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Innes, L., & McBride, S. (2008). Negative versus positive reinforcement: An evaluation of training strategies for rehabilitated horses. Appl. Anim. Behav. Sci., 112(3–4), 357–368.
Abstract: Rescued equids are often exposed to rehabilitation and training (or retraining) programmes to improve their physical and psychological well-being as well as to facilitate the re-homing process. Training uses either positive or negative reinforcement learning procedures and it is considered here that, there may be welfare implications associated with using the latter technique as it has the potential to overlay acute stress on animals with a chronic stress life history. The aim of this study, therefore, was to compare these training strategies (negative versus positive reinforcement) on equine behaviour and physiology as the first step in establishing an optimal rehabilitation approach (from a welfare perspective) for equids that have been subjected to chronic stress in the form of long-term neglect/cruelty. Over a 7-week period, 16 ponies (aged 6–18 months) were trained using either positive (‘positive’) (n = 8) or negative reinforcement (‘negative’) (n = 8) techniques to lead in hand, stand to be groomed, traverse an obstacle course and load into a trailer. Heart rate was measured (5 s intervals) on days 1 and 4 of each training week, ‘Pre’- (1 h), ‘During’ (0.5 h) and ‘Post’- (1 h) training session. Ethograms (10.00–20.00 h) outside of the training period were also compiled twice weekly. In addition, weekly arena tests (as a measure of reactivity) were also performed 1 week before and during the 7 weeks of training. Results showed significant differences between the two training schedules for some measures during the latter stages of the trial and suggested that animals trained under a positive reinforcement schedule were more motivated to participate in the training sessions and exhibited more exploratory or ‘trial and error’ type behaviours in novel situations/environments. In this context, the incorporation of positive reinforcement schedules within a rehabilitation programme may be of benefit to the animal from a welfare perspective.
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Jameson, K. A., Appleby, M. C., & Freeman, L. C. (1999). Finding an appropriate order for a hierarchy based on probabilistic dominance. Anim. Behav., 57(5), 991–998.
Abstract: Methods of ranking individuals in a dominance hierarchy that use transitivity of relationships may obscure irregularities. Furthermore, these methods use only a small proportion of the information available from dominance encounters. This paper presents an intuitively appealing and easily implemented alternative to existing methods for ordering dominance data, developed from the work of Batchelder et al. (1992Journal of Mathematical Psychology36, 185-212). The procedure presented here is based on a mathematical model of paired comparisons and it involves only simple estimation procedures. We illustrate its use with data on dominance among red deerCervus elaphus, stags. The results indicate that dominance relationships are well characterized by the scale values that the model provides, and, because the method provides predictions for all pairings of animals, dominance predictions also exist for pairs of animals that have yet to be observed. Moreover, the dominance outcomes predicted by the model using the order scale are highly correlated with actual dominance observations at all levels. Overall, the procedure described provides a solution to the problem of identifying an appropriate order for a near-linear dominance hierarchy.
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Janson, C. H. (1990). Social correlates of individual spatial choice in foraging groups of brown capuchin monkeys, Cebus apella. Anim. Behav., 40(5), 910–921.
Abstract: Individuals in a foraging group of wild bronwn capuchin monkeys choose different spatial positions relative to the rest of the group. Markov analysis of sequencess of individual spatial positions demonstrated significant differnces between individuals, which coul be categorized a posteriori into four homogenous subgroups. An individual's spatial position was related primarily to the amount of aggression it received from the group's dominant male, but also varied with its sex. Spatial choice varied with changes in an individual's social status, but did not vary consistently with seasonal differences in food availability. These results support the hypothesis that individuals compete for preferred spatial positions within a foraging group.
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Janson, C. H. (1990). Ecological consequences of individual spatial choice in foraging groups of brown capuchin monkeys, Cebus apella. Anim. Behav., 40(5), 922–934.
Abstract: Individuals in a foraging group of brown capuchin monkeys choose different spatial positions relative to the rest of the group. An individual's choice of spatial positiion affects its foraging success and perceived predation risk (as measured by vigilance behaviour). The two most dominant group members preferred to forage where their expected forwaging success was greatest. Juveniles chose to forage where their perceived predation risk was least, not where they would achieve the highest foraging success. The positions used by non-dominant adults neither maximized foraging success nor minimized predation risk. It is likely that subordinate adults accept spatial positions with suboptimal ecological consequences to avoid the costs of frequent confrontations with the dominant members of the group over foraging sites in poreferred positions.
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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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Jennings, D. J., Carlin, C. M., & Gammell, M. P. (2009). A winner effect supports third-party intervention behaviour during fallow deer, Dama dama, fights. Anim. Behav., 77(2), 343–348.
Abstract: Male ungulates engage in intense competition for access to females during the breeding season. Although fights are generally dyadic level encounters, they are on occasion disrupted by the intervention of third-party males. We investigated these third-party interventions using predictions derived from Dugatkin's model (Dugatkin 1998, Proceedings of the Royal Society of London, Series B, 265, 433-437) of intervention behaviour. The model argues that when an individual successfully defeats an opponent there is an increase in the probability of winning a subsequent contest: a winner effect. Third-party intervention behaviour is predicted to occur as it serves to prevent either member of a competing dyad from successfully defeating his opponent, achieving a winner effect and subsequently becoming a threat to the intervener. Consistent with model predictions, our results show that intervening males held significantly higher rank than males that did not intervene and were also more likely to be dominant to both of the competing males. Intervening males did not selectively target competitors based on rank, nor did they target males based on overall dyadic rates of aggression between the intervener and competing males. Furthermore, interveners were more likely to have won their interaction immediately prior to intervention and were also likely to win their interaction subsequent to intervention when compared with contest success of the two competing males. Our results are consistent with predictions that support a winner effect for intervention behaviour in fallow deer fights.
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Jennings, D. J., Gammell, M. P., Carlin, C. M., & Hayden, T. J. (2004). Effect of body weight, antler length, resource value and experience on fight duration and intensity in fallow deer. Anim. Behav., 68(1), 213–221.
Abstract: We tested predictions of evolutionary game theory focusing on fight duration and intensity during contests between European fallow deer, Dama dama L. We examined the relation between contest duration and intensity and resource-holding potential (RHP; body weight and antler size), in an effort to reveal the assessment rules used by competing males. We examined other potential determinants of duration and intensity: resource value (the oestrous female) and experience of agonistic interactions. Asymmetry in body weight or antler length of contestants was not correlated with fight duration. Body weight and antler length of the fight winner or loser were also not correlated with fight duration. Neither were the body weight of the heavier or lighter animal or the antler length of the animal that had longer or shorter antlers. A measure of intensity (the jump clash) was positively related to the body weight of the losing animal and the lighter member of the dyad. These results are consistent with the hypothesis that opponents escalate contest intensity based on assessment of their own ability rather than through mutual assessment. There was no evidence that resource value is an important factor in either fight duration or intensity in this population. As the number of fights between pairs of males increased, there was a decrease in fight duration. Fights were longer when at least one member of a competing pair of males had previously experienced a victory.
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Jennings, D. J., Gammell, M. P., Carlin, C. M., & Hayden, T. J. (2003). Is the parallel walk between competing male fallow deer, Dama dama, a lateral display of individual quality? Anim. Behav., 65(5), 1005–1012.
Abstract: During competitive encounters protagonists are expected to use signals of individual quality particularly if there is a risk of injury or death. Lateral presentation of body profile, by which information regarding phenotypic characteristics associated with individual quality are displayed, may represent such a strategy. During aggressive interactions, male fallow deer frequently engage in parallel walking which is assumed to represent a mutual display of quality, as mediated by exposure of the maximal profile of the body or antlers. We examined the context and role of the parallel walk during competitive encounters to investigate whether there was evidence that dyads of competing males were assessing differences in phenotypic characteristics. There was no evidence to support the hypotheses that the parallel walk is a lateral display of body size or weaponry or that its use is associated with a reduced level of escalated or risky behaviours during fighting. Total time spent fighting was not shorter when a parallel walk was present than when there was no parallel walk. The parallel walk was highly associated with fighting and it was more likely to be initiated by the subsequent loser. Furthermore, parallel walking frequently followed bouts of fighting and as such may represent a strategy that permits an animal the opportunity to decide whether to continue fighting. Parallel walking was also associated with a failure to resolve contests in favour of one animal indicating that it may be a means of withdrawing from further fighting without incurring a loss in dominance status. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.
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Jezierski, T., Jaworski, Z., & Górecka, A. (1999). Effects of handling on behaviour and heart rate in Konik horses: comparison of stable and forest reared youngstock. Appl. Anim. Behav. Sci., 62(1), 1–11.
Abstract: Thirty foals and young Konik horses born in 3 consecutive years and reared up to weaning either in a forest reserve (R) or conventional stable (S) were compared with respect to behavioural reactions and heart rate (HR) during handling manipulations. The foals were randomly allocated within sex and rearing group to one of two handling treatments. Intensively handled (IH) foals received a 10-min handling, 5 days/week, beginning at the age of 2 weeks (S foals) or 10 months (R foals), and lasting up to the age of 24 months. During handling IH foals were haltered, touched, rubbed and their feet were picked up; non-handled (NH) foals were not handled except for routine or emergency veterinary care. The horses were tested at the age of approximately 6 months (S only) and 12, 18 and 24 months of age. In a test comprising catching the horse on a paddock, leading away from and towards the stable, picking up feet and being approached by an unfamiliar person, the horses' behaviour was scored and the HR was recorded telemetrically. The IH horses scored better as far as manageability behaviour is concerned (P<0.001) and demonstrated lower HR than the NH ones and the S horses scored better than R ones (P<0.001). Fillies demonstrated higher HR than colts (P=0.007). Youngstock of all groups tended to be less manageable at the age of 24 months than at 18 months. Differences between youngstock stemming from particular harems from the reserve seem to be related to differences in accidental contact with people visiting the forest reserve.
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Johnsson, J. I., & Akerman, A. (1998). Watch and learn: preview of the fighting ability of opponents alters contest behaviour in rainbow trout. Anim. Behav., 56(3), 771–776.
Abstract: The costs associated with initial conflicts could be reduced if animals can assess the fighting ability of possible future opponents by watching their contest success against other individuals. We tested this hypothesis by conducting repeated dyadic dominance trials on size-matched juvenile rainbow trout,Oncorhynchus mykiss. In the first trial a dyadic contest was `observed' by a single fish separated by a transparent divider. In the second trial, the observer was paired against either the `familiar' dominant fish or an unfamiliar dominant fish from the first trial. We predicted that observers should settle conflicts with previewed opponents faster and with less aggression than those with unfamiliar fish. This prediction was supported for observers that lost against a previewed competitor, since these fish reduced their aggression more rapidly than did unfamiliar observers. Familiar observers that won, however, showed a more rapid increase in aggression compared with unfamiliar winning observers. This suggests that, regardless of whether an observer challenges the initial dominant, this `decision' is taken more rapidly in conflicts with preassessed contestants, because of the a priori information about their fighting ability. Since preassessment could save energy and allow effort to be concentrated on contests with a high payoff/probability of winning, selection may favour preview strategies when contest competition over resources is important for fitness.
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