:
3106:
3033:
2973:
2955:
2914:
2869:
2781:
2755:Nature
2734:
2726:
2718:
2615:
2533:
2523:
2500:
2492:
2428:
2385:
2377:
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2282:
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2249:
2202:
2169:783193
2167:
2132:
2124:
2074:
2064:
2022:
2014:
1968:
1960:
1921:
1911:
1872:
1862:
1813:
1805:
1769:
1734:
1724:
1716:
1677:
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1572:
1564:
1489:
1479:
1430:
1420:
1340:
1305:
1270:
1195:
1185:
1144:
1119:
1080:
1072:
1028:
1020:
814:Cattle
802:Horses
548:Three
535:ponies
455:estrus
87:humans
4720:S2CID
4690:(PDF)
4540:S2CID
4503:S2CID
4463:S2CID
4413:S2CID
4363:S2CID
4326:S2CID
4306:(PDF)
4267:S2CID
4216:S2CID
4050:S2CID
3999:S2CID
3912:22556
3817:S2CID
3741:S2CID
3698:S2CID
3652:S2CID
3601:S2CID
3555:S2CID
3337:S2CID
3317:(PDF)
3165:S2CID
2971:S2CID
2941:(PDF)
2867:S2CID
2779:S2CID
2732:S2CID
2613:S2CID
2579:(PDF)
2556:(PDF)
2498:S2CID
2383:S2CID
2200:S2CID
2165:S2CID
2130:S2CID
2094:(PDF)
2020:S2CID
1966:S2CID
1811:S2CID
1767:S2CID
1338:S2CID
1268:S2CID
1117:S2CID
1078:S2CID
1026:S2CID
780:Birds
650:Many
154:. In
4670:PMID
4619:PMID
4578:ISBN
4559:ISBN
4532:PMID
4495:PMID
4355:PMID
4288:ISBN
4259:PMID
4241:)".
4208:PMID
4200:ISSN
4164:PMID
4146:ISSN
4107:PMID
4089:ISSN
4042:PMID
4034:ISSN
3991:PMID
3953:PMID
3917:PMID
3858:PMID
3809:PMID
3774:OCLC
3764:ISBN
3733:PMID
3690:PMID
3644:PMID
3593:PMID
3454:PMID
3395:PMID
3292:ISSN
3255:2019
3230:2019
3199:ISBN
3157:PMID
3122:PMID
3104:ISSN
3031:ISBN
2912:PMID
2833:2020
2808:2020
2724:PMID
2716:ISSN
2645:2020
2531:OCLC
2521:ISBN
2490:ISSN
2455:2017
2426:PMID
2375:ISSN
2290:OCLC
2280:ISBN
2257:PMID
2122:ISSN
2072:PMID
2012:PMID
1958:PMID
1919:PMID
1870:PMID
1803:PMID
1732:PMID
1714:ISSN
1675:PMID
1580:PMID
1562:ISSN
1487:PMID
1428:PMID
1368:2020
1303:PMID
1193:PMID
1142:ISBN
1070:PMID
1050:)".
1018:PMID
891:and
686:Bats
613:lion
611:One
398:>
198:fMRI
95:bond
4712:doi
4660:PMC
4652:doi
4640:493
4609:hdl
4601:doi
4524:doi
4487:doi
4483:277
4455:doi
4434:doi
4405:doi
4384:doi
4347:doi
4318:doi
4251:doi
4192:doi
4154:PMC
4138:doi
4097:PMC
4081:doi
4077:371
4026:doi
3983:doi
3979:277
3945:doi
3941:277
3907:PMC
3897:doi
3848:doi
3801:doi
3725:doi
3682:doi
3636:doi
3624:807
3585:doi
3547:doi
3517:doi
3513:130
3490:doi
3486:116
3444:PMC
3434:doi
3385:PMC
3377:doi
3329:doi
3282:doi
3191:doi
3149:doi
3112:PMC
3094:doi
3001:doi
2963:doi
2902:hdl
2894:doi
2859:doi
2771:doi
2759:241
2708:doi
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