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Kelly, C. D. (2006). Fighting for harems: assessment strategies during male-male contests in the sexually dimorphic Wellington tree weta. Anim. Behav., 72(3), 727–736.
Abstract: Females often aggregate at particular sites for feeding or shelter, thus giving adult males the opportunity to defend harems and increase male reproductive success. Rival males compete for control of harems via ritualized displays or direct combat using weaponry. Contests for harems or the resources required by females can be settled based on asymmetries in fighting ability or resource ownership. Males that accurately assess a rival's fighting ability prior to engaging in potentially costly combat should be favoured by selection. Game theory and optimality models provide three models to explain how individuals decide to persist in or flee from a fight. These models are the energetic war of attrition, the sequential assessment model and the cumulative assessment model. Using staged contests in the laboratory, I tested predictions of these models using the Wellington tree weta, Hemideina crassidens, a sexually dimorphic insect native to New Zealand. Male H. crassidens use their enlarged mandibles as weapons in fights for access to adult females that reside in cavities in trees. My results supported a prediction common to each assessment model: contest duration was negatively correlated with the asymmetry in opponent's weapon size. The sequential assessment model of contest settlement was partially supported but the strongest support was for the cumulative assessment model. Predictions of the latter model were supported because: (1) fights are apparently settled based on own-size assessment; (2) fights occur in a single phase and escalate; and (3) contests involve physical combat and injury. I suggest that, in nocturnal species, cumulative assessment will generally be most applicable.
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Flack, J. C., Girvan, M., de Waal, F. B. M., & Krakauer, D. C. (2006). Policing stabilizes construction of social niches in primates. Nature, 439(7075), 426–429.
Abstract: All organisms interact with their environment, and in doing so shape it, modifying resource availability. Termed niche construction, this process has been studied primarily at the ecological level with an emphasis on the consequences of construction across generations. We focus on the behavioural process of construction within a single generation, identifying the role a robustness mechanism--conflict management--has in promoting interactions that build social resource networks or social niches. Using 'knockout' experiments on a large, captive group of pigtailed macaques (Macaca nemestrina), we show that a policing function, performed infrequently by a small subset of individuals, significantly contributes to maintaining stable resource networks in the face of chronic perturbations that arise through conflict. When policing is absent, social niches destabilize, with group members building smaller, less diverse, and less integrated grooming, play, proximity and contact-sitting networks. Instability is quantified in terms of reduced mean degree, increased clustering, reduced reach, and increased assortativity. Policing not only controls conflict, we find it significantly influences the structure of networks that constitute essential social resources in gregarious primate societies. The structure of such networks plays a critical role in infant survivorship, emergence and spread of cooperative behaviour, social learning and cultural traditions.
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Aureli, F., Cords, M., & van Schaik, C. P. (2002). Conflict resolution following aggression in gregarious animals: a predictive framework. Anim. Behav., 64(3), 325–343.
Abstract: Knowledge of how animals manage their conflicts is critical for understanding the dynamics of social systems. During the last two decades research on gregarious animals, especially primates, has focused on the mechanisms of conflict management, mainly on friendly postconflict reunions (also called `reconciliation') in which former opponents exchange affiliative behaviour soon after an aggressive conflict. Our aim in this paper is to present a framework in which the costs and benefits of friendly postconflict reunions, both for each individual opponent and for their mutual relationship, are used to predict the patterning of postconflict resolution mechanisms in other gregarious animals. The framework predicts the occurrence of postconflict reunions in species that live in stable social units, have individualized relationships, and experience postconflict hostility, but especially in those in which intragroup aggression disrupts valuable relationships. The critical issue is whether aggressive conflicts occur between cooperative partners and whether the level of aggression is sufficient to jeopardize the benefits associated with such valuable relationships. We conclude by proposing four research priorities to evaluate the role of friendly reunions in negotiating relationships and the way they are themselves influenced by asymmetries in partner value and biological market effects. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.
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Call, J., Aureli, F., & de Waal, F. B. M. (2002). Postconflict third-party affiliation in stumptailed macaques. Anim. Behav., 63(2), 209–216.
Abstract: Stumptailed macaques, Macaca arctoides, are characterized by high levels of postconflict affiliative contacts between opponents. We investigated the occurrence of postconflict affiliative contacts between opponents and third parties that were not involved in the original conflict. We collected 10-min focal observations during postconflict and control periods in which we recorded all aggressive and affiliative behaviours between opponents and third parties. We distinguished three types of third parties depending on the relationship with the focal animal: own kin, opponent's kin and individuals unrelated to both opponents. We analysed the interactions with third parties separately, while distinguishing two classes of affiliative behaviours: (1) allogrooming and contact sitting and (2) sociosexual behaviours (e.g. genital inspection). The macaques showed differences between postconflict and control periods in their affiliative contacts with third parties. Aggressors received more postconflict grooming and contact sitting from their opponents' kin, received more sociosexual behaviour from their own kin and unrelated individuals, and directed more sociosexual behaviour to unrelated individuals. Victims received and directed less postconflict grooming from and towards their own kin. They received more postconflict sociosexual behaviour from all partners except their own kin and directed more sociosexual behaviour to all partners except the opponent's kin. This study establishes the occurrence of multiple postconflict triadic affiliation in stumptailed macaques, and is the first to show that victims receive contacts from third parties in a cercopithecine species, a behaviour previously described only in chimpanzees. It also highlights the importance of analysing the different affiliative behaviours separately in postconflict situations. Otherwise, many of the patterns we report, especially those involving victims, would have been missed.
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Koski, S. E., & Sterck, E. H. M. (2007). Triadic postconflict affiliation in captive chimpanzees: does consolation console? Anim. Behav., 73(1), 133–142.
Abstract: Consolation is a triadic postconflict interaction between a conflict participant and an uninvolved third party. The term consolation implies stress alleviation. Consequently, consolation may be an effective mechanism to alleviate postconflict stress. However, this assumption has not been tested. We tested whether consolation alleviates postconflict stress in captive chimpanzees, Pan troglodytes. In addition, we examined whether consolation is a substitute postconflict interaction for reconciliation. We collected 643 postconflict-matched control pairs on aggressees and 576 on aggressors. Consolation occurred equally frequently with aggressees and aggressors. However, we found no evidence that consolation alleviated stress, regardless of the identity of the consoler. In addition, consolation was also directed to conflict participants with no evident postconflict stress. Furthermore, we found no evidence for consolation being a substitute for reconciliation. The occurrence of consolation did not depend on the occurrence of reconciliation and consolation was not more prevalent with the sex class that reconciled less often or had the highest postconflict stress levels. We conclude that consolation is a postconflict interaction in its own right, the function of which is not likely to be connected to stress alleviation of the consoled individual. We propose that the function of triadic postconflict affiliation, previously labelled as consolation, should be reassessed with regard to the third parties' reasons to affiliate with conflict opponents.
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Mills, D. S., & Riezebos, M. (2005). The role of the image of a conspecific in the regulation of stereotypic head movements in the horse. Appl. Anim. Behav. Sci., 91(1-2), 155–165.
Abstract: Weaving is a common locomotor stereotypy in horses, which has been shown to be reduced by the presence of a mirror in the stable; however, it remains unclear whether this effect is due to the reflected image of a horse or some other property associated with the introduction of a mirror. To investigate this further, the response of six known weaving horses to each of three different types of poster was recorded. The poster images consisted of a life-size image of a horse's face (true image), a version of the same image cut into 54 squares, which were then randomly rearranged (pixilated image) and a blank (white image) display of the same size as the other posters. Four observation periods were included in each of the two days of observation per treatment condition: 08:00-08:30 h, 10:00-10:30 h, 12:00-12:30 h and 16:00-17:00 h. The first observation period was followed by the provision of concentrate feed and forage, the second observation period by exercise, the third by further forage and the fourth by concentrate feed and forage. The horses' activities and positions in the box were scanned at 1 min intervals and the number of observations registering each activity and position was calculated as a percentage of the total number of observations for each observation period. Weaving was significantly less when the horses were provided with the image of a horse's face (mean percentage of observations +/- S.E.M.; 5.56 +/- 1.57), compared to both the pixilated (14.85 +/- 3.06) and white (20.52 +/- 4.12) images. Nodding was significantly less when provided with the true image of a horse (0.45 +/- 0.15) compared to the pixilated image (2.15 +/- 0.67). The pattern of weaving throughout the day differed from that of nodding, and together these results suggest that the two behaviours are not analogous. The presence of the image of the horse's face was associated with a significant increase in alertness (F112,2 = 11.31, p < 0.001) and recorded time spent looking at the poster(F112,2 = 3.46, p < 0.05). The timing of stereotypic head movements in the horse in this study suggests that they are not associated with boredom, or lack of stimulation but rather acute frustration.
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Sondergaard, E., & Halekoh, U. (2003). Young horses' reactions to humans in relation to handling and social environment. Appl. Anim. Behav. Sci., 84(4), 265–280.
Abstract: Forty Danish warmblood colts in two replicates were used to investigate the effect of housing and handling in the rearing period on the reactions to humans. The horses entered the experiment after weaning and were housed either individually (n=16) or in groups of three (n=24). Half of the horses from each housing group were handled three times per week for a period of 10 min. Approach tests were performed in the home environment when the horses were 6, 9, 12, 18, 21, and 24 months old, and an Arena and Human Encounter test was performed in a novel environment when the horses were 12 and 24 months old, respectively. In the home environment, single-housed horses approached sooner and were more easily approached by a human than group-housed horses where no effect of handling was observed. Horses approached sooner and were more easily approached with increasing age. In the Arena and Human Encounter test, single-housed horses expressed less restless behaviour, more explorative behaviour, and less vocalisation than group-housed horses. Handled horses showed lower increase in heart rate during the test than non-handled horses. There was no difference between the number of times single or group-housed horses touched an unfamiliar person in the Arena and Human Encounter test but handled horses approached sooner than non-handled horses. It is concluded that the social environment affected the way horses reacted to humans when tested in the home environment but not in a novel environment. In contrast, handling affected the reactions to humans when tested in the novel environment but not in the home environment. However, handled horses also reacted less to the novel environment in general, thus indicating that handling is a mean of avoiding potential dangerous situations.
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Rietmann, T. R., Stuart, A. E. A., Bernasconi, P., Stauffacher, M., Auer, J. A., & Weishaupt, M. A. (2004). Assessment of mental stress in warmblood horses: heart rate variability in comparison to heart rate and selected behavioural parameters. Appl. Anim. Behav. Sci., 88(1-2), 121–136.
Abstract: The aim of the study was to investigate whether heart rate variability (HRV) could assess alterations of the autonomic nervous system (ANS) at different levels of excitement. The behavioural and physiological responses of 20 warmblood horses to a challenging ground exercise task were studied. Prior to the experiment, the horses were evaluated at rest and during forward walking (FW). The horses were then forced to move backwards continuously during 3 min according to a standardised protocol (BW1). Subsequently, the horses were exposed to two training sessions, after which the backward walking (BW2) was re-evaluated. Heart rate (HR) and HRV-parameters such as the standard deviation of the beat-to-beat intervals (SDRR), the low (LF; sympathetic tone) and high frequency (HF) component of HRV (HF; parasympathetic tone) and their ratio (LF/HF; index representing the sympatho-vagal balance) were sampled at rest, and during FW, BW1 and BW2. Stress-related behaviour during BW1 and BW2 was determined from video recordings. The results of the different evaluations were compared to each other. Compared to rest and FW, the first backward experiment induced a significant rise in HR, LF and LF/HF and a significant decrease of HF. SDRR decreased from both FW and rest with only the latter reaching significance. In BW2 after the training sessions, HR and the parameters of the sympathetic branch of the ANS (LF, LF/HF) were decreased and the vagal tone (HF) increased compared to BW1; all changes were significant. The duration of stress indicating behavioural patterns revealed also a significant decrease of excitement after the training, when backward walking did not differ from forward walking in any parameter. Correlations between HRV-parameters and stress indicating behaviour as well as HR were found. We conclude that the HRV-parameters LF and HF are valuable measures to quantify sympatho-vagal balance, which allows a more precise assessment of the responses of HR and SDRR to mental stress during low intensity exercise.
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Sibbald, A. M., Elston, D. A., Smith, D. J. F., & Erhard, H. W. (2005). A method for assessing the relative sociability of individuals within groups: an example with grazing sheep. Appl. Anim. Behav. Sci., 91(1-2), 57–73.
Abstract: We describe a method for quantifying relative sociability within a group of animals, which is defined as the tendency to be close to others within the group and based on the identification of nearest neighbours. The method is suitable for groups of animals in which all individuals are visible and identifiable and has application as a tool in other areas of behavioural research. A sociability index (SI) is calculated, which is equivalent to the relative proportion of time that an individual spends as the nearest neighbour of other animals in the group and is scaled to have an expectation of 1.0 under the null hypothesis of random mixing. Associated pairs, which are animals seen as nearest neighbours more often than would be expected by chance, are also identified. The method tests for consistency across a number of independent observation periods, by comparison with values obtained from simulations in which animal identities are randomised between observation periods. An experiment is described in which 8 groups of 7 grazing sheep were each observed for a total of 10, one-hour periods and the identities and distances away of the 3 nearest neighbours of each focal animal recorded at 5-min intervals. Significant within-group differences in SIs were found in four of the groups (P < 0.001). SIs calculated using the nearest neighbour, two nearest neighbours or three nearest neighbours, were generally highly correlated within all groups, with little change in the ranking of animals. There were significant negative correlations between SIs and nearest neighbour distances in five of the groups. It was concluded that there was no advantage in recording more than one neighbour to calculate the SI. Advantages of the SI over other methods for measuring sociability and pair-wise associations are discussed.
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Heitor, F., do Mar Oom, M., & Vicente, L. (2006). Social relationships in a herd of Sorraia horses: Part II. Factors affecting affiliative relationships and sexual behaviours. Behav. Process., 73(3), 231–239.
Abstract: The influence of age, dominance rank, kinship and aggressiveness over affiliative relationships and sexual behaviours were analysed in a herd of Sorraia horses, Equus caballus, kept under extensive management. Subjects were 10 adult mares 5-18 years old that had known each other since birth, and a stallion introduced into the group for breeding for the first time. Kinship coefficient and dominance rank were the most important factors affecting affiliative relationships. Bonds were reciprocal and stronger among mares with higher kinship. Mares spent more time in proximity to close-ranking and lower-ranking females. Mares with stronger affiliative relationships or higher relatedness were not less aggressive towards each other. Affiliative relationships between the stallion and the mares were not reciprocal: lower-ranking mares formed stronger bonds with the stallion but he preferred the less genetically related mares for proximity. However, the stallion was involved in sexual behaviours more frequently with the mares that were more genetically related to him. These results suggest that kinship beyond close relatives may affect affiliative relationships both among familiar and among unfamiliar horses. However, the influence of kinship does not imply that horses possess a kin recognition system and alternative explanations are discussed.
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