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Ahrendt, L.P.; Labouriau, R.; Malmkvist, J.; Nicol, C.J.; Christensen, J.W. |
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Development of a standard test to assess negative reinforcement learning in horses |
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2015 |
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Applied Animal Behaviour Science |
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Appl. Anim. Behav. Sci. |
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169 |
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38-42 |
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Algometry; Horse behaviour; Learning performance; Operant conditioning; Pressure-release; Horse training |
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Most horses are trained by negative reinforcement. Currently, however, no standardised test for evaluating horses' negative reinforcement learning ability is available. The aim of this study was to develop an objective test to investigate negative reinforcement learning in horses. Twenty-four Icelandic horses (3 years old) were included in this study. The horses were tested in a pressure-release task on three separate days with 10, 7 and 5 trials on each side, respectively. Each trial consisted of pressure being applied on the hindquarter with an algometer. The force of the pressure was increased until the horse moved laterally away from the point of pressure. There was a significant decrease in required force over trials on the first test day (P<0.001), but not the second and third day. The intercepts on days 2 and 3 differed significantly from day 1 (P<0.001), but not each other. Significantly stronger force was required on the right side compared to the left (P<0.001), but there was no difference between first and second side tested (P=0.56). Individual performance was evaluated by median-force and the change in force over trials on the first test day. These two measures may explain different characteristics of negative reinforcement learning. In conclusion, this study presents a novel, standardised test for evaluating negative reinforcement learning ability in horses. |
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0168-1591 |
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Equine Behaviour @ team @ |
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6650 |
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Valderrabano-Ibarra, C.; Brumon, I.; Drummond, H. |
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Development of a linear dominance hierarchy in nestling birds |
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2007 |
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Animal Behaviour. |
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Anim. Behav. |
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74 |
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6 |
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1705-1714 |
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agonistic behaviour; blue-footed booby; dominance; hatch asynchrony; hierarchy; Sula nebouxii; trained winning |
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Theoreticians propose that trained winning and losing are important processes in creating linear animal dominance hierarchies, and experiments have shown that both processes can occur in animals, but their actual roles in creating natural hierarchies are unknown. We described agonism in 18 broods of three blue-footed boobies, Sula nebouxii, a species for which trained winning and losing have been demonstrated, to infer how these processes generate and maintain a natural hierarchy. Ranks in the linear hierarchy that emerged in every brood were initially assigned by asymmetries in age, size and maturity, which led to differences between broodmates in levels of expressed and received aggression and, consequently, to differences in the training of their aggressiveness and submissiveness. Later, ranks appeared to be maintained by the chicks' acquired aggressive and submissive tendencies combined with ongoing effects of persisting differences in size and maturity. Our results suggest that trained winning and trained losing are important in the construction of booby hierarchies but that these two axes of learning are largely independent. Increase in submissiveness occurs over a period of about 10-20 days, and the level of submissiveness reached varies with the amount of aggression received. After training, submissiveness is apparently maintained by a lower level of aggression and increasing use of threats. Threats become increasingly effective as chicks age, but are never as effective as attacks. |
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Equine Behaviour @ team @ |
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4318 |
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Taillon, J.; Cote, S.D. |
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Social rank and winter forage quality affect aggressiveness in white-tailed deer fawns |
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2007 |
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Animal Behaviour. |
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Anim. Behav. |
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74 |
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2 |
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265-275 |
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aggressiveness; agonistic behaviour; body mass loss; dominance status; forage intake; Odocoileus virginianus; resource competition; social behaviour; white-tailed deer; winter forage quality |
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Achieving a high social rank may be advantageous for individuals at high population densities, because dominance status may determine the priority of access to limited resources and reduce individual loss of body mass. The establishment of dominance relationships between individuals involves variable levels of aggressiveness that can be influenced by resource availability. The relationship between social rank and aggressiveness and the impacts of resource abundance on aggressiveness are, however, poorly understood, but may be relevant to understand the mechanisms determining dominance relationships between individuals. We experimentally simulated, in seminatural enclosures, a deterioration of winter forage quality induced by a high-density deer population and examined the effects of (1) social dominance and (2) diet quality on aggressiveness, forage intake and body mass loss of white-tailed deer, Odocoileus virginianus, fawns during two winters. Within diet-quality treatments, fawns were consistently organized into linear hierarchies and showed clear dominance relationships. Dominants initiated more interactions and showed higher aggressiveness than subordinates, but subordinates had higher forage intake than dominants throughout winter. Social rank did not influence cumulative body mass loss of fawns. During both winters, fawns fed the control diet maintained their aggressiveness level, whereas fawns fed the poor-quality diet decreased it. Our experimental approach revealed that white-tailed deer responded to a reduction in winter forage quality by modifying their aggressiveness, indicating that ungulates may show plasticity not only in their foraging behaviour in response to decreased resources but also in their social behaviour. |
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Equine Behaviour @ team @ |
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4203 |
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Lingle, S.; Rendall, D.; Wilson, W.F.; DeYoung, R.W.; Pellis, S.M. |
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Altruism and recognition in the antipredator defence of deer: 2. Why mule deer help nonoffspring fawns |
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2007 |
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Animal Behaviour. |
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Anim. Behav. |
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73 |
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5 |
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907-916 |
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aggressive defence; altruism; behavioural discrimination; cooperation; motivational constraint; mule deer; Odocoileus hemionus; Odocoileus virginianus; recognition error; white-tailed deer |
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Both white-tailed deer, Odocoileus virginianus, and mule deer, O. hemionus, females defend fawns against coyotes, Canis latrans, but only mule deer defend nonoffspring conspecific and heterospecific fawns. During a predator attack, females may have to decide whether to defend a fawn while having imperfect information on its identity obtained from hearing a few distress calls. Although imperfect recognition can influence altruistic behaviour, few empirical studies have considered this point when testing functional explanations for altruism. We designed a series of playback experiments with fawn distress calls to test alternative hypotheses (by-product of parental care, kin selection, reciprocal altruism) for the mule deer's defence of nonoffspring, specifically allowing for the possibility that females mistake these fawns for their own. White-tailed deer females approached the speaker only when distress calls of white-tailed deer fawns were played and when their own fawn was hidden, suggesting that fawn defence was strictly a matter of parental care in this species. In contrast, mule deer females responded similarly and strongly, regardless of the caller's identity, the female's reproductive state (mother or nonmother) or the presence of their own offspring. The failure of mule deer females to adjust their responses to these conditions suggests that they do not defend nonoffspring because they mistake them for their own fawns. The lack of behavioural discrimination also suggests that kin selection, reciprocal altruism and defence of the offspring's area are unlikely to explain the mule deer's defence of nonoffspring. We identify causal and functional questions that still need to be addressed to understand why mule deer defend fawns so indiscriminately. |
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Equine Behaviour @ team @ |
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4211 |
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Cloutier, S.; Newberry, R.C.; Honda, K. |
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Comparison of social ranks based on worm-running and aggressive behaviour in young domestic fowl |
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2004 |
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Behavioural Processes |
Abbreviated Journal |
Behav. Process. |
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65 |
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1 |
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79-86 |
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Aggression; Social behaviour; Dominance; Play; Chickens; Animal welfare |
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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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2090 |
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Jennings, D.J. |
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Limited evidence that visual lateralization is associated with fitness in rutting male fallow deer |
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2014 |
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Animal Behaviour |
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Anim. Behav. |
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92 |
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85-91 |
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aggression; contest behaviour; Dama dama; fallow deer; lateral display; lateralization; mating success; third-party intervention behaviour |
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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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0003-3472 |
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Equine Behaviour @ team @ |
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5802 |
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Austin, N.P.; Rogers, L.J. |
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Limb preferences and lateralization of aggression, reactivity and vigilance in feral horses, Equus caballus |
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2012 |
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Animal Behaviour |
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Anim. Behav. |
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83 |
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1 |
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239-247 |
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aggression; behavioural asymmetry; Equus caballus; eye preference; feral horse; limb preference; reactivity; side bias |
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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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0003-3472 |
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Equine Behaviour @ team @ |
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5651 |
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Cozzi, A.; Sighieri, C.; Gazzano, A.; Nicol, C.J.; Baragli, P. |
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Post-conflict friendly reunion in a permanent group of horses (Equus caballus) |
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2010 |
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Behavioural Processes |
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Behav. Process. |
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85 |
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2 |
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185-190 |
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aggression; affiliative interactions; conflict resolution; horse; post conflict; behaviour; reconciliation |
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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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0376-6357 |
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Equine Behaviour @ team @ |
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5168 |
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Flauger, B.; Krueger, K. |
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Aggressionslevel und Platzangebot bei Pferden (Equus caballus) [ Aggression level and enclosure size in horses (Equus caballus)] |
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2013 |
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Pferdeheilkunde |
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29 |
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4 |
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495-504 |
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Aggression / Verletzungsgefahr / Sozialverhalten / Gruppenhaltung / Pferdehaltung / Eingliederung von Pferden [aggression / injury risk / social behaviour / group housing / horse management / introduction of horses] |
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Viele Pferdebesitzer bevorzugen aus Angst vor aggressiven Interaktionen und Verletzungsgefahr der Tiere untereinander die Einzelhaltung, obwohl von Tierschutzorganisationen die Gruppenhaltung für Pferde empfohlen wird. In dieser Studie beobachteten wir während des alltäglichen Soziallebens als auch bei der Eingliederung von neuen Gruppenmitgliedern das Sozialverhalten, insbesondere das Aggressionsverhalten, von elf Gruppen domestizierter Pferde (Equus caballus) verschiedener Größe und Zusammensetzung. Während des alltäglichen Soziallebens hatten die Gruppe und der Paddock-Typ (Gras / kein Gras) keinen Einfluss auf die Verhaltensweisen, wohingegen die Paddockgröße unter 10000 m2 einen signifikanten Einfluss auf die submissiven Verhaltensweisen (GzLM; n=56; t=-2.061, P=0.044) und einen nicht signifikanten Einfluss auf die aggressiven Verhaltensweisen (GzLM; n=56; t=-1.782, P=0.081) hatte. Allerdings verringerten sich sowohl die aggressiven als auch die submissiven Verhaltensweisen mit steigendem Platzangebot bis zu 10000 m2 (Spearman rank Korrelation; n=56; aggressive Verhaltensweisen: r = -0.313, P = 0.019; submissive Verhaltensweisen: r = -0.328, P = 0.014). Während den Eingliederungen reduzierten sich die Aggressionen pro Stunde mit der Vergrößerung des Platzangebotes (Spearman rank Korrelation; n=28; r=-0.402, P=0.034). Dies zeigte sich noch deutlicher, wenn Beobachtungen mit einem Platzangebot von über 10000 m2 ausgeschlos- sen wurden (Spearman rank Korrelation; n=23; r=-0.549, P=0.007). Während des alltäglichen Soziallebens näherte sich der Aggressionslevel der Nulllinie an, wenn das Platzangebot pro Pferd mehr als 331 m2 betrug. Deshalb empfehlen wir zur Reduzierung des Aggressionslevels und des Verletzungsrisikos von sozial gehaltenen Pferdegruppen ein Platzangebot von mindestens 331 m2 pro Pferd.
[Even though animal welfare organisations propose group housing for horse welfare, many owners stable horses individually for fear of aggressive interactions and injury risks. In the present study we observed social behaviour, and especially aggressiveness, in eleven domestic horse groups (Equus caballus) of different size and composition, in basic social situations and when new group members were introduced. During basic social situations, the group and the type of paddock (grass / no grass) had no effect on any of the behaviours, where- as the enclosure size below 10,000 m2 had a significant effect on submissive behaviour (GzLM; n=56; t=-2.061, P=0.044) and an insignificant effect on aggressive behaviour (GzLM; n=56; t=-1.782, P=0.081). However, aggressive and submissive behaviour dimi- nished with the increase of enclosure sizes up to 10,000 m2 (Spearman rank correlation; n = 56; aggressive behaviour: r = -0.313, P=0.019; submissive behaviour: r=-0.328, P=0.014). During introductions, aggression levels per hour decreased with any increase of enclosure size (Spearman rank correlation; n=28; r=-0.402, P=0.034) and even more when enclosure sizes above 10,000 m2 were excluded (Spearman rank correlation; n=23; r=-0.549, P=0.007). During basic social situations the aggression level approached zero when the space allowance was more than 331 m2 per horse. We therefore recommend keeping horse groups in an enclosure with at least 331 m2 per horse to reduce aggression and injuries.] |
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Equine Behaviour @ team @ |
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5714 |
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Dall, Sasha R. X; Houston, Alasdair I.; McNamara, John M. |
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The behavioural ecology of personality: consistent individual differences from an adaptive perspective |
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2004 |
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Ecology Letters |
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Ecol. Letters |
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7 |
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734-739 |
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Adaptive individual differences, behavioural ecology, behavioural syndromes, evolutionary game theory, life history strategies, personality differences, state-dependent dynamic programming |
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Individual humans, and members of diverse other species, show consistent differences in
aggressiveness, shyness, sociability and activity. Such intraspecific differences in
behaviour have been widely assumed to be non-adaptive variation surrounding
(possibly) adaptive population-average behaviour. Nevertheless, in keeping with recent
calls to apply Darwinian reasoning to ever-finer scales of biological variation, we sketch
the fundamentals of an adaptive theory of consistent individual differences in behaviour.
Our thesis is based on the notion that such .personality differences. can be selected for if
fitness payoffs are dependent on both the frequencies with which competing strategies
are played and an individual`s behavioural history. To this end, we review existing models
that illustrate this and propose a game theoretic approach to analyzing personality
differences that is both dynamic and state-dependent. Our motivation is to provide
insights into the evolution and maintenance of an apparently common animal trait:
personality, which has far reaching ecological and evolutionary implications. |
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