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Conradt, L., Krause, J., Couzin, I. D., & Roper, T. J. (2009). “Leading According to Need” in Self-Organizing Groups. Am Nat, 173(3), 304–312.
Abstract: Self‐organizing‐system approaches have shed significant light on the mechanisms underlying synchronized movements by large groups of animals, such as shoals of fish, flocks of birds, or herds of ungulates. However, these approaches rarely consider conflicts of interest between group members, although there is reason to suppose that such conflicts are commonplace. Here, we demonstrate that, where conflicts exist, individual members of self‐organizing groups can, in principle, increase their influence on group movement destination by strategically changing simple behavioral parameters (namely, movement speed, assertiveness, and social attraction range). However, they do so at the expense of an increased risk of group fragmentation and a decrease in movement efficiency. We argue that the resulting trade‐offs faced by each group member render it likely that group movements are led by those members for which reaching a particular destination is most crucial or group cohesion is least important. We term this phenomenon leading according to “need” or “social indifference,” respectively. Both kinds of leading can occur in the absence of knowledge of or communication about the needs of other group members and without the assumption of altruistic cooperation. We discuss our findings in the light of observations on fish and other vertebrates.
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Dyer, J. R. G., Johansson, A., Helbing, D., Couzin, I. D., & Krause, J. (2009). Leadership, consensus decision making and collective behaviour in humans. Phil. Trans. Biol. Sci., 364(1518), 781–789.
Abstract: This paper reviews the literature on leadership in vertebrate groups, including recent work on human groups, before presenting the results of three new experiments looking at leadership and decision making in small and large human groups. In experiment 1, we find that both group size and the presence of uninformed individuals can affect the speed with which small human groups (eight people) decide between two opposing directional preferences and the likelihood of the group splitting. In experiment 2, we show that the spatial positioning of informed individuals within small human groups (10 people) can affect the speed and accuracy of group motion. We find that having a mixture of leaders positioned in the centre and on the edge of a group increases the speed and accuracy with which the group reaches their target. In experiment 3, we use large human crowds (100 and 200 people) to demonstrate that the trends observed from earlier work using small human groups can be applied to larger crowds. We find that only a small minority of informed individuals is needed to guide a large uninformed group. These studies build upon important theoretical and empirical work on leadership and decision making in animal groups.
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Harcourt, J. L., Ang, T. Z., Sweetman, G., Johnstone, R. A., & Manica, A. (2009). Social feedback and the emergence of leaders and followers. Curr Biol, 19(3), 248–252.
Abstract: In many animal groups, certain individuals consistently appear at the forefront of coordinated movements [1-4]. How such leaders emerge is poorly understood [5, 6]. Here, we show that in pairs of sticklebacks, Gasterosteus aculeatus, leadership arises from individual differences in the way that fish respond to their partner's movements. Having first established that individuals differed in their propensity to leave cover in order to look for food, we randomly paired fish of varying boldness, and we used a Markov Chain model to infer the individual rules underlying their joint behavior. Both fish in a pair responded to each other's movements-each was more likely to leave cover if the other was already out and to return if the other had already returned. However, we found that bolder individuals displayed greater initiative and were less responsive to their partners, whereas shyer individuals displayed less initiative but followed their partners more faithfully; they also, as followers, elicited greater leadership tendencies in their bold partners. We conclude that leadership in this case is reinforced by positive social feedback.
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Cameron, E. Z., Setsaas, T. H., & Linklater, W. L. (2009). Social bonds between unrelated females increase reproductive success in feral horses. Proc. Natl. Acad. Sci. U.S.A., 106(33), 13850–13853.
Abstract: In many mammals, females form close social bonds with members of their group, usually between kin. Studies of social bonds and their fitness benefits have not been investigated outside primates, and are confounded by the relatedness between individuals in primate groups. Bonds may arise from kin selection and inclusive fitness rather than through direct benefits of association. However, female equids live in long-term social groups with unrelated members. We present 4 years of behavioral data, which demonstrate that social integration between unrelated females increases both foal birth rates and survival, independent of maternal habitat quality, social group type, dominance status, and age. Also, we show that such social integration reduces harassment by males. Consequently, social integration has strong direct fitness consequences between nonrelatives, suggesting that social bonds can evolve based on these direct benefits alone. Our results support recent studies highlighting the importance of direct benefits in maintaining cooperative behavior, while controlling for the confounding influence of kinship.
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Mitani, J. C. (2009). Male chimpanzees form enduring and equitable social bonds. Anim. Behav., 77(3), 633–640.
Abstract: Controversy exists regarding the nature of primate social relationships. While individual primates are frequently hypothesized to form enduring social bonds with conspecifics, recent studies suggest that relationships are labile, with animals interacting only over short periods to satisfy their immediate needs. Here I use data collected over 10 years on a community of chimpanzees, Pan troglodytes, at Ngogo, Kibale National Park, Uganda, to investigate whether male chimpanzees establish long-term social relationships and to determine the factors that affect variation in relationship quality and the stability of social bonds. Kinship and dominance rank influenced the quality of relationships. Maternal brothers and males of the same dominance rank class groomed each other more equitably than did unrelated males and males that were dissimilar in rank. In addition, males that formed strong social bonds groomed more equitably than did males that displayed weaker bonds. Social bonds were stable over time, with relationships in one year predicting those in subsequent years. Kinship and the quality of social relationships affected bond stability. Maternal half siblings and males that groomed each other equitably maintained longer-lasting bonds than did nonkin and males that groomed each other unevenly. Virtually all of the males established at least one enduring relationship with another individual. The most enduring bonds formed between a few pairs of maternal brothers and dyads that maintained balanced grooming interactions. These results indicate that male chimpanzees maintain long-lasting and equitable social bonds whose formation is affected by maternal kinship and the quality of social relationships.
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Langergraber, K., Mitani, J., & Vigilant, L. (2009). Kinship and social bonds in female chimpanzees (Pan troglodytes). Am. J. Primatol., 71(10), 840–851.
Abstract: A large body of theoretical and empirical research suggests that kinship influences the development and maintenance of social bonds among group-living female mammals, and that human females may be unusual in the extent to which individuals form differentiated social relationships with nonrelatives. Here we combine behavioral observations of party association, spatial proximity, grooming, and space use with extensive molecular genetic analyses to determine whether female chimpanzees form strong social bonds with unrelated individuals of the same sex. We compare our results with those obtained from male chimpanzees who live in the same community and have been shown to form strong social bonds with each other. We demonstrate that party association is as good a predictor of spatial proximity and grooming in females as it is in males, that the highest party association indices are consistently found between female dyads, that the sexes do not differ in the long-term stability of their party association patterns, and that these results cannot be explained as a by-product of the tendency of females to selectively range in particular areas of the territory. We also show that close kin (i.e. mother-daughter and sibling dyads) are very rare, indicating that the vast majority of female dyads that form strong social bonds are not closely related. Additional analyses reveal that “subgroups” of females, consisting of individuals who frequently associate with one another in similar areas of the territory, do not consist of relatives. This suggests that a passive form of kin-biased dispersal, involving the differential migration of females from neighboring communities into subgroups, was also unlikely to be occurring. These results show that, as in males, kinship plays a limited role in structuring the intrasexual social relationships of female chimpanzees.
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Kurvers, R. H. J. M., Eijkelenkamp, B., van Oers, K., van Lith, B., van Wieren, S. E., Ydenberg, R. C., et al. (2009). Personality differences explain leadership in barnacle geese. Anim. Behav., 78(2), 447–453.
Abstract: Personality in animal behaviour describes the observation that behavioural differences between individuals are consistent over time and context. Studies of group-living animals show that movement order among individuals is also consistent over time and context, suggesting that some individuals lead and others follow. However, the relationship between leadership and personality traits is poorly studied. We measured several personality traits and leadership of individual barnacle geese, Branta leucopsis. We measured body size and scored the dominance of individuals living in a stable group situation before subjecting them to an open-field test, an activity test, a novel-object test, and a leadership test in which the order of the movement of individuals in pairs towards a feeding patch was scored. We found high repeatability for activity and novel-object scores over time. Leadership was strongly correlated with novel-object score but not with dominance rank, activity or exploration in an open field. These results provide evidence that leadership is closely related to some aspects of personality. Interestingly, an individual's arrival at the food patch was affected not only by the novel-object score of the focal individual, but also by the novel-object score of the companion individual, indicating that movement patterns of individuals living in groups are affected by the personality traits of other group members and suggesting that movement patterns of a group may be shaped by the mix of personality types present in the group.
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Eagle, N., & Pentland, A. (2009). Eigenbehaviors: identifying structure in routine. Behav. Ecol. Sociobiol., 63(7), 1057-1066.
Abstract: Longitudinal behavioral data generally contains a significant amount of structure. In this work, we identify the structure inherent in daily behavior with models that can accurately analyze, predict, and cluster multimodal data from individuals and communities within the social network of a population. We represent this behavioral structure by the principal components of the complete behavioral dataset, a set of characteristic vectors we have termed eigenbehaviors. In our model, an individual’s behavior over a specific day can be approximated by a weighted sum of his or her primary eigenbehaviors. When these weights are calculated halfway through a day, they can be used to predict the day’s remaining behaviors with 79% accuracy for our test subjects. Additionally, we demonstrate the potential for this dimensionality reduction technique to infer community affiliations within the subjects’ social network by clustering individuals into a behavior space spanned by a set of their aggregate eigenbehaviors. These behavior spaces make it possible to determine the behavioral similarity between both individuals and groups, enabling 96% classification accuracy of community affiliations within the population-level social network. Additionally, the distance between individuals in the behavior space can be used as an estimate for relational ties such as friendship, suggesting strong behavioral homophily amongst the subjects. This approach capitalizes on the large amount of rich data previously captured during the Reality Mining study from mobile phones continuously logging location, proximate phones, and communication of 100 subjects at MIT over the course of 9 months. As wearable sensors continue to generate these types of rich, longitudinal datasets, dimensionality reduction techniques such as eigenbehaviors will play an increasingly important role in behavioral research.
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Fischhoff, I., Dushoff, J., Sundaresan, S., Cordingley, J., & Rubenstein, D. (2009). Reproductive status influences group size and persistence of bonds in male plains zebra (Equus burchelli). Behav. Ecol. Sociobiol., 63(7), 1035-1043.
Abstract: Animal groups arise from individuals’ choices about the number, characteristics, and identity of associates. Individuals make these choices to gain benefits from their associations. As the needs of an individual change with its phenotype, so too we expect the nature of its associations to vary. In this paper, we investigate how the social priorities of male plains zebra (Equus burchelli) depend on reproductive state. An adult male is either a bachelor, and lacking mating access, or a stallion defending a harem. Multiple harems and bachelor males aggregate in larger herds. Herds frequently split and merge, affording males opportunities to change associates. Over a 4-year period, we sampled the herd associations in a population of 500–700 zebras. To isolate the effects of reproductive state on male social behavior, we account for potential confounding factors: changes in population size, grouping tendencies, and sampling intensity. We develop a generally applicable permutation procedure, which allows us to test the null hypothesis that social behavior is independent of male status. Averaging over all individuals in the population, we find that a typical bachelor is found in herds containing significantly more adults, bachelors, and stallions than the herds of a typical stallion. Further, bachelors’ bonds with each other are more persistent over time than those among stallions. These results suggest that bachelors form cohesive cliques, in which we may expect cooperative behaviors to develop. Stallion–stallion associations are more diffuse, and less conducive to long-term cooperation.
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Franks, D., James, R., Noble, J., & Ruxton, G. (2009). A foundation for developing a methodology for social network sampling. Behav. Ecol. Sociobiol., 63(7), 1079-1088.
Abstract: Researchers are increasingly turning to network theory to understand the social nature of animal populations. We present a computational framework that is the first step in a series of works that will allow us to develop a quantitative methodology of social network sampling to aid ecologists in their social network data collection. To develop our methodology, we need to be able to generate networks from which to sample. Ideally, we need to perform a systematic study of sampling protocols on different known network structures, as network structure might affect the robustness of any particular sampling methodology. Thus, we present a computational tool for generating network structures that have user-defined distributions for network properties and for key measures of interest to ecologists. The user defines the values of these measures and the tool will generate appropriate network randomizations with those properties. This tool will be used as a framework for developing a sampling methodology, although we do not present a full methodology here. We describe the method used by the tool, demonstrate its effectiveness, and discuss how the tool can now be utilized. We provide a proof-of-concept example (using the assortativity measure) of how such networks can be used, along with a simulated egocentric sampling regime, to test the level of equivalence of the sampled network to the actual network.
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