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Argue, C. K., & Clayton, H. M. (1993). A preliminary study of transitions between the walk and trot in dressage horses. Acta Anat (Basel), 146(2-3), 179–182.
Abstract: The object of this study was to determine the limb support sequence during the transitions from walk to trot and from trot to walk in dressage horses under saddle and to test the null hypothesis that the limb support sequence during the transitions is not related to the level of training. Sixteen dressage horses training at novice to FEI Grand Prix level were videotaped performing an average of 9 transitions each from walk to trot and from trot to walk. The 30-Hz videotapes were viewed in slow motion, and based on the limb support sequence the transitions were categorized into two types. In type 1 transitions there were no intermediate steps between the walk and trot sequences. Type 2 transitions were characterized by intermediate steps, including a single support phase. The Kendall rank-order correlation coefficient showed that a higher level of training was positively associated with an increased percentage of type 1 transitions for both walk-to-trot transitions (p < or = 0.05) and trot-to-walk transitions (p < or = 0.01). No significant preference for initiating or completing the trot on the left or right diagonal was found using the binomial test for individual horses and the Wilcoxon signed-ranks test for the group.
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Meunier, H., Leca, J. B., Deneubourg, J. L., & Petit, O. (2006). Group movement decisions in capuchin monkeys: the utility of an experimental study and a mathematical model to explore the relationship between individual and collective behaviours. Behaviour, 143, 1511–1527.
Abstract: In primate groups, collective movements are typically described as processes dependent on leadership mechanisms. However, in some species, decision-making includes negotiations and distributed leadership. These facts suggest that simple underlying processes may explain certain decision mechanisms during collective movements. To study such processes, we have designed experiments on white-faced capuchin monkeys (Cebus capucinus) during which we provoked collective movements involving a binary choice. These experiments enabled us to analyse the spatial decisions of individuals in the group. We found that the underlying process includes anonymous mimetism, which means that each individual may influence all members of the group. To support this result, we created a mathematical model issued from our experimental data. A totally anonymous model does not fit perfectly with our experimental distribution. A more individualised model, which takes into account the specific behaviour of social peripheral individuals, revealed the validity of the mimetism hypothesis. Even though white-faced capuchins have complex cognitive abilities, a coexistence of anonymous and social mechanisms appears to influence their choice of direction during collective movements. The present approach may offer vital insights into the relationships between individual behaviours and their emergent collective acts.
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Robbins, M. M., Robbins, A. M., Gerald-Steklis, N., & Steklis, H. D. (2005). Long-term dominance relationships in female mountain gorillas: strength, stability and determinants of rank. Behaviour, 142(6), 779–809.
Abstract: A common practice in studies of social animals is to rank individuals according to dominance status, which has been shown to influence access to limited resources and stability of social relationships, and may in turn correlate with reproductive success. According to the socioecological model for primates, most female dominance relationships are either nepotistic or virtually undetectable (egalitarian), with nepotistic species being philopatric, and dispersing females being egalitarian. Female mountain gorillas (Gorilla beringei beringei) disperse, and they have been characterized as being egalitarian, but previous studies have not examined their dominance relationships from a long-term perspective. We evaluated 15 matrices of displacement/supplantation interactions that spanned 30 years of observations in the Virunga Volcanoes region, and included 51 female mountain gorillas in six groups. Only 4% of displacements were directed against higher ranking females, and when matrices had less than 5% unknown dyads, linearity indices were consistently greater than 0.95. Therefore, previous results suggesting undetectable dominance relationships may have reflected an insufficient quantity of data for this species, rather than actual nonlinearity in its hierarchies. Dominance depended on age and group tenure rather than nepotism, yet some females maintained a high ranking for most of adulthood (15-25 years). Most rank shifts occurred through changes in group composition, rather than switches in established relationships. These results fit within growing evidence for linear individualistic hierarchies in some primates, often coupled with dispersal, as commonly found in ungulates. In light of these results, we propose that the dominance relationships of female mountain gorilla are best characterized as “Dispersal-Individualistic” instead of the previously suggested “Dispersal-Egalitarian”.
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Stevens, J., Vervaecke, H., de Vries, H., & Van Elsacker, L. (2005). The influence of the steepness of dominance hierarchies on reciprocity and interchange in captive groups of bonobos (Pan paniscus). Behaviour, 142(7), 941–960.
Abstract: Biological market models explain variability in reciprocity and interchange between groups. In groups with a shallow dominance gradient, grooming will be mostly exchanged for itself (i.e. exchange will occur). In groups with steep dominance hierarchies, interchange is expected: individuals will groom higher ranking individuals to get access to limited resources or commodities such as support in conflicts, and grooming will be traded for these commodities. We examine patterns of reciprocity in grooming and support, and of interchange of grooming for support or for tolerance in six captive groups of bonobos. We test whether differences between groups in patterns of reciprocity and interchange can be attributed to differences in a measure of steepness of dominance hierarchies, which is based on dyadic agonistic interactions. We found that grooming was reciprocal in some, but not all groups. Support was highly reciprocal, but this was a side effect of dominance in most groups. Interchange between grooming and support was observed in some groups. Corroborating earlier findings, this was a side effect of individuals preferring high ranking individuals as grooming and support partners, possibly because these high-ranking individuals provide more efficient support in conflicts. There was no evidence for interchange of grooming for tolerance. Variability in grooming reciprocity was explained by differences in steepness of dominance hierarchies, as predicted by the biological market models. In groups with a shallow dominance hierarchy, grooming was more reciprocal. This was not true for reciprocity in support. There was some evidence that individuals groomed dominants more frequently in groups with a steep dominance hierarchy. The variation in interchange relations between grooming and support did not depend on the steepness of dominance hierarchies. We suggest that grooming in itself is a valuable commodity in bonobos, especially under captive conditions, which can be exchanged reciprocally. Bonobos may interchange grooming for another value equivalent, with food sharing as a very likely candidate. This interchange effects seem more dependent on potential to monopolise food than on steepness of dominance hierarchies per se.
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Granquist, S. M., Thorhallsdottir, A. G., & Sigurjonsdottir, H. (2012). The effect of stallions on social interactions in domestic and semi feral harems. Applied Animal Behaviour Science, 141(1–2), 49–56.
Abstract: Earlier research indicates that stallions may supress interactions of their harem members, leading to less stable hierarchies and friendship bonds in harems compared to non-stallion groups. In this paper, the effect of the presense of a stallion on the social behaviour of mares was studied by comparing six harems containing stallions to four mixed sex groups not containing stallions. Both temporary and permanent harems were studied, giving the possibility to investigate the effect of group stability on social interactions. A significant linear hierarchy was found in all non-stallion groups that were used for comparison, while the hierarchies were only found to be linear in three of the six harems containing stallions (Landaus h', p < 0.05). Aggression rate was lower (t-test, p < 0.05) and fewer friendship bonds (G-test, p < 0.0001) were found within the harems, compared to the groups without stallions. Stallions seldom intervene directly in interactions between harem members. Thus, our results give support to the hypothesis that stallions may suppress interactions of harem members, but in a more indirect way than with direct interference. In addition, our results give support for earlier findings that aggression rate may be affected by group stability. We found a higher aggression rate in the temporary harems compared to the permanent harems (Kruskal–Wallis, p < 0.05) and in the temporary non-stallion group compared to the permanent non-stallion group. The results have significance for further research on social structure of mammals, and may be applied in management of domestic animals.
Keywords: Horses; Hierarchy; Icelandic horse; Social behaviour; Social bonds
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Sigurjónsdóttir, H., van Dierendonck, M. C., Snorrason, S., & Thórhallsdóttir, A. G. (2003). Social relationships in a group of horses without a mature stallion. Behaviour, 140(6), 783–804.
Abstract: 1. The social relationships in a group of Icelandic horses without a mature stallion were studied. The horses were all familiar to each other. Mutual grooming and play relationships, spatial associations, dominance-subordinate relations and the effect of kinship on these relationships were analysed.TAGSTARTBRTAGEND 2. The social structure was clearly dominated by the behaviour of the adult mares. The horses preferred to form bonds within their social class (sex/age) and they kept close proximity with their friends. The group was effectively divided into two social subgroups, adult mares as one group and adult geldings and sub-adults as another group. The sub-adults and adult geldings formed associations, which were based on mutual grooming and play, while the adult mares did not play. Differences between the sexes were evident. Males played more than the females, had more playing partners and were more popular as playmates.TAGSTARTBRTAGEND 3. Aggression rates were low. The dominance hierarchy was linear. Adult mares ranked higher than adult geldings, sub-adults and the foals. Rank was significantly correlated with age. The closer the adult mares were in rank, the more they groomed with each other. Such relationships were not found amongst the other social group.TAGSTARTBRTAGEND 4. Kinship was calculated between all pairs of animals for up to 4 or 5 generations. Allogrooming and play frequencies and proximity were all positively correlated with kinship. Adult mares, which were close in the dominance hierarchy, were on average more related than those further apart.TAGSTARTBRTAGEND 5. The social relationships in the Icelandic herd were, to some extent, different from relationships reported from unmanaged and feral horse-herds with mature stallions and bachelors. Our results suggest that adult mares groom more in groups without a stallion. Furthermore, they have more preferred partners than in natural harems and their partners are other adult mares, not their weaned offspring as seems to be the case in feral herds. The sub-adults also seem to be more socially active in the absence of stallions. Interestingly, in the Icelandic group, the adult mares showed stallion like behaviours, like mounting and protecting foals. Only by studying the behaviour and the nature of the relationships of horses in groups of different compositions, can we expect to gain a comprehensive understanding about individual social strategies and cognitive capabilities of the species. Such knowledge is valuable for management and welfare of the horse.
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Van de Weerd, H. A., Seaman, S., Wheeler, K., Goddard, P., & Mclean, B. (2012). Use of artificial drinkers by unhandled semi-feral ponies. Appl. Anim. Behav. Sci., 139(1-2), 86–95.
Abstract: This study investigated drinking behaviour of unhandled, semi-feral Dartmoor ponies. Aspects studied were drinking behaviour, latency to drink from novel unfamiliar drinkers after transport, preferences for different types of artificial water drinkers, effects of mixing with unfamiliar ponies and group size, on drinking behaviour, and the effect of a simulated market on the latency to drink. Ponies were tested in groups of three or six animals, or as individuals in test pens that were equipped with three water drinkers: bucket, automatic drinking bowl, flowing water trough. Behaviour was recorded using time-lapse video. An individual pony drank on average 10 l per day. Ponies also drank, but at a lower rate, during the night. The latencies to drink after 4.5 h of transport showed large variation, but most ponies drank within the first hour after being transported (all groups 80.5 ± 32.94 min, mean ± SEM). In the individual choice tests, the preferred drinkers were the bucket and the flowing water trough, but not the automatic drinking bowl (drinking time 25.2 ± 4.66, 11.5 ± 4.26, 2.4 ± 2.23 min for bucket, trough and bowl respectively, mean ± SEM; paired t-tests, bowl versus other drinkers, all tests p < 0.02). A possible reason for the avoidance of the automatic bowl was the noise it made when filling. After mixing a group of three ponies with a group of three unfamiliar animals, the ponies did not express their individual drinker preferences anymore. The use of the previously preferred bucket decreased significantly and the use of the initially, non-preferred, bowl increased significantly. This was likely caused by the fact that ponies were either intentionally or accidentally obstructing drinkers in certain areas of the pen and unfamiliar ponies did not want to push past them. In the simulated market, the differences in latencies to drink between ponies in the home pen and market groups did not reach significance. No significant effect of group size (groups of three versus six ponies) on drinking behaviour was detected. The results have implications for situations where only automatic water bowls are provided, such as during pony sales at livestock markets. Preventing ponies from expressing their drinking choice, either by offering non-preferred drinkers or by mixing with unfamiliar animals, could adversely affect their welfare especially if this happens in conjunction with other stressful events such as transport and markets, and potentially weaning.
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Vervaecke, H., de Vries, H., & van Elsacker, L. (2000). The Pivotal Role Of Rank In Grooming And Support Behavior In A Captive Group Of Bonobos (Pan Paniscus). Behaviour, 137(11), 1463–1485.
Abstract: We investigated dyadic grooming relationships in a captive group of bonobos (Pan paniscus) and questioned what social function grooming fulfils in the 'market of services and favors'. Hereto we examined which of two theoretical models – grooming for support (Seyfarth, 1977, 1980) or grooming according to the similarity principle (de Waal & Luttrell, 1986) – best accounted for the observed grooming distribution. Similarity in traits did not correlate with increased grooming or close proximity among the individuals. Therefore, the similarity hypothesis was rejected. Seyfarth's model of rank-related grooming was largely confirmed. The animals distributed their grooming according to the rank of the receivers. We found an exchange between grooming and receipt of support. There was more grooming up than down the hierarchy. However, not all predictions about rank-related competition over grooming were confirmed. We found that dyadic grooming reciprocity indeed increased with decreasing rank distance. Yet, there was no increase of grooming within the dyad with decreasing rank distance and high ranking individuals were not competed over at the highest rates. The observed correlation between grooming and support received represents an important fit with Seyfarth's prediction, but does not allow for conclusions about underlying causal processes. Other causal explanations, besides the 'groom to receive support' hypothesis, that could explain a similar correlation are discussed.
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Vervaecke, H., Vries, H. D., & Elsacker, L. V. (1999). An Experimental Evaluation Of The Consistency Of Competitive Ability And Agonistic Dominance In Different Social Contexts In Captive Bonobos. Behaviour, 136(4), 423–442.
Abstract: Bonobos have been described as a relatively egalitarian and female dominant species. The exact nature and quality of their dominance relationships and the existence of female dominance are current topics of dispute. We investigated the consistency across social contexts, the stability in time, and the degree of expression of the competitive feeding ability and agonistic dominance in a captive group of bonobos. First, we examined whether the competitive feeding ranks and agonistic ranks differed in different dyadic contexts, triadic contexts and the whole group context. For some pairs of animals the dominance relationships with respect to competitive feeding altered with different group compositions. The agonistic dominance relationships changed accordingly. The competitive feeding ranks and agonistic ranks in the experiments correlated strongly with each other. The alpha position was occupied by a female, but not all females outranked all males. We suggest that females can profit from each others presence to gain inter-sexual dominance. Second, although the agonistic rank order in the whole group remained the same over at least five years, some dyadic competitive feeding ranks changed over time, resulting in a stronger female intersexual dominance. Third, the degree of expression of the behaviors used to quantify dyadic competitive and agonistic dominance was not high, in line with the popular 'egalitarian' epithet. Notwithstanding its low consistency across contexts, the dominance hierarchy in the whole group has a strong predictive value for other social relationships such as grooming. Given this strong effect of rank on other behaviours and given the strong dependency of rank on social context, the choice of the right party members may be a crucial factor in the fission-fusion processes of free-ranging bonobos.
Keywords: BONOBO PAN PANISCUS; RANK ORDERS; FEEDING SCORES; AGONISTIC RANKS; PEERING
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Uchiyama, H., Ohtani, N., & Ohta, M. (2011). Three-dimensional analysis of horse and human gaits in therapeutic riding. Appl. Anim. Behav. Sci., 135(4), 271–276.
Abstract: Therapeutic horse riding or hippotherapy is used as an intervention for treating individuals with mental and physical disabilities. Equine-assisted interventions are based on the hypothesis that the movement of the horse's pelvis during horseback riding resembles human ambulation, and thus provides motor and sensory inputs similar to those received during human walking. However, this hypothesis has not been investigated quantitatively and qualitatively. This study aimed to verify the hypothesis by conducting a three-dimensional analysis of the horse's movements while walking and human ambulation. Using four sets of equipments, we analysed the acceleration patterns of walking in 50 healthy humans and 11 horses. In addition, we analysed the exercise intensity by comparing the heart rate, breathing rate, and blood pressure of 127 healthy individuals before and after walking and horse riding. The acceleration data series of the stride phase of horse walking were compared with those of human walking, and the frequencies (in Hz) were analysed by Fast Fourier transform. The acceleration curves of human walking overlapped with those of horse walking, with the frequency band of human walking corresponding with that of horse walking. Exercise intensity, as measured by the heart rate and breathing rate, was not significantly different between horse riding and human walking. The levels of diastolic blood pressure were slightly higher during horse riding than during walking, but were lower during both conditions compared with those in normal conditions (P < 0.01). The present study shows that, although not completely matched, the accelerations of the horse and human walking are comparable quantitatively and qualitatively. Horse riding at a walking gait could generate motor and sensory inputs similar to those produced by human walking, and thus could provide optimum benefits to persons with ambulatory difficulties.
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