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Kudo, H., & Dunbar, R. I. M. (2001). Neocortex size and social network size in primates. Anim. Behav., 62(4), 711–722.
Abstract: Primates use social grooming to service coalitions and it has been suggested that these directly affect the fitness of their members by allowing them to reduce the intrinsic costs associated with living in large groups. We tested two hypotheses about the size of grooming cliques that derive from this suggestion: (1) that grooming clique size should correlate with relative neocortex size and (2) that the size of grooming cliques should be proportional to the size of the groups they have to support. Both predictions were confirmed, although we show that, in respect of neocortex size, there are as many as four statistically distinct grades within the primates (including humans). Analysis of the patterns of grooming among males and females suggested that large primate social groups often consist of a set of smaller female subgroups (in some cases, matrilinearly based coalitions) that are linked by individual males. This may be because males insert themselves into the interstices between weakly bonded female subgroups rather than because they actually hold these subunits together.
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Boesch, C. (1994). Cooperative hunting in wild chimpanzees. Anim. Behav., 48(3), 653–667.
Abstract: A model for the evolution of cooperation shows that two conditions are necessary for cooperation to be stable: a hunting success rate that is low for single hunters and increases with group size, and a social mechanism limiting access to meat by non-hunters. Testing this model on TaI chimpanzees, Pan troglodytes, showed that (1) it pays for individuals to hunt in groups of three or four rather than alone or in pairs, and (2) cooperation is stable because hunters gain more at these group sizes than cheaters, owing to a meat-sharing pattern in which hunting, dominance and age, in that order, determine how much an individual gets. In addition, hunters provide cheaters (about 45% of the meat eaters) with the surplus they produce during the hunts. Thus, cooperation in Tai male chimpanzees is an evolutionarily stable strategy, and its success allows cheating to be an evolutionarily stable strategy for Tai female chimpanzees. In Gombe chimpanzees, cooperation is not stable, first, because hunting success is very high for single hunters, and second, because no social mechanism exists that limits access to meat by non-hunters. The analysis showed that some assumptions made when discussing cooperation in other social hunters might be wrong. This might downgrade our general perception of the importance of cooperation as an evolutionary cause of sociality.
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Lansade, L., & Bouissou, M. - F. (2008). Reactivity to humans: A temperament trait of horses which is stable across time and situations. Appl. Anim. Behav. Sci., 114(3-4), 492–508.
Abstract: The aim of this study was to test the existence of a “reactivity-to-humans” trait which is stable over time and across situations. For this purpose, four test situations were repeated on the same animals at 8 months, 1.5 years (N = 110 horses) and 2.5 years of age (N = 54 horses). These situations involved a “familiar passive human test” during which a familiar person stayed motionless in the test pen, a “familiar active human test” and an “unfamiliar active human test” during which a familiar or unfamiliar person attempted to touch the horse, and a “halter-fitting and heart-rate measurement test”, during which the horse was haltered and equipped with a surcingle and then had its heart-rate measured. Many behavioural variables were recorded during the tests, and correlations between ages and variables measured during other situations were investigated. When correlations were found, we concluded that the variables corresponded to temperament traits which are stable over time and across situations. The horses tested were divided into four groups according to breed (Anglo-Arab and Welsh) and year of birth (2001 and 2002). Data for each group were analysed separately with Spearman rank correlations. Regarding stability over time, there was a strong correlation between ages in the frequency of licking/nibbling the passive human, the time taken by a familiar or unfamiliar person to touch the horse, and the time taken to fit a halter (e.g. the time taken to touch the horse's muzzle was significantly correlated between ages in 7 out of the 8 possible cases: 0.40 <= R <= 0.67). Regarding stability across situations, results indicate that the more frequently a horse sniffed, licked or nibbled a passive human, the easier it was for either a familiar or unfamiliar active human to touch and halter it. Moreover, a horse which could be touched easily by a familiar human could also be touched and haltered easily by an unfamiliar human. These behaviours, which are linked directly to humans, therefore indicate good stability over time and across situations. In conclusion, this stability suggests the existence of a “reactivity-to-humans” temperament trait, whether the human is passive or active, familiar or unfamiliar. When the animals do not have frequent contact with humans, this reactivity is stable over time and can be measured as early as 8 months of age using the behavioural variables cited above.
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Cameron, E. Z., Linklater, W. L., Stafford, K. J., & Minot, E. O. (2008). Maternal investment results in better foal condition through increased play behaviour in horses. Anim. Behav., 76(5), 1511–1518.
Abstract: Play behaviour is widespread in mammals, but benefits to play have been difficult to demonstrate. Physical training is one of the many proposed hypotheses, suggesting that males and females should play differently, that increased maternal investment should lead to increases in play, and that increases in play should result in physical advantages. In a population of feral horses, Equus caballus, males and females did not differ in their play behaviour except that males initiated more of their play bouts than females. Maternal condition influenced play behaviour only in males, with sons of mothers in good condition playing more. However, when we controlled for maternal effects by comparing a son and a daughter of the same mother, daughters played more when their mother was in poor condition and sons played more when their mother was in good condition. Mothers of foals that played more lost more condition. Therefore, the difference in play behaviour could not be explained by offspring sex or maternal condition alone, but play behaviour mirrored variation in maternal investment. In addition, those individuals that played more survived better and had better body condition as yearlings despite weaning earlier. Since increased activity has been linked to enhanced musculoskeletal development in domestic horses, we suggest that play provides a link between increased maternal investment, increased body condition and future reproductive success in feral horses, and probably in other species.
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Boesch, C. (1991). Teaching among wild chimpanzees. Anim. Behav., 41(3), 530–532.
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Griffin, A. S. (2008). Social learning in Indian mynahs, Acridotheres tristis: the role of distress calls. Anim. Behav., 75(1), 79–89.
Abstract: Socially acquired predator avoidance is a phenomenon in which individuals acquire an avoidance response towards an initially neutral stimulus after they have experienced it together with the antipredator signals of social companions. Earlier research has established that alarm calls used for intraspecific communication are effective stimuli for triggering acquisition. However, animals produce a large range of other antipredator responses that might engage antipredator learning. Here, I examine the effects of conspecific distress calls, a signal that is produced by birds when restrained by a predator, and that appears to be directed towards predators, rather than conspecifics, on predator avoidance learning in Indian mynahs, Acridotheres tristis. Distress calls reflect high levels of alarm in the caller and should, therefore, mediate robust learning. Experiment 1 revealed that subjects performed higher rates of head movements in response to a previously unfamiliar avian mount after it had been presented simultaneously with playbacks of conspecific distress vocalizations. Experiment 2 revealed that increased rates of head saccades resembled the spontaneous response evoked by a novel stimulus more closely than it resembled the response evoked by a perched raptor, suggesting that distress calls inculcated a visual exploratory response, rather than an antipredator response. While it is usually thought that the level of acquisition in learners follows a simple relationship with the level of alarm shown by demonstrators, the present results suggest that this relationship may be more complex. Antipredator signals with different functions may have differential effects on learners.
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Hewitt, S. E., Macdonald, D. W., & Dugdale, H. L. (2009). Context-dependent linear dominance hierarchies in social groups of European badgers, Meles meles. Anim. Behav., 77(1), 161–169.
Abstract: A social hierarchy is generally assumed to exist in those mammalian societies in which the costs and benefits of group living are distributed unevenly among group members. We analysed infrared closed-circuit television footage, collected over 3 years in Wytham Woods, Oxfordshire, to test whether social groups of European badgers have dominance hierarchies. Analysis of directed aggression between dyads revealed linear dominance hierarchies in three social-group-years, but patterns within social groups were not consistent across years. Dominance hierarchies were significantly steeper than random in five out of six social-group-years. In those social-group-years where a linear hierarchy was determined, there was an effect of sex on dominance rank, with females gaining significantly higher rank than males in two social-group-years. Overall, rank was not related to age, nor did it appear to affect the likelihood of an individual being wounded, or an individual's breeding status. The latter resulted from nonorthogonality between sex and breeding status, as there were only two breeding males. Overall, hierarchies were primarily dominated by breeding females, and may occur when breeding competition arises. Relatedness, unreciprocated allogrooming and sequential allomarking were not consistently related to levels of directed aggression across social-group-years. We suggest that dominance structures within European badger groups may be context dependent, with future study required to complete our understanding of where, and when, they arise.
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Palagi, E., Antonacci, D., & Norscia, I. (2008). Peacemaking on treetops: first evidence of reconciliation from a wild prosimian (Propithecus verreauxi). Anim. Behav., 76(3), 737–747.
Abstract: Reconciliation is defined as the first postconflict affinitive contact between former opponents. While reconciliation in anthropoid primates has been widely investigated, few studies have focused on postconflict mechanisms in prosimians, and only in captivity. Unlike anthropoids, Malagasy prosimians show female dominance, lack of sexual dimorphism and seasonal breeding. However, they share features with anthropoids such as cohesive societies, female philopatry and individual recognition. Comparing social prosimians with anthropoids is crucial for understanding the evolution of reconciliation dynamics. Here we present the first study on reconciliation in a wild prosimian. We focused on the Propithecus verreauxi (sifaka) of the Berenty forest (southern Madagascar). We examined postconflict behaviour in the light of theoretical expectations based on potential costs and benefits of the individuals involved. Our results indicate that P. verreauxi can evaluate possible risks and benefits of engaging in postconflict reunions. Victims were most likely to interact affinitively with the aggressor after low-intensity aggression. Moreover, only the conflicts occurring outside the feeding context were reconciled. Such results are consonant with the fact that, in P. verreauxi, social dominance is translated more into feeding priority than into a framework of despotic relationships. In agreement with the valuable relationship hypothesis, P. verreauxi were more likely to reconcile with valuable partners: reconciliation preferentially occurred between subordinates and top-ranking individuals, and between animals sharing good relationships (high levels of affinitive behaviours). Over the short term, reconciliation in P. verreauxi seems to have an important role in reducing the probability of further attacks by the aggressor.
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Whitehead, H. (2008). Precision and power in the analysis of social structure using associations. Anim. Behav., 75(3), 1093–1099.
Abstract: I develop guidelines for assessing the precision and power of statistical techniques that are frequently used to study nonhuman social systems using observed dyadic associations. Association indexes estimate the proportion of time that two individuals are associated. Binomial approximation and nonparametric bootstrap methods produce similar estimates of the precision of association indexes. For a mid-range (0.4-0.9) association index to have a standard error of less than 0.1 requires about 15 observations of the pair associated, and for it to be less than 0.05, this rises to 50 observations. The coefficient of variation among dyads of the proportion of time that pairs of individuals are actually associated describes social differentiation (S), and this may be estimated from association data using maximum likelihood. With a poorly differentiated population (S~0.2), a data set needs about five observed associations per dyad to achieve a correlation between true and estimated association indexes of r=~0.4. It requires about 10 times as much data to achieve a representation with r=~0.8. Permutation tests usually reject the null hypothesis that individuals have no preferred associates when S2H>5, where H is the mean number of observed associations per individual. Thus most situations require substantial numbers of observations of associations to give useful portrayals of social systems, and sparse association data inform only when social differentiation is high.
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Wey, T., Blumstein, D. T., Shen, W., & Jordán, F. (2008). Social network analysis of animal behaviour: a promising tool for the study of sociality. Anim. Behav., 75(2), 333–344.
Abstract: Social animals live and interact together, forming complex relationships and social structure. These relationships can have important fitness consequences, but most studies do not explicitly measure those relationships. An approach that explicitly measures relationships will further our understanding of social complexity and the consequences of both direct and indirect interactions. Social network analysis is the study of social groups as networks of nodes connected by social ties. This approach examines individuals and groups in the context of relationships between group members. Application of social network analysis to animal behaviour can advance the field by identifying and quantifying specific attributes of social relationships, many of which are not captured by more common measures of sociality, such as group size. Sophisticated methods for network construction and analysis exist in other fields, but until recently, have seen relatively little application to animal systems. We present a brief history of social network analysis, a description of basic concepts and previous applications to animal behaviour. We then highlight relevance and constraints of some network measures, including results from an original study of the effect of sampling on network parameter estimates, and we end with promising directions for research. By doing so, we provide a prospective overview of social network analysis' general utility for the study of animal social behaviour.
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