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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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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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Visser, E. K., van Reenen, C. G., van der Werf, J. T. N., Schilder, M. B. H., Knaap, J. H., Barneveld, A., et al. (2002). Heart rate and heart rate variability during a novel object test and a handling test in young horses. Physiol. Behav., 76(2), 289–296.
Abstract: Forty-one Dutch Warmblood immature horses were used in a study to quantify temperamental traits on the basis of heart rate (HR) and heart rate variability (HRV) measures. Half of the horses received additional training from the age of 5 months onwards; the other half did not. Horses were tested at 9, 10, 21 and 22 months of age in a novel object and a handling test. During the tests, mean HR and two heart variability indices, e.g. standard deviation of beat-to-beat intervals (SDRR) and root mean square of successive beat-to-beat differences (rMSSD), were calculated and expressed as response values to baseline measures. In both tests, horses showed at all ages a significant increase in mean HR and decrease in HRV measures, which suggests a marked shift of the balance of the autonomic nervous system towards a sympathetic dominance. In the novel object test, this shift was more pronounced in horses that had not been trained. Furthermore, statistical analysis showed that the increase in mean HR could not be entirely explained by the physical activity. The additional increase in HR, the nonmotor HR, was more pronounced in the untrained horses compared to the trained. Hence, it is suggested that this nonmotor HR might be due to the level of emotionality. HR variables showed consistency between years, as well as within the second year. These tests bring about a HR response in horses, part of which may indicate a higher level of emotionality; and horses show individual consistency of these HR variables over ages. Therefore, it is concluded that mean HR and HRV measures used with these tests quantify certain aspects of a horse's temperament.
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Visser, E. K., van Reenen, C. G., Hopster, H., Schilder, M. B. H., Knaap, J. H., Barneveld, A., et al. (2001). Quantifying aspects of young horses' temperament: consistency of behavioural variables. Appl. Anim. Behav. Sci., 74(4), 241–258.
Abstract: Performance of horses, whether in sports or in leisure, depends on both physical abilities as well as temperament. The aim of the present work was to measure individual variation and consistency of behavioural variables, related to temperament, in young horses of the same breed and age, and reared under controlled housing conditions and management. A total of 41 Dutch Warmblood horses were tested at 9, 10, 21 and 22 months of age in two behavioural tests, i.e. the novel object test and the handling test. In the novel object test horses were confronted with an open umbrella that was lowered from the ceiling. In the handling test horses were led by a human to cross a bridge. Per test, behavioural variables in the following behavioural classes were observed: locomotor activity, latency times, postural expressions and vocalisations. Within years, all behavioural variables in the handling test, and all but two in the novel object test were positively correlated (0.36<Rs<0.81, P<0.05). For both tests, at 9, 10, 21 and 22 months of age, a principal component analysis (PCA) was carried out to examine whether there were indications for underlying components of these individual behavioural variables that could possibly serve as measures for temperamental traits. The first component in the novel object test could be regarded as `flightiness' and the second as `sensitiveness'. In the handling test, the first component was suggested to relate to `patience', the second component to `willingness to perform'. The temperamental trait `flightiness' (novel object test) as well as the temperamental trait `patience' (handling test) were positively correlated within both years (0.36<Rs<0.65, P<0.05). For the traits `sensitiveness' (novel object test) and `willingness to perform' (handling test) a positive correlation was only found within the first year (0.44<Rs<0.57, P<0.01). A few individual behavioural variables showed consistency over years. Additionally, just one out of four temperamental traits, namely `flightiness', proved to be consistent over years (Rs=0.49, P<0.01). The temperamental trait `patience' showed a trend between years (Rs=0.31, 0.05<P<0.1). It is concluded that the behavioural tests employed in the present study can be used to reliably identify individual behavioural variables and temperamental traits in young horses. Long-term consistency, i.e. between subsequent years, could not be demonstrated convincingly. Nevertheless, future work may indicate that employing the same approach and considering an even longer time period or different phases of the horse's life, long-term consistency does exist.
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Visser, E. K., Van Reenen, C. G., Engel, B., Schilder, M. B. H., Barneveld, A., & Blokhuis, H. J. (2003). The association between performance in show-jumping and personality traits earlier in life. Appl. Anim. Behav. Sci., 82(4), 279–295.
Abstract: For a horse to succeed in a show-jumping career, the individual has to possess both excellent physical abilities as well as a suitable personality to perform under challenging conditions. Forty-one Dutch Warmblood horses were used to develop personality tests and correlations between test variables and early training performances in jumping were studied. In behavioural tests, during the first 2 years of the horses' lives, personality aspects like emotionality, reactivity to human and learning abilities were quantified. At the age of 3, horses were broken and received early training in show-jumping. The inter-relationship between several performance variables measured during this early training phase were studied using principal component analysis (PCA). Variables measured in the different personality tests (novel-object test, handling test, avoidance-learning test and a reward-learning test) showed no correlations, suggesting that these tests all triggered different aspects of a horse's personality. This study indicates that it is possible to predict a substantial part of the show-jumping performance of an individual horse later in life by personality traits earlier in life.
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Hausberger, M., & Muller, C. (2002). A brief note on some possible factors involved in the reactions of horses to humans. Appl. Anim. Behav. Sci., 76(4), 339–344.
Abstract: In order to investigate relationships of adult horses to humans, we developed a simple evaluation test and scores based on observations. The first reactions of 224 adult horses to the presence of an experimenter were observed and scored. All these horses belonged to the same riding school, had the same general housing conditions and were all geldings. The evaluation was based on the horse's posture. Individual differences that could be related to some extent to the breed but also to human factors emerged clearly. French saddlebreds showed more often friendly behaviour than Angloarabs, whereas thoroughbreds were more indifferent. Clear variations occurred between groups of horses that depended on different caretakers. In this school, one caretaker is responsible for the whole daily management of a group of horses and is probably a very important factor in their well-being. The effects of this daily relation to a human seemed to be involved in the reactions to a strange person. Further studies are required to investigate what, in practice, may be determinant.
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Anderson, M. C., & Shettleworth, S. J. (1977). Behavioral adaptation to fixed-interval and fixed-time food delivery in golden hamsters. J Exp Anal Behav, 27(1), 33–49.
Abstract: Food-deprived golden hamsters in a large enclosure received food every 30 sec contingent on lever pressing, or free while their behavior was continuously recorded in terms of an exhaustive classification of motor patterns. As with other species in other situations, behavior became organized into two main classes. One (terminal behaviors) increased in probability throughout interfood intervals; the other (interim behaviors) peaked earlier in interfood intervals. Which class an activity belonged to was independent of whether food was contingent on lever pressing. When food was omitted on some of the intervals (thwarting), the terminal activities began sooner in the next interval, and different interim activities changed in different ways. The interim activities did not appear to be schedule-induced in the usual sense. Rather, the hamsters left the area of the feeder when food was not due and engaged in activities they would normally perform in the experimental environment.
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Cooper, J. J., Ashton, C., Bishop, S., West, R., Mills, D. S., & Young, R. J. (2003). Clever hounds: social cognition in the domestic dog (Canis familiaris). Appl. Anim. Behav. Sci., 81(3), 229–244.
Abstract: This paper reviews the reasons why domestic dogs make good models to investigate cognitive processes related to social living and describes experimental approaches that can be adopted to investigate such processes in dogs. Domestic dogs are suitable models for investigating social cognition skills for three broad reasons. First, dogs originated from wolves, social animals that engage in a number of co-operative behaviours, such as hunting and that may have evolved cognitive abilities that help them predict and interpret the actions of other animals. Second, during domestication dogs are likely to have been selected for mental adaptations for their roles in human society such as herding or companionship. Third, domestic dogs live in a human world and “enculturation” may facilitate the development of relevant mental skills in dogs. Studies of social cognition in animals commonly use experimental paradigms originally developed for pre-verbal human infants. Preferential gaze, for example, can be used as a measure of attention or “surprise” in studies using expectancy violation. This approach has been used to demonstrate simple numerical competence in dogs. Dogs also readily use both conspecific and human social signals (e.g. looking or pointing) as information sources to locate hidden rewards such as food or favourite toys. Such abilities make dogs particularly good models for investigating perspective-taking tasks, where animals are required to discriminate between apparently knowledgeable and apparently ignorant informants.
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Hogue, M. - E., Beaugrand, J. P., & Lague, P. C. (1996). Coherent use of information by hens observing their former dominant defeating or being defeated by a stranger. Behav. Process., 38(3), 241–252.
Abstract: This study examines the role of observation during the formation of triads in female domestic hens. Results indicate that during hierarchy formation, a hen observing agonistic interactions and conflict settlement between its former dominant and a stranger uses this information when in turn confronted by the latter. Under a first condition (E, N = 15 triads), bystanders witnessed their prior dominant being defeated by a stranger before being introduced to them. In a second condition (C1, N = 16 triads), bystanders witnessed the victory of their prior dominant over a stranger. In a third condition (C2, N = 15 triads), bystanders witnessed two strangers establishing a dominance relationship before being introduced to their prior dominant and to a stranger the former had just defeated. The behavioural strategies of bystanders depended on the issue of the conflict they had witnessed. Bystanders of the E condition behaved as having no chance of defeating the stranger. They never initiated an attack against it, and upon being attacked, readily submitted in turn to the stranger. On the contrary, bystanders of the C1 condition behaved as having some chances against the stranger. They initiated attacks in 50% of cases, and won 50% of conflicts against the stranger. Under condition C2, bystanders first initiated contact with the strangers in only 27% of cases, which approximates the average of their chances for defeating the stranger. However, bystanders finally defeated the strangers in 40% of cases. These results suggest that bystanders of conditions E and C1 gained some information on the relationship existing between their prior dominant and the stranger and that they used it coherently, perhaps through transitive inference, thus contributing to the existence of transitive relationships within the triads. Alternate explanations are examined.
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Bond, A. B., Kamil, A. C., & Balda, R. P. (2003). Social complexity and transitive inference in corvids. Anim. Behav., 65(3), 479–487.
Abstract: The social complexity hypothesis asserts that animals living in large social groups should display enhanced cognitive abilities along predictable dimensions. To test this concept, we compared highly social pinyon jays,Gymnorhinus cyanocephalus , with relatively nonsocial western scrub-jays, Aphelocoma californica, on two complex cognitive tasks relevant to the ability to track and assess social relationships. Pinyon jays learned to track multiple dyadic relationships more rapidly and more accurately than scrub-jays and appeared to display a more robust and accurate mechanism of transitive inference. These results provide a clear demonstration of the association between social complexity and cognition in animals. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.
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