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Lloyd, A. S., Martin, J. E., Bornett-Gauci, H. L. I., & Wilkinson, R. G. (2007). Evaluation of a novel method of horse personality assessment: Rater-agreement and links to behaviour. Appl. Anim. Behav. Sci., 105(1-3), 205–222.
Abstract: The efficacy of questionnaire-based personality assessment has been shown in a variety of animal and human personality studies. There has been a recent increase in questionnaire-based studies focussing on equine personality but with a lack of comparability to studies on other species. The aim of this study was to test the reliability of an assessment method originally developed for primates and demonstrate reliability using three criteria (1) assessments by independent observers must agree with one another, (2) these assessments must predict behaviours and real-world outcomes and (3) observer ratings must be shown to reflect genuine attributes of the individuals rated, not merely the observer's implicit personality theories about how traits co-vary. The personality of 61 horses (Equus caballus) was assessed using a questionnaire constructed of 30 behaviourally defined adjectives (BDAs). Horses were each assessed by three judges, in addition to a total of 2 h behaviour observations recorded per horse. Rater agreement was demonstrated for 72.1% of the horses and 25 of the BDAs. Principal component analysis was carried out on the rating data and revealed six underlying personality components that were labelled “dominance”, “anxiousness”, “excitability”, “protection”, “sociability” and “inquisitiveness”. Component scores for horses were correlated against behavioural observations for the same horses and revealed significant correlations with 20 of the recorded behaviour variables. Correlations between specific components and their associated behaviours were logical and consistent with the types of behaviours that would be expected to be linked with such personality types. The data were shown to meet all three criteria and provided strong evidence that the assessment method was reliably measuring horse personality.
Keywords: Personality; Horse; Individual differences; Behaviour; Reliability
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Lloyd, A. S., Martin, J. E., Bornett-Gauci, H. L. I., & Wilkinson, R. G. (2008). Horse personality: Variation between breeds. Appl. Anim. Behav. Sci., 112(3-4), 369–383.
Abstract: Anecdotal evidence from horse owners and handlers suggests the existence of breed typical behaviour and personality in horses. This is further supported by current research on heritability of personality characteristics in a range of species. The Horse Personality Questionnaire (HPQ) is a 25-item rating method that has previously been shown to be reliable for the assessment of personality in horses. Principal component analysis on HPQ data has identified six underlying personality components in horses. These are Dominance, Anxiousness, Excitability, Protection, Sociability and Inquisitiveness. Using the HPQ a survey of 1223 horses of eight different breeds was completed. Data were analysed to explore any differences in personality between breeds across the six personality components. Breed differences in personality were identified, and it was noted that variability between breeds varied between personality components. Anxiousness and Excitability showed the most variation between breeds, whilst Dominance and Protection showed the least variance. The results identified breed typical personalities that were comparable to results from previous studies as well as anecdotal evidence provided by the popular equine literature. The results are discussed in terms of the potential selection pressures that may have resulted in these differences.
Keywords: Horse personality; Breed; Assessment; Questionnaire survey; Animal
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Livoreil, B., & Giraldeau, L. (1997). Patch departure decisions by spice finches foraging singly or in groups. Anim. Behav., 54(4), 967–977.
Abstract: The marginal value theorem predicts that when resources are clumped in space, a forager can maximize its rate of intake by deciding to leave a patch when its current feeding rate falls below the average for the habitat. A group version of the model predicts that when rate-maximizing group members share a patch, they should leave sooner, and each with less gain, than single animals exploiting the same patch. We tested these predictions in the laboratory by measuring patch departure decisions of spice finches, Lonchura punctulataexploiting food patches alone or in groups of three under two habitats that require different travel times. As predicted, group members left the patch sooner and with fewer seeds than single foragers. Unlike the model's assumptions, however, birds did not share the patch equally, and their exploitation curves could not be simply derived from those of single foragers. Grouping decreased the effect of travel time on patch exploitation. Moreover, within each group the bird expected to leave first delayed its departure although it collected fewer seeds than the others. This delayed departure could aim to maintain group membership. We noted an increased variability in seed number collected by group members compared with single foragers, which could be a cost of group foraging.Copyright 1997 The Association for the Study of Animal Behaviour1997The Association for the Study of Animal Behaviour
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Lisa Nash, H., Song, G. K., & Price, E. O. (1987). Head partitions facilitate feeding by subordinate horses in the presence of dominant pen-mates. Appl. Anim. Behav. Sci., 19(1-2), 179–182.
Abstract: The purpose of the following study was to determine if head partitions would facilitate feeding by subordinate horses in the presence of dominant pen-mates. Six pairs of mares, each with established dominant-subordinate relationships, were allowed to compete for feed in a 112-cm trough following 24 h of deprivation. Time spent feeding by each mare in each pair was recorded with a wire-mesh partition, a solid plywood partition or no partition dividing the trough. Differences in feeding times between dominant and subordinate mares were greatest in the absence of a partition and least (P<0.05) with the wire barrier in place. Differences in feeding times with the solid plywood barrier were not significantly different from either of the other treatment conditions. It was concluded that head partitions on a trough facilitate feeding by subordinate horses in the presence of dominant pen-mates and thus provide a more equitable distribution of food resources.
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Linklater, W. L., Cameron, E. Z., Minot, E. O., & Stafford, K. J. (1999). Stallion harassment and the mating system of horses. Anim. Behav., 58(2), 295–306.
Abstract: Feral horse, Equus caballus, breeding groups, called bands, usually include one but sometimes up to five stallions. We found that mares were loyal to single-stallion (SS) or multistallion (MS) bands or were social dispersers (maverick mares, Mv). The spacing and social behaviour of mares and stallions in single- and multistallion bands was measured. Indices of mare well-being were also measured including activity budgets (feeding: MS>SS=Mv; resting: MS<SS=Mv), band and mare travel (MS>SS), maternal effort in maintaining contact with foals (MS=Mv>SS), parasite levels in faeces (MS>Mv>SS), body condition (MS=Mv<SS), fecundity (Mv<MS<SS) and offspring mortality (Mv<MS<SS). We present evidence suggesting that the poorer well-being of maverick mares and multistallion band mares results from greater harassment by stallions. Stallion and mare behaviour and poor reproductive success in multistallion bands were not consistent with explanations for the existence of such bands based on cooperation or alternative mating strategies. We suggest an alternative explanation. Stable relationships between mares and a single stallion may enhance reproductive success by reducing aggression between individuals. Therefore, we propose that there is strong selection pressure for stable, long-term stallion-mare relationships, called consort relationships. We propose the consort hypothesis, that multistallion bands are an artefact of selection for stable relationships that occasionally result in more than one such relationship forming, because mares solicit more than one stallion and stallion dominance changes during band formation. Copyright 1999 The Association for the Study of Animal Behaviour.
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Linklater, W. L., & Cameron, E. Z. (2000). Tests for cooperative behaviour between stallions. Anim. Behav., 60(6), 731–743.
Abstract: Breeding groups with multiple stallions occur sympatrically with single-stallion breeding groups in feral horse, Equus caballus, populations. Mutualism and reciprocal altruism between stallions have been proposed to explain the origin and functioning of multistallion bands. However, empirical support for these hypotheses is contradictory and incomplete. Furthermore, there are no explicit tests of the predictions that each hypothesis makes about stallion behaviour and social structure. We compared nine multistallion and 18 single-stallion bands in the Kaimanawa Ranges, New Zealand. Compared with agonistic behaviours, affiliative behaviours were relatively unimportant in the relationships between stallions within bands. The number of stallions in the band did not have a positive influence on mare group size, stability, home range quality or reproductive success in bands. Furthermore, there was a positive relationship between aggression ('intolerance') by the dominant towards subordinate stallions and the subordinates' effort in mare group defence ('helping') but a negative relationship between helping effort by subordinates and their proximity to, and mating with, the bands' mares. Therefore, the predictions of the mutualism and reciprocal altruism hypotheses were not supported. Indeed, for some of the predictions we found the opposite outcomes to be true. Multistallion bands had significantly poorer reproductive success, and dominant stallions were less tolerant of subordinates that helped most and reduced their access to mares. Nevertheless, in all other respects Kaimanawa stallions in multistallion bands behaved like those described elsewhere. Thus, we reject cooperative hypotheses for multimale breeding groups in horses and discuss the mate parasitism and consort hypotheses as better alternatives. Copyright 2000 The Association for the Study of Animal Behaviour.
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Lingle, S., Rendall, D., Wilson, W. F., DeYoung, R. W., & Pellis, S. M. (2007). Altruism and recognition in the antipredator defence of deer: 2. Why mule deer help nonoffspring fawns. Anim. Behav., 73(5), 907–916.
Abstract: 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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Lingle, S., Rendall, D., & Pellis, S. M. (2007). Altruism and recognition in the antipredator defence of deer: 1. Species and individual variation in fawn distress calls. Anim. Behav., 73(5), 897–905.
Abstract: Mule deer, Odocoileus hemionus, females actively defend fawns against predators, including nonoffspring conspecific fawns and heterospecific white-tailed deer, O. virginianus, fawns. We hypothesized that the defence of nonoffspring fawns was due to a recognition error. During a predator attack, females may have to decide whether to defend a fawn with imperfect information on its identity obtained from hearing only a few distress calls. We examined fawn distress calls to determine whether calls made by the two species and by different individuals within each species were acoustically distinctive. The mean and maximum fundamental frequencies of mule deer fawns were nearly double those of white-tailed deer fawns, with no overlap, enabling us to classify 100% of calls to the correct species using a single trait. A large proportion of calls was also assigned to the correct individual using a multivariate analysis (66% and 70% of mule deer and white-tailed deer fawns, respectively, chance = 6% and 10%); however, there was considerable statistical uncertainty in the probability of correct classification. We observed fawns approach conspecific females in an attempt to nurse; females probed most offspring fawns with their noses before accepting them, and always probed nonoffspring fawns before rejecting them, suggesting that close contact and olfactory information were needed to unequivocally distinguish nonoffspring from offspring fawns. Taken together, these results suggest that acoustic variation alone would probably be sufficient to permit rapid and reliable species discrimination, but it may not be sufficient for mothers to unequivocally distinguish their own fawn from conspecific fawns.
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Lindberg, A. C., Kelland, A., & Nicol, C. J. (1999). Effects of observational learning on acquisition of an operant response in horses. Appl. Anim. Behav. Sci., 61(3), 187–199.
Abstract: The effect of observational learning on the acquisition of an operant response was examined in eighteen riding horses and ponies. The test horses were randomly divided into three groups of six and individually exposed to one of three treatments. An additional horse was trained as a demonstrator, to perform the operant response. The observer horses watched either the demonstrator performing the bin-opening response (Group D+B); the demonstrator standing passively (Group D); or the operant bin in the absence of the demonstrator (Group B). Observers had access to and were free to interact with an identical bin during testing. Observers in Groups D+B and D were socially familiar with the demonstrator. Each test horse was tested once a day for 10 days. An ANOVA revealed no significant differences between treatment groups in the number of responses or the time taken to reach the learning criterion. However, there were highly significant differences between breed types, with non-warmbloods performing more bouts of opening the bin and feeding (p=0.02), feeding from the bin sooner (p=0.01) and reaching the criterion for learning sooner than warmbloods (p=0.05). There was also a significant negative linear relationship between horses' ages and time spent investigating the bin, with younger horses performing more investigative behaviour (y=-3.08x+106.86; p=0.02).
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Ligout, S., Porter, R. H., & Bon, R. (2002). Social discrimination in lambs: persistence and scope. Appl. Anim. Behav. Sci., 76(3), 239–248.
Abstract: Social recognition among familiar unrelated lambs was assessed in a series of tests. Lambs and their mothers were housed together in small groups for 1 week (Original groups; O) then reorganized into new groupings (Recent-groups; R) for the remainder of the experiment. During test series 1, lambs that were paired with a familiar O-group partner, from which they had been separated for 5 days, emitted fewer distress bleats than did those tested with an unfamiliar partner. This same effect was not evident when the test was repeated several hours later, indicating that the animals had become habituated to the testing procedures. Two days later, when given the choice between an O- versus a R-partner (test series 2), lambs did not display a preference for either of the stimulus lambs. However, in an additional two-choice test (test series 3) the subject lambs responded discriminatively to a recent familiar partner that was simultaneously present with an unfamiliar lamb. Overall, the results suggest that lambs are capable of developing discriminative relationships with age-mates from different sub-groups, and that such social discrimination persists over a separation period lasting at least several days. It is not clear whether lambs recognize several individual conspecifics per se or discriminate between members of higher order social categories (e.g. familiar versus unfamiliar individuals). Proximal and distal social discrimination may be mediated by different combinations of sensory modalities.
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