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Heird, J. C., Lokey, C. E., & Cogan, D. C. (1986). Repeatability and comparison of two maze tests to measure learning ability in horses. Appl. Anim. Behav. Sci., 16(2), 103–119.
Abstract: Sixteen Quarter Horses were randomly divided into two groups after sorting by age and sex. After a 10-day preconditioning period, each animal was scored for emotionality and trainability. Each group then completed a series of learning tasks in a modified T-maze for 20 consecutive days. Group P/D was initially tested on a simple place-learning task, while Group D/P was trained in a visual discrimination task. The groups were tested alternately on the two tasks with 10-day extinction periods between each task. Upon reaching a criterion of 11 of 12 correct responses (the last 8 responses consecutive), a horse was retired for the day. If this criterion was not attained, the horse completed 20 trials. Learning occurred at a faster rate on the discrimination tasks compared to the gradual learning curves observed on place tasks. Animals learned more rapidly and reached higher levels of performance as the series of tasks progressed. Trainability and emotionality scores tended to predict the final level of learning achieved. Correlations of performance ranks within emotionality and training groups were higher between tasks of the same type than between the different tasks. Rank correlations between odd and even days on each task indicated that the within-group rankings were more consistent on the discrimination task than on the place task.
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Heird, J. C., Whitaker, D. D., Bell, R. W., Ramsey, C. B., & Lokey, C. E. (1986). The effects of handling at different ages on the subsequent learning ability of 2-year-old horses. Appl. Anim. Behav. Sci., 15(1), 15–25.
Abstract: Forty weanling Quarter Horses, from the Pitchfork Ranch, Dickens, Texas, were randomly assigned to one of five groups based on the amount of handling they would receive. The five groups were: (1) limited handling; (2) 1 week of handling; (3) 2 weeks of handling; (4) 3 weeks of handling; (5) continuous handling for 18 months. As 2-year-olds, the horses were tested in a simple place-learning T-maze after being preconditioned. A maximum of 30 daily trials were conducted for 20 consecutive days, with feed location alternating between sides on successive days. If a horse met the criterion of 11 out of 12 correct responses with 8 correct responses being consecutive, it was retired without completing the 30 trials. Group 1 made fewer correct responses during the 20 days than any other group except Group 3. All groups achieved learning by Day 10, but the most-handled group reached a consistently higher percentage of correct responses earlier than the less-handled groups. Group 5, the group receiving the most handling, was the least emotional, as determined by the animals' reactivity to a novel stimulus, and received the highest scores for trainability after being ridden. Preconditioning trainability scores, or estimates of the horses' ability to learn prior to conditioning and testing, tended to predict maze-learning ability. These scores were also significantly correlated with post-maze training scores.
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Heitor, F., & Vicente, L. (2008). Maternal care and foal social relationships in a herd of Sorraia horses: Influence of maternal rank and experience. Appl. Anim. Behav. Sci., 113(1-3), 189–205.
Abstract: The influence of maternal rank and experience on patterns of maternal care and social relationships of foals were investigated in a managed herd of Sorraia horses, Equus caballus. Social interactions and spatial relationships of 13 foals (seven females and six males) born to seven mares were examined from birth to 10 months of life, within the three major periods of foal development. Conflict over suckling between dam and foal was not generally affected by rank and experience, but higher-ranking mothers allowed more suckling during late lactation than lower-ranking mothers. Foals of higher-ranking mares spent more time in proximity to the mother during socialization. Maternal rank and experience did not significantly affect maternal protectiveness, foal independence from the mother or the development of affiliative relationships between foals and group members. Foals of higher-ranking mares received lower frequencies of aggression from other horses only in the first month of life. Dominance relationships among foals depended mainly on aggressiveness and were not associated with maternal rank. The large variability in maternal behaviour, the absence of a significant association between maternal rank and body condition at parturition and the stable social environment within this herd may partly account for the reported results.
Keywords: Horse; Rank; Experience; Maternal care; Social relationships
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Heleski, C. R., Shelle, A. C., Nielsen, B. D., & Zanella, A. J. (2002). Influence of housing on weanling horse behavior and subsequent welfare. Appl. Anim. Behav. Sci., 78(2-4), 291–302.
Abstract: Weaning foals marks a stressful event in horses' lives. Limited research exists regarding different housing methods post-weaning and the long-term implications on horse behavior and welfare. The purpose of this study was to monitor behavior and physiological stress markers in horses weaned individually in solid partition box stalls versus horses weaned in small groups and housed in paddocks. Both treatment groups underwent maternal deprivation stress, but the stalled weanlings had the additive effects of social isolation which prevented them from performing social behaviors. Quarter Horse weanlings from the Michigan State University, Merillat Equine Center, average age 4.5 months, were weaned in 13.4 m2 box stalls (n=6) or in groups of three in a 992 m2 paddock, which had very limited grazing forage and an open shelter available (n=6). Subjects were fed concentrate and hay to National Research Council recommendations. A time budget for 31 observed behaviors was developed. Behavioral observations were made 2 days per week, approximately 6 h per day, for the duration of the 56 days study. Instantaneous samples were recorded every 5 min on each observation day, with equal division between the two treatment groups (n=35 scans per horse per observation day). Focal data were recorded continuously between scans to provide a more detailed ethogram. On each observation day, fecal samples were collected to measure 11,17-dioxoandrostanes, an indicator of glucocorticoid metabolite concentration. Regarding the fecal 11,17-dioxoandrostanes, there was no discernible treatment difference either immediately post-weaning or at the conclusion of the 56 days study. Interestingly, all 12 weanlings showed a 4 week post-weaning increase in 11,17-dioxoandrostanes. The reason for this peak was unclear. Behavioral observations demonstrated a significantly different time budget in paddock-housed weanlings than in stall-housed weanlings (P<0.0001). Paddock-housed weanlings displayed a time budget more like a feral horse time budget, showing more time spent moving and less time spent lying. Paddock-housed weanlings, who had the option of selectively engaging in a broader range of behaviors, showed strong motivation to graze and be near conspecifics. Stalled weanlings spent significantly more time engaged in aberrant behaviors: licking or chewing the stall/shed wall, kicking at the stall/shed wall, pawing, and bucking/rearing bouts (P<0.03). Based on the variety of behaviors shown, the ability to engage in strongly preferred behaviors, and freedom from aberrant behavior, we conclude that the paddock-reared, group-housed weanlings had better welfare. However, there was insufficient evidence to conclude that the stalled weanlings had poor welfare.
Keywords: Horse behavior; Weaning; Housing; Welfare; Time budget
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Hemelrijk, C. K. (2000). Towards the integration of social dominance and spatial structure. Anim. Behav., 59(5), 1035–1048.
Abstract: My aim was to show how individual-oriented (or artificial life) models may provide an integrative background for the development of theories about dominance by including effects of spatial structure. Dominance interactions are thought to serve two different, contrasting functions: acquisition of high rank and reduction of aggression. The model I present consists of a homogeneous virtual world inhabited by artificial agents whose actions are restricted to grouping and dominance interactions in which the effects of winning and losing are self-reinforcing. The two functions are implemented as strategies to initiate dominance interactions and the intensity of aggression and dominance perception (direct or memory based) are varied experimentally. Behaviour is studied by recording the same behavioural units as in real animals. Ranks appear to differentiate more clearly at high than at low intensity of aggression and also more in the case of direct than of memory-based rank perception. Strong differentiation of rank produces a cascade of unexpected effects that differ depending on which function is implemented: for instance, a decline in aggression, spatial centrality of dominants and a correlation between rank and aggression. Insight into the origination of these self-organized patterns leads to new hypotheses for the study of the social behaviour of real animals.
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Hemelrijk, C. K. (1990). Models of, and tests for, reciprocity, unidirectionality and other social interaction patterns at a group level. Anim. Behav., 39(6), 1013–1029.
Abstract: Research on reciprocity is impaired by confusing definitions and often wrongly used statistical tests. Here, two models of the mechanism on which reciprocity may be based are discussed and an initial step towards a new fremework for its analysis is presented. A distinction is made between reciprocity and interchange. In the case of reciprocity, for one kind of act the same kind is received in return. In interchange, however, two different kinds of acts are bartered. Three types of reciprocity/interchange in social actions among all pairs of group-members are distinguished ([`]qualitative', [`]relative' and [`]absolute') on the basis of the precision of the reciprocity/interchange. Permutation procedures for association between matrices (such as the Mantel Z and two other newly derived tests) are used as a statistical test for detecting reciprocity/interchange. A rough comparison of the power of the two new tests is included. The tests can be applied to all kinds of group-living animals and to all sorts of social behaviour. The distinction between the three types of reciprocity/interchange and the matching statistical methods are also useful for defining and detecting other patterns in social interactions, like unidirectionality and associations between different kinds of social behaviour. The influence on social interactions of variables like dominance rank, age and sex can be analysed in the three forms by testing correlations between invented matrices which represent the influence of these variables (the so-called hypothesis matrices) and social interaction matrices. These methods are extended for two categories of individuals, thus allowing the investigation of, for example, reciprocity between males and females. The methods are illustrated with examples of coalition formation and grooming behaviour among captive chimpanzees, Pan troglodytes.
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Henry, S., Hemery, D., Richard, M. - A., & Hausberger, M. (2005). Human-mare relationships and behaviour of foals toward humans. Appl. Anim. Behav. Sci., 93(3-4), 341–362.
Abstract: We studied experimentally whether horse dams influenced foals' relationships with humans. We investigated the influence of the establishment of positive human-mare relationships on foals' behaviour toward humans. Forty-one foals and their dams were involved in this experiment. Half of the mares were softly brushed and fed by hand during a short period (total of 1.25 h) during the first 5 days of their foals' lives (experimental group, n = 21). The other mares were not handled experimentally and their foals received no contact with the experimenter (control group, n = 20). The reactions of both experimental and control foals were recorded under various conditions, first, for 5 min in the presence of a motionless experimenter, when foals were 15 and 30-35 days old, then in an approach test when they were 15 days old and in a saddle-pad tolerance test when they were 30-35 days old. Finally, approach-stroking tests were performed successively by the familiar experimenter when foals were 11-13 months old and by an unfamiliar person when they were 13-15 months old. Several observations strongly suggest that mares can influence their foals' behaviour toward humans: (1) during the handling procedure, experimental foals of protective mares were further from the handler than foals of calm mares (p < 0.001); (2) experimental foals remained, at all ages, closer to the experimenter (p < 0.05) and initiated more physical contacts (sniffing, licking, etc.) with the experimenter (p < 0.05) than control foals; (3) avoidance and flight responses of experimental foals were considerably reduced during approaches by the experimenter (p < 0.01) and they accepted saddle-pads on their backs more easily (p < 0.01) and more quickly (p < 0.01) than control foals. Lastly, the consequences of handling mares had effects that lasted at least until foals were one year old (p < 0.05) and became generalized from experimenter to unfamiliar humans, who could approach and stroke experimental foals rapidly during a test (p < 0.05). This is the first report of an attempt to use observation of mother by foals to facilitate human-foal relationships. The procedure is simple, takes little time and can easily be applied to any dam-foal pair, as it is not intrusive and presents no risks of disrupting mare-foal bonds.
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Henson, S. M., Dennis, B., Hayward, J. L., Cushing, J. M., & Galusha, J. G. (2007). Predicting the dynamics of animal behaviour in field populations. Anim. Behav., 74(1), 103–110.
Abstract: Many species show considerable variation in behaviour among individuals. We show that some behaviours are largely deterministic and predictable with mathematical models. We propose a general differential equation model of behaviour in field populations and use the methodology to explain and predict the dynamics of sleep and colony attendance in seabirds as a function of environmental factors. Our model explained over half the variability in the data to which it was fitted, and it predicted the dynamics of an independent data set. Differential equation models may provide new approaches to the study of behaviour in animals and humans.
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Henzi, S. P., Lycett, J. E., & Weingrill, T. (1998). Mate guarding and risk assessment by male mountain baboons during inter-troop encounters. Anim. Behav., 55(6), 1421–1428.
Abstract: Aggressive herding of females is a frequent but not invariant response by male savannah baboons,Papio cynocephalus, to encounters with other troops. While males in some troops are consistently more likely to herd than those in others, not all inter-troop encounters result in herding, even within particular troops. This suggests that males assess the risk of male invasion posed by each encounter and respond accordingly. We used data from baboon troops in the Drakensberg mountains to determine the rules males follow in deciding whether to herd. Consistent differences between troops were explained only by the adult sex ratio. Males were more likely to herd if the sex ratio of their own troop was female biased, a finding that is concordant with the observation that males are more likely to immigrate into troops where the sex ratio is more female biased than the population average. Differences within troops were a consequence only of encounter distance, with herding being more likely at closer distances. We found a negative correlation between the angle of approach to the other troop and the subsequent angle of deflection. We interpret this to mean that herding functions to increase the distance between the interacting troops, thereby curtailing opportunities for strange males to inspect the troop and determine its sex ratio. In this way, possibly unlike those in other populations, the decision rules of these male baboons are geared to protecting longer-term reproductive prospects.
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Hewitt, S. E., Macdonald, D. W., & Dugdale, H. L. (2009). Context-dependent linear dominance hierarchies in social groups of European badgers, Meles meles. Anim. Behav., 77(1), 161–169.
Abstract: A social hierarchy is generally assumed to exist in those mammalian societies in which the costs and benefits of group living are distributed unevenly among group members. We analysed infrared closed-circuit television footage, collected over 3 years in Wytham Woods, Oxfordshire, to test whether social groups of European badgers have dominance hierarchies. Analysis of directed aggression between dyads revealed linear dominance hierarchies in three social-group-years, but patterns within social groups were not consistent across years. Dominance hierarchies were significantly steeper than random in five out of six social-group-years. In those social-group-years where a linear hierarchy was determined, there was an effect of sex on dominance rank, with females gaining significantly higher rank than males in two social-group-years. Overall, rank was not related to age, nor did it appear to affect the likelihood of an individual being wounded, or an individual's breeding status. The latter resulted from nonorthogonality between sex and breeding status, as there were only two breeding males. Overall, hierarchies were primarily dominated by breeding females, and may occur when breeding competition arises. Relatedness, unreciprocated allogrooming and sequential allomarking were not consistently related to levels of directed aggression across social-group-years. We suggest that dominance structures within European badger groups may be context dependent, with future study required to complete our understanding of where, and when, they arise.
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