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Johnstone, R. A. (2001). Eavesdropping and animal conflict. Proc. Natl. Acad. Sci. U.S.A., 98(16), 9177–9180.
Abstract: Fights between pairs of animals frequently take place within a wider social context. The displays exchanged during conflict, and the outcome of an encounter, are often detectable by individuals who are not immediately involved. In at least some species, such bystanders are known to eavesdrop on contests between others, and to modify their behavior toward the contestants in response to the observed interaction. Here, I extend Maynard Smith's well known model of animal aggression, the Hawk-Dove game, to incorporate the possibility of eavesdroppers. I show that some eavesdropping is favored whenever the cost of losing an escalated fight exceeds the value of the contested resource, and that its equilibrium frequency is greatest when costs are relatively high. Eavesdropping reduces the risk of escalated conflict relative to that expected by chance, given the level of aggression in the population. However, it also promotes increased aggression, because it enhances the value of victory. The net result is that escalated conflicts are predicted to occur more frequently when eavesdropping is possible.
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Jennings, D. J. (2014). Limited evidence that visual lateralization is associated with fitness in rutting male fallow deer. Anim. Behav., 92, 85–91.
Abstract: Under certain models of animal competition, individuals are expected to gather information about opponent quality in order to determine whether they should fight or withdraw. However, the ability to process complex information differs between individuals and across brain hemispheres: a feature of vertebrate cognition known as lateralization that is not anticipated by contest models. I investigated the relationship between aggressive behaviour and mating success during the fallow deer, Dama dama, rut and a measure of lateralization derived from eye preference during parallel walking. Results show that there was no relationship between the tendency to escalate to fighting or predictability in the tendency to engage in fighting and lateralization. Conversely, there was a quadratic relationship between third-party intervention behaviour and lateralization: the greater the tendency to intervene in ongoing fights the lower the degree of lateralization. However, individuals that showed lateralization for right-eye use were least likely to be targeted by the intervening male; thus lateralization is beneficial in this context because targeted males are highly likely to lose this subsequent encounter. The relationship between lateralization and mating success was also nonlinear: males that showed little evidence for an eye bias during lateral displays had the greatest mating success. Taken together, individuals that showed lateralization benefited from avoiding being targeted after third-party intervention; conversely, individuals that showed little evidence for lateralization actively intervened during ongoing fights and had higher mating success. These results suggest that, although lateralization does appear to confer a fitness advantage on individuals, this is not as extensive as anticipated.
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Duboscq, J., Agil, M., Engelhardt, A., & Thierry, B. (2014). The function of postconflict interactions: new prospects from the study of a tolerant species of primate. Anim. Behav., 87, 107–120.
Abstract: Aggression can generate anxiety, create uncertainty about its aftermath and jeopardise social relationships. Postconflict interactions serve as conflict management strategies to mitigate these consequences. Whereas postconflict interactions are well characterized in many animals, their functions are still insufficiently investigated. Four functional hypotheses have been proposed: stress reduction, relationship repair, self-protection and benign intent. We aimed to test these hypotheses in females of a tolerant macaque species, the crested macaque, Macaca nigra, under natural conditions, for three postconflict interactions: reconciliation, affiliation and aggression with third parties. Our results provide meaningful contrasts compared with findings in other species. We found no evidence that aggression had consequences for individuals' behavioural indicators of anxiety, although it increased the likelihood of secondary aggression with third parties. There was little evidence for the stress reduction hypothesis as the occurrence of any of the three postconflict interactions investigated had little effect on the measured behavioural indicators of anxiety. Conflict and dyad characteristics also had limited influence on anxiety. The relationship repair function was only partly validated: dyads with stronger bonds or that exchanged more support did not reconcile more often, but dyads with attributes related to the symmetry, stability and predictability (i.e. security) within relationships did. Patterns of initiation and directionality of postconflict interactions in this study population suggest that reconciliation may constitute the signalling of appeasement and benign intent. Furthermore, we found that aggression towards third parties may serve as a source of self-protection and reassertion of the females' social status. The distinctive pattern of postconflict management strategies revealed in wild female crested macaques appears to be related to their typically tolerant social style. These results demonstrate the usefulness of concomitantly studying aggression, postconflict interactions and their functions, to understand conflict management strategies comprehensively, while taking into account the level of social tolerance characterizing the studied society.
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Cozzi, A., Sighieri, C., Gazzano, A., Nicol, C. J., & Baragli, P. (2010). Post-conflict friendly reunion in a permanent group of horses (Equus caballus). Behav. Process., 85(2), 185–190.
Abstract: Gregarious animals living in permanent social groups experience intra-group competition. Conflicts over resources can escalate into costly aggression and, in some conditions, non-dispersive forms of conflict resolution may be favoured. Post-conflict friendly reunions, hence reconciliation, have been described in a variety of species. The aim of this study was to explore, for the first time, the occurrence of reconciliation in a group of domestic horses (Equus caballus) and learn more about strategies used to maintain group cohesion. The behaviour of seven horses living as permanent group in an enclosure for at least 2 years was observed by video for 108 h from June to August 2007. We used a Post-Conflict/Matched Control method to assess the existence of reconciliation and third-party affiliation. Behaviours recorded Post-Conflict, or during Matched Control periods, were classified as affiliative based on previous descriptions of visual communication patterns in horses. The proportion of attracted pairs over total post-conflict situations was significantly greater than the proportion of dispersed pairs, both during dyadic interactions (p < 0.001) and during triadic interactions (p = 0.002). The results of the present study show that both dyadic reconciliation and third-party post-conflict affiliative interactions form important social mechanisms for managing post-conflict situations in horses.
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Austin, N. P., & Rogers, L. J. (2012). Limb preferences and lateralization of aggression, reactivity and vigilance in feral horses, Equus caballus. Anim. Behav., 83(1), 239–247.
Abstract: Observational field studies were conducted on two remote populations of feral horses in Australia to determine whether lateralization is a characteristic of Equus caballus as a species or results from handling by humans. Group 1 had been feral for two to five generations and Group 2 for 10–20 generations. In both groups, left-side biases were present during agonistic interactions and in reactivity and vigilance. Therefore, as in other vertebrates, the right hemisphere appears to be specialized to control agonistic behaviour and responses to potential threats. The leftwards bias was stronger in measures of behaviour involving more aggression and reactivity. Preferences to place one forelimb in front of the other during grazing were also determined. No population bias of forelimb preference was found, suggesting that such limb preferences present in domestic horses may be entrained. Since stronger individual limb preferences were found in immature than in adult feral horses, limb preference may be modified by maturation or experience in the natural habitat. Stronger limb preference was associated significantly with elevated attention to the environment but only in younger feral horses. No sex differences in lateralization were found. The findings are evidence that horses show visual lateralization, as in other vertebrates, not dependent on handling by humans. Limb preference during grazing, by contrast, does appear to depend on experience.
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Drummond, H. (2006). Dominance in vertebrate broods and litters. Quarterly Review of Biology, 81(1), 3–32.
Abstract: Drawing on the concepts and theory of dominance in adult vertebrates, this article categorizes the relationships of dominance between infant siblings, identifies the behavioral mechanisms that give rise to those relationships, and proposes a model to explain their evolution. Dominance relationships in avian broods can be classified according to the agonistic roles of dominants and subordinates as “aggression-submission,” “aggression-resistance, ” “aggression-aggression,” “aggression-avoidance,” “rotating dominance,” and “flock dominance.” These relationships differ mainly in the submissiveness/pugnacity of subordinates, which is pivotal, and in the specificity/generality of the learning processes that underlie them. As in the dominance hierarchies of adult vertebrates, agonistic roles are engendered and maintained by several mechanisms, including differential fighting ability, assessment, trained winning and losing (especially in altricial species), learned individual relationships (especially in precocial species), site-specific learning, and probably group-level effects. An evolutionary framework in which the species-typical dominance relationship is determined by feeding mode, confinement, cost of subordination, and capacity for individual recognition, can be extended to mammalian litters and account for the aggression-submission and aggression-resistance observed in distinct populations of spotted hyenas and the “site-specific dominance” (teat ownership) of some pigs, felids, and hyraxes. Little is known about agonism in the litters of other mammals or broods of poikilotherms, but some species of fish and crocodilians have the potential for dominance among broodmates. Copyright © 2006 by The University of Chicago. All rights reserved.
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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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Schmidt, R., Amrhein, V., Kunc, H. P., & Naguib, M. (2007). The day after: effects of vocal interactions on territory defence in nightingales. T. J. Anim. Ecol., 76(1), 168–173.
Abstract: 1. Models on territory acquisition and tenure predict that territorial animals benefit by adjusting territorial defence behaviour to previous challenges they had experienced within the socially complex environment of communication networks. 2. Here, we addressed such issues of social cognition by investigating persisting effects of vocal contests on territory defence behaviour in nightingales Luscinia megarhynchos (Brehm). 3. Using interactive playback during nocturnal song of subjects, a rival was simulated to countersing either aggressively (by song overlapping) or moderately (by song alternating) from outside the subjects' territory. Thereby, the time-specific singing strategy provided an experimentally controlled source of information on the motivation of an unfamiliar rival. 4. Expecting that nightingales integrate information with time, the same rival was simulated to return as a moderately singing intruder on the following morning. 5. The results show that the vigour with which male nightingales responded to the simulated intrusion of an opponent during the day depended on the nature of the nocturnal vocal interaction experienced several hours before. 6. Males that had received the song overlapping playback the preceding night approached the simulated intruder more quickly and closer and sang more songs near the loudspeaker than did males that had received a song alternating playback. 7. This adjustment of territory defence strategies depending on information from prior signalling experience suggests that integrating information with time plays an important part in territory defence by affecting a male's decision making in a communication network.
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Jonart, L. M., Hill, G. E., & Badyaev, A. V. (2007). Fighting ability and motivation: determinants of dominance and contest strategies in females of a passerine bird. Anim. Behav., 74(6), 1675–1681.
Abstract: The communication of aggressive motivation or fighting ability has important fitness consequences for competing animals. Selection should favour rapid and honest communication between opponents to settle dominance relationships while avoiding prolonged and intense fighting. We investigated factors that influence fighting strategies and contest outcomes in female house finches, Carpodacus mexicanus, specifically focusing on the following questions. (1) What social contexts trigger an aggressive response? (2) Does body size and condition contribute to female fighting ability? (3) Do contextual factors, such as mate presence, nest status, nest proximity, and site experience contribute to fighting motivation? (4) Does contest intensity and duration increase as the differences in fighting ability between opponents decrease? (5) What is the relative contribution of fighting ability and aggressive motivation to the outcome of a contest? We found that aggression was triggered most frequently by female intrusions in the vicinity of nest and by extrapair female intrusions on an established pair. Female fighting and contest outcomes were strongly influenced by body condition and body size, and females were more motivated to initiate fights and won more contests when their mates were present. Females at the later breeding stages and those fighting closer to their nests were dominant and won more fights compared to females at earlier breeding stages or further from their nests. Females initiated a greater proportion of contests against opponents with similar local familiarity and breeding history. Escalated and prolonged contests, while rare, occurred exclusively between females of the most similar body size and condition. When differences in body condition between opponents are not easily perceived, contestants might escalate contests for more reliable assessments of relative fighting ability. Finally, body condition was not a strong determinant of contest outcome in the contexts with easily assessed differences in the resource value (e.g. mate presence), but without these motivational differences, body condition was the ultimate determinant of contest outcomes.
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Heitor, F., do Mar Oom, M., & Vicente, L. (2006). Social relationships in a herd of Sorraia horses Part I. Correlates of social dominance and contexts of aggression. Behav. Process., 73(2), 170–177.
Abstract: Factors related to dominance rank and the functions of aggression were studied in a herd of Sorraia horses, Equus caballus, under extensive management. Subjects were 10 adult mares 5-18 years old and a stallion introduced into the group for breeding. Dominance relationships among mares were clear, irrespective of rank difference, and remained stable after introduction of the stallion. The dominance hierarchy was significantly linear and rank was positively correlated with age and total aggressiveness. Higher-ranking mares received lower frequency and intensity of agonistic interactions. Nevertheless, higher-ranking dominants were not more likely to elicit submission from their subordinates than lower-ranking dominants. Neither close-ranking mares nor mares with less clear dominance relationships were more aggressive towards each other. Agonistic interactions seemed to be used more importantly in regulation of space than to obtain access to food or to reassert dominance relationships. Contexts of aggression were related to mare rank. The results suggest that dominance relationships based on age as a conventional criterion were established to reduce aggressiveness in a herd where the costs of aggression are likely to outweigh the benefits.
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