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Beaver, B. V. (1986). Aggressive behavior problems. Vet Clin North Am Equine Pract, 2(3), 635–644.
Abstract: Accurate diagnosis of the cause of aggression in horses is essential to determining the appropriate course of action. The affective forms of aggression include fear-induced, pain-induced, intermale, dominance, protective, maternal, learned, and redirected aggressions. Non-affective aggression includes play and sex-related forms. Irritable aggression and hypertestosteronism in mares are medical problems, whereas genetic factors, brain dysfunction, and self-mutilation are also concerns.
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Cameron, E. Z., Linklater, W. L., Stafford, K. J., & Minot, E. O. (2008). Maternal investment results in better foal condition through increased play behaviour in horses. Anim. Behav., 76(5), 1511–1518.
Abstract: Play behaviour is widespread in mammals, but benefits to play have been difficult to demonstrate. Physical training is one of the many proposed hypotheses, suggesting that males and females should play differently, that increased maternal investment should lead to increases in play, and that increases in play should result in physical advantages. In a population of feral horses, Equus caballus, males and females did not differ in their play behaviour except that males initiated more of their play bouts than females. Maternal condition influenced play behaviour only in males, with sons of mothers in good condition playing more. However, when we controlled for maternal effects by comparing a son and a daughter of the same mother, daughters played more when their mother was in poor condition and sons played more when their mother was in good condition. Mothers of foals that played more lost more condition. Therefore, the difference in play behaviour could not be explained by offspring sex or maternal condition alone, but play behaviour mirrored variation in maternal investment. In addition, those individuals that played more survived better and had better body condition as yearlings despite weaning earlier. Since increased activity has been linked to enhanced musculoskeletal development in domestic horses, we suggest that play provides a link between increased maternal investment, increased body condition and future reproductive success in feral horses, and probably in other species.
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Cloutier, S., Newberry, R. C., & Honda, K. (2004). Comparison of social ranks based on worm-running and aggressive behaviour in young domestic fowl. Behav. Process., 65(1), 79–86.
Abstract: Worm-running is behaviour in which a chick runs carrying a worm-like object while flock mates follow and attempt to grab the object from its beak. We hypothesised that social ranks based on worm-running frequency are stable over time and are positively correlated with social ranks based on success in aggressive interactions when older. At 8-12 days of age, we scored worm-running in 17 groups of 12 female White Leghorn chicks during three 10-min tests. Based on instantaneous scans at 5-s intervals, the bird carrying the `worm' most often was placed in rank one and so on down the rank order. These tests were repeated at 68-70 days of age. An aggression index for each bird was calculated as the number of aggressive acts given, divided by the number given and received, during three 1-h observation periods when the birds were 68-70 days. Ranks obtained in worm-running tests were positively correlated over the two age periods (P<0.05) but were not correlated with ranks based on the aggression index (P>0.05). Our results indicate that worm-running ranks are not predictive of success in aggressive interactions. Instead, worm-running fits some criteria for play.
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de Waal, F. B., & Johanowicz, D. L. (1993). Modification of reconciliation behavior through social experience: an experiment with two macaque species. Child Dev, 64(3), 897–908.
Abstract: Reconciliation, defined as a friendly reunion between former opponents shortly after an aggressive encounter, is common in the stumptail macaque (Macaca arctoides) but rare in the rhesus macaque (M. mulatta). Juveniles of the two species were cohoused for 5 months, after which they were observed with conspecifics only. Control rhesus monkeys, matched in age and sex to the experimental subjects, went through the same procedure without exposure to the other species. A threefold increase in the proportion of reconciled fights was measured in the rhesus subjects. The difference emerged gradually during cohousing with the tutor species and was sustained following removal of this species. Other behavior, such as grooming and aggression, decreased over time. It is suggested that the social attitude of the subjects was affected through contact with a species characterized by a more relaxed dominance style.
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Dugatkin, L. A., & Bekoff, M. (2003). Play and the evolution of fairness: a game theory model. Behav. Process., 60(3), 209–214.
Abstract: Bekoff [J. Consci. Stud. 8 (2001) 81] argued that mammalian social play is a useful behavioral phenotype on which to concentrate in order to learn more about the evolution of fairness. Here, we build a game theoretical model designed to formalize some of the ideas laid out by Bekoff, and to examine whether `fair' strategies can in fact be evolutionarily stable. The models we present examine fairness at two different developmental stages during an individual's ontogeny, and hence we create four strategies--fair at time 1/fair at time 2, not fair at time 1/not fair at time 2, fair at time 1/not fair at time 2, not fair at time 1/fair at time 2. Our results suggest that when considering species where fairness can be expressed during two different developmental stages, acting fairly should be more common than never acting fairly. In addition, when no one strategy was evolutionarily stable, we found that all four strategies we model can coexist at evolutionary equilibrium. Even in the absence of an overwhelming database from which to test our model, the general predictions we make have significant implications for the evolution of fairness.
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Flack, J. C., Jeannotte, L. A., & de Waal, F. B. M. (2004). Play signaling and the perception of social rules by juvenile chimpanzees (Pan troglodytes). J Comp Psychol, 118(2), 149–159.
Abstract: Prescriptive social rules are enforced statistical regularities. The authors investigated whether juvenile chimpanzees (Pan troglodytes) recognize and use enforced statistical regularities to guide dyadic play behavior. They hypothesized (a) that proximity of adults, especially mothers of younger play partners, to play bouts will increase the play signaling of older partners and (b) that when juvenile-juvenile play bouts occur in proximity to adults, older partners will play at a lower intensity than when no adults are present. They found that older and younger partners increase their play signaling in the presence of the mothers of younger partners, particularly as the intensity of play bouts increases. In contrast to their hypothesis, older partners played more roughly when the mothers of younger partners were in proximity. These results suggest that juvenile chimpanzees increase play signaling to prevent termination of the play bouts by mothers of younger partners.
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Funk, M. S. (2002). Problem solving skills in young yellow-crowned parakeets (Cyanoramphus auriceps). Anim. Cogn., 5(3), 167–176.
Abstract: Despite the long divergent evolutionary history of birds and mammals, early avian and primate cognitive development have many convergent features. Some of these features were investigated with a series of tasks designed to assess human infant development. The tasks were presented to young parakeets to assess their means-end problem solving abilities. Examples of these early skills are: attaining and playing with objects, retrieving rewards through use of a stick or rake, or by pulling in rewards on supports or on the ends of strings. Twelve such tasks were presented to 11 young yellow-crowned parakeets ( Cyanoramphus auriceps) to investigate their natural abilities; there was no attempt to train them to do those tasks that they did not spontaneously perform. Six of the birds were parent-raised and five were hand-raised. The birds completed 9 of the 12 tasks, demonstrating all the Piagetian sensorimotor circular reactions, but they failed to hand-watch (“claw-watch”), to stack objects, or to fill a container. Their ordinality on the tasks differed from that of human infants in that locomotion to obtain objects occurred earlier in the avian sequence of development and the mid-level tasks were performed by the two groups of avian subjects in a mixed order perhaps indicating that these abilities may not emerge in any particular order for these birds as they supposedly do for human infants. The hand-raised group needed fewer sessions to complete these means-end tasks.
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Gaunet, F. (2010). How do guide dogs and pet dogs (Canis familiaris) ask their owners for their toy and for playing? Anim. Cogn., 13(2), 311–323.
Abstract: Abstract When apes are not fully understood by humans, they persist with attempts to communicate, elaborating their behaviours to better convey their meaning. Such abilities have never been investigated in dogs. The present study aimed to clarify any effect of the visual attentional state of the owner on dogs’ (Canis familiaris) social-communicative signals for interacting with humans, and to determine whether dogs persist and elaborate their behaviour in the face of failure to communicate a request. Gaze at a hidden target or at the owner, gaze alternation between a hidden target and the owner, vocalisations and contacts in 12 guide and 12 pet dogs were analysed (i) when the dogs were asked by their owners (blind or sighted) to fetch their inaccessible toy and (ii) when the dogs were subsequently given an unfamiliar object (apparent unsuccessful communication) or their toy (apparent successful communication). No group differences were found, indicating no effect of the visual status of the owner on the dogs’ socio-communicative modes (i.e. no sensitivity to human visual attention). Results, however, suggest that the dogs exhibited persistence (but not elaboration) in their “showing” behaviours in each condition, except that in which the toy was returned. Thus, their communication was about a specific item in space (the toy). The results suggest that dogs possess partially intentional non-verbal deictic abilities: (i) to get their inaccessible toy, the dogs gazed at their owners as if to trigger their attention; gaze alternation between the owner and the target direction, and two behaviours directed at the target were performed, apparently to indicate the location of the hidden toy; (ii) after the delivery of the toy, the dogs behaved as if they returned to the play routine, gazing at their owner whilst holding their toy. In conclusion, this study shows that dogs possess partially intentional non-verbal deictic abilities: they exhibit successive visual orienting between a partner and objects, apparent attention-getting behaviours, no sensitivity to the visual status of humans for communication, and persistence in (but no elaboration of) communicative behaviours when apparent attempts to “manipulate” the human partner fail.
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Hanggi, E. B., & Ingersoll, J. F. (2009). Long-term memory for categories and concepts in horses (Equus caballus). Anim. Cogn., 13(3), 451–462.
Abstract: Three horses (Equus caballus) with a history of performing cognitive tasks including discrimination learning, categorization, and concept use were tested to evaluate their long-term memory (LTM) in three experiments. In addition, use of LCD multi-displays for stimulus presentation was incorporated into cognition testing protocol for the first time with horses. Experiment 1 tested LTM for discrimination learning that originally occurred 6 years earlier. Five sets of stimuli were used and the two horses tested showed no decrement in performance on four of the sets; however, both horses did score below chance on one set. Experiment 2 examined long-term categorization recall 10 years after horses had demonstrated the ability to make stimulus selections based on shared characteristics within a given category. The horse tested for LTM after the decade-long interval immediately and consistently applied the previously learned categorization rule to not only familiar but also novel sets of stimuli. Experiment 3 tested another horse for LTM for a relative size concept. This horse had originally demonstrated concept rule use in order to select stimuli based on their relative size to one another. More than 7 years later and without further training, this horse reliably applied the previously established size concept to both familiar and novel sets of stimuli. These findings are the first reports of long-term categorical and conceptual memory in horses and are consistent with observations of domestic and wild horses, which indicate that behavioral and ecological events may be remembered for long periods of time. These studies also demonstrate the adaptive nature of horses with regard to their ability to generalize over several different testing conditions.
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Hausberger, M., & Fureix, C. (2012). On the significance of puzzling behaviours: what do yawning and adult play tell us about horse (Equus caballus) welfare? In K. Krueger (Ed.), Proceedings of the 2. International Equine Science Meeting (Vol. in press). Wald: Xenophon Publishing.
Abstract: Some behaviours remain a mystery and their determinants are still uncertain. Here we propose a critical interpretation of two puzzling behaviours, i.e. yawning and adult play, recorded in large samples of domestic horses kept in sub-optimal conditions (e.g. time-restricted feeding practices, social isolation). Yawning is involved in behavioural state changes, especially in quiet contexts of motor relaxation, but seems also to be triggered by stress and emotional contexts. Thus, we investigated specific correlates of yawning and stereotypic behaviours in two large samples of working horses (n1 = 87, n2 = 59). A clear co-occurrences of yawning and stereotypic behaviours appeared: stereotypic horses yawned more than the non-stereotypic horses (Fisher test, p < 0.02), yawning increased at the same time periods as stereotypic behaviours did (Wilcoxon tests, p < 0.05 to 0.001) and yawning frequency was even sometimes positively correlated with stereotypic behaviour frequencies (Spearman correlations, p < 0.005). Play, more typical of young stages in healthy individuals, occurs rarely at adult stages but then more often in captive/domestic animals (which can be challenging to welfare) than in animals living in natural conditions. Thus, we tested the hypothesis that adult play may reflect altered welfare states in horses, where, as in several species, play rarely occurs at adult stages in natural conditions. We observed the behaviour (in particular social play) of 29 adult riding school horses during occasional outings in a paddock and measured several stress indicators (health-related, behavioural and physiological) when these horses were in their home individual boxes. Results revealed that the number of horses and rates of adult play appeared very high compared to field report data and that most stress indicators measured differed between “players” and “non-players”, revealing that most “playful” animals were more prone to suffer from chronic stress contrarily to “non-playful” horses (Fisher, Chi-square, Mann-Whitney and Spearman correlation tests, p < 0.05 to 0.001). Altogether, these results show that yawning and adult play can both reflect altered welfare states in horses, highlighting that these puzzling behaviours should not be considered, as currently thought, as reliable indicators of good welfare states and / or expression of positive emotions.
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