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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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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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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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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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Udell, M. A. R., Dorey, N. R., & Wynne, C. D. L. (2008). Wolves outperform dogs in following human social cues. Anim. Behav., 76(6), 1767–1773.
Abstract: Domestic dogs, Canis familiaris, have been shown capable of finding hidden food by following pointing gestures made with different parts of the human body. However, previous studies have reported that hand-reared wolves, C. lupus, fail to locate hidden food in response to similar points in the absence of extensive training. The failure of wolves to perform this task has led to the proposal that the ability to understand others' intentions is a derived character in dogs, not present in the ancestral population (wolves). Here we show that wolves, given the right rearing environment and daily interaction with humans, can use momentary distal human pointing cues to find food without training, whereas dogs tested outdoors and dogs at an animal shelter do not follow the same human points. In line with past studies, pet dogs tested indoors were successful in following these points. We also show that the reported failure of wolves in some past studies may be due to differences in the testing environment. Our findings indicate that domestication is not a prerequisite for human-like social cognition in canids, and show the need for additional research on the role of rearing conditions and environmental factors in the development of higher-level cognitive abilities.
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Stueckle, S., & Zinner, D. (2008). To follow or not to follow: decision making and leadership during the morning departure in chacma baboons. Anim. Behav., 75(6), 1995–2004.
Abstract: To benefit from group living, group members need to keep the group cohesive by coordinating time and direction of travelling. Self-organization and leadership are two means of coordination and two types of decision can be made on the group level: combined and consensus. We studied the initiation process of group movements during the morning departure of a group of chacma baboons, Papio hamadryas ursinus, from its sleeping site in De Hoop Nature Reserve, South Africa. Findings from other female-bonded primate groups led us to hypothesize that females should play a major role in the decision-making process. Approximately 75% of the adults made a start attempt, with 62 of 92 attempts being by males. There was no sex difference in the probability of being successful when initiating an attempt. Lactating females initiated fewer than pregnant or cycling females. Thus, at least for this group of chacma baboons, leadership appeared to be distributed and the decision about the timing of departure and travel direction seemed to be a partially shared consensus decision with adult males contributing more to the decision outcome, with a slightly more prominent role of the dominant male. Our results do not support the [`]leading females' hypothesis. No behavioural patterns that might serve as specialized signals leading to a more successful recruitment of other group members were observed. The departure process appeared to be coordinated merely through individuals setting an example by moving off.
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Lusseau, D., Whitehead, H., & Gero, S. (2008). Incorporating uncertainty into the study of animal social networks. Anim. Behav., 75(5), 1809–1815.
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Schwab, C., Bugnyar, T., Schloegl, C., & Kotrschal, K. (2008). Enhanced social learning between siblings in common ravens, Corvus corax. Anim. Behav., 75(2), 501–508.
Abstract: It has been suggested that social dynamics affect social learning but empirical support for this idea is scarce. Here we show that affiliate relationships among kin indeed enhance the performance of common ravens, Corvus corax, in a social learning task. Via daily behavioural protocols we first monitored social dynamics in our group of captive young ravens. Siblings spent significantly more time in close proximity to each other than did nonsiblings. We subsequently tested birds on a stimulus enhancement task in model-observer dyads composed of both siblings and nonsiblings. During demonstration the observer could watch the model manipulating one particular object (target object) in an adjacent room. After removing the model, the observer was confronted with five different objects including the former target object. Observers from sibling dyads handled the target object for significantly longer periods of time as compared with the other four available objects, whereas observers from nonsibling dyads did not show a preference for the target object. Also, siblings matched the model's decision to cache or not to cache objects significantly more often than did nonsiblings. Hence, siblings were likely to attend to both, the behaviour of the model (caching or noncaching) and object-specific details. Our results support the hypothesis that affiliate relations between individuals affect the transmission of information and may lead to directed social learning even when spatial proximity has been experimentally controlled for.
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Amant, R. S., & Horton, T. E. (2008). Revisiting the definition of animal tool use. Anim. Behav., 75(4), 1199–1208.
Abstract: Benjamin Beck's definition of tool use has served the field of animal cognition well for over 25 years (Beck 1980, Animal Tool Behavior: the Use and Manufacture of Tools, New York, Garland STPM). This article proposes a new, more explanatory definition that accounts for tool use in terms of two complementary subcategories of behaviours: behaviours aimed at altering a target object by mechanical means and behaviours that mediate the flow of information between the tool user and the environment or other organisms in the environment. The conceptual foundation and implications of the new definition are contrasted with those of existing definitions, particularly Beck's. The new definition is informally evaluated with respect to a set of scenarios that highlights differences from Beck's definition as well as those of others in the literature.
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Boogert, N. J., Reader, S. M., Hoppitt, W., & Laland, K. N. (2008). The origin and spread of innovations in starlings. Anim. Behav., 75(4), 1509–1518.
Abstract: There are numerous reports of novel learned behaviour patterns in animal populations, yet the factors influencing the invention and spread of these innovations remain poorly understood. Here we investigated to what extent the pattern of spread of innovations in captive groups of starlings, Sturnus vulgaris, could be predicted by knowledge of individual and social group variables, including association patterns, social rank orders, measures of neophobia and asocial learning performance. We presented small groups of starlings with a series of novel extractive foraging tasks and recorded the latency for each bird to contact and solve each task, as well as the orders of contacting and solving. We then explored which variables best predicted the observed diffusion patterns. Object neophobia and social rank measures characterized who was the first of the group to contact the novel foraging tasks, and the subsequent spread of contacting tasks was associated with latency to feed in a novel environment. Asocial learning performance, measured in isolation, predicted who was the first solver of the novel foraging tasks in each group. Association patterns did not predict the spread of solving. Contact latency and solving duration were negatively correlated, consistent with social learning underlying the spread of solving. Our findings indicate that we can improve our understanding of the diffusion dynamics of innovations in animal groups by investigating group-dependent and individual variables in combination. We introduce novel methods for exploring predictors of the origin and spread of behavioural innovations that could be widely applied.
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