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Suagee-Bedore, J. K., Linden, D. R., & Bennett-Wimbush, K. (2021). Effect of Pen Size on Stress Responses of Stall-Housed Horses Receiving One Hour of Daily Turnout. J. Equine Vet. Sci., 98, 103366.
Abstract: Group turnout provides important socializing opportunities for horses, particularly those that are primarily stalled. A high percentage of equine injuries occur during group turnout, which could partly be due to the physical constraints of fencing. To investigate appropriate paddock sizes for group turnouts, horses (n = 12) from a single herd were divided into groups of 4, stalled for 24 hours, and then turned out for 1 hour into one of three differently sized pens: 342, 263, and 184 m2 per horse. Groups rotated through pens across 3 days, receiving one treatment per day. Blood was sampled for cortisol concentrations at 08:00 hours each morning, and then at 15 and 60 minutes into the turn out sessions, and 60 minutes after return to individual stalls. Groups rotated through three turnout times: 09:00, 12:00, 14:00 hours. Counts of agonistic behaviors (chasing, contact biting, and kicking) and low-level threats (pinned ears, tail swishing, bite and kick threats) were recorded. When turned out in pens that provided 342 m2 per horse, horses exhibited reduced plasma cortisol concentrations by 15 minutes after turnout and at 1 hour after return to their stalls (P < .05). Horses in pens providing 184 m2 per horse exhibited greater agonistic (P < .001) and low-level threat (P < .01) behaviors than horses in larger pens. These data provide insight into appropriate pen sizes for horses from established herds. Providing at least 342 m2 per horse may reduce the chance of injury in horses accustomed to group turnout.
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de Waal, F. B. M., & Luttrell, L. M. (1988). Mechanisms of social reciprocity in three primate species: Symmetrical relationship characteristics or cognition? Ethology and Sociobiology, 9(2–4), 101–118.
Abstract: Agonistic intervention behavior was observed in captive groups of chimpanzees (Pan troglodytes), rhesus monkeys (Macaca mulatta), and stumptail monkeys (M. arctoides). Reciprocity correlations of interventions were determined while removing from the data the effects of several symmetrical relationship characteristics, that is, matrillineal kinship, proximity relations, and same-sex combination. It was considered likely that if significant reciprocity persisted after controlling for these characteristics, the reciprocity was based on cognitive mechanisms. Statistical significance was tested by means of recently developed matrix permutation procedures. All three species exhibited significant reciprocity with regard to beneficial interventions, even after controlling for symmetrical traits. Harmful interventions were, however, reciprocal among chimpanzees only. This species showed a “revenge system”, that is, if A often intervened against B, B did the same to A. In contrast, both macaque species showed significantly inversed reciprocity in their harmful interventions: if A often intervened against B, B rarely intervened against A. Further analysis indicates that the strict hierarchy of macaques prevents them from achieving complete reciprocity. Compared to chimpanzees, macaques rarely intervene against higher ranking group members. The observed contrast can be partially explained on the basis of differences in available space, as indicated by a comparison of indoor and outdoor living conditions for the chimpanzee colony. Yet, even when such spatial factors are taken into account, substantial behavior differences between chimpanzees and macaques remain.
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Bonanni, R., Cafazzo, S., Valsecchi, P., & Natoli, E. (2010). Effect of affiliative and agonistic relationships on leadership behaviour in free-ranging dogs. Anim. Behav., 79(5), 981–991.
Abstract: Consensus decisions about the nature and timing of group activities allow animals to maintain group cohesiveness, but also entail costs because individuals often differ with respect to their optimal activity budgets. Two mechanisms whereby animals reach a consensus include ‘consistent leadership’, in which a single dominant individual makes the decision, and ‘variable leadership’ in which several group members contribute to the decision outcome. Sharing of consensus decisions is expected to reduce consensus costs to most group members. Both patterns are thought to emerge from the complexity of social relationships of group members. We investigated the distribution of leadership during group departures in two packs of free-ranging dogs, Canis lupus familiaris, and tested how its distribution between individuals was affected by dominance rank-related affiliative and agonistic relationships. Although leadership was not entirely concentrated on a single group member, both packs had a limited number of habitual leaders. In the largest pack, the pattern of leadership changed from ‘variable’ to nearly ‘consistent’ after its size had shrunk. Habitual leaders were usually old and high-ranking individuals. However, high-ranking dogs that received affiliative submissions in greeting ceremonies were more likely to lead than dominant dogs receiving submissions only in agonistic contexts. During resting times, habitual followers associated more closely with habitual leaders than with other followers. These results suggest that in social species collective movements may arise from the effort of subordinates to maintain close proximity with specific valuable social partners.
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Valderrabano-Ibarra, C., Brumon, I., & Drummond, H. (2007). Development of a linear dominance hierarchy in nestling birds. Anim. Behav., 74(6), 1705–1714.
Abstract: 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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Motch, S. M., Harpster, H. W., Ralston, S., Ostiguy, N., & Diehl, N. K. (2007). A note on yearling horse ingestive and agonistic behaviours in three concentrate feeding systems. Appl. Anim. Behav. Sci., 106(1-3), 167–172.
Abstract: The objective of this study was to compare behaviours of yearling horses fed concentrates under each of three management systems. Over two consecutive years, 16 yearling horses (n = 8/year; 4 fillies, 4 geldings, full siblings between years) were observed over a 60-day trial period/year at 15:30 h each day. The experimental design consisted of three factors (sex, feeder type, and year); repeated measures on feeder type: tire feeders (control system), individual tub feeders, and manger feeders. Frequency of agonistic interaction was affected by feeder type and sex. Fillies performed more than three times the total number of agonistic behaviours per feeding session as geldings. In both years, horses spent the most time eating and had the fewest agonistic interactions when fed in tire feeders.
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McDonnell, S. M., & Haviland, J. C. S. (1995). Agonistic ethogram of the equid bachelor band. Appl. Anim. Behav. Sci., 43(3), 147–188.
Abstract: An ethogram of agonistic and related behaviors among equid bachelor band members was developed. Several key English-language studies on equids were reviewed to derive a preliminary inventory of specific behaviors to be included in the ethogram. A bachelor band of domestic pony stallions pastured together was observed for approximately 50 daylight hours to obtain detailed descriptions of each behavior, enable photographic and video documentation of behaviors, and identify any behaviors to be added to the preliminary inventory. An initial draft of the ethogram was sent to 65 equine researchers for review. Twenty-eight critical reviews were received and their suggestions considered for the final draft. A total of 49 elemental behaviors including five distinct vocalizations was included in the ethogram. Three complex behavioral sequences were also included. Most of the behaviors catalogued from the direct observation of pastured pony stallions were also found in the equid literature. For many, references to these behaviors specifically among males or bachelor band members were not found. The results offer a practical tool for quantitative research and other studies of equid inter-male behavior as well as for teaching of equid behavior, and should facilitate progress toward development of a complete ethogram for the horse and other equids.
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Hoff, M. P., Powell, D. M., Lukas, K. E., & Maple, T. L. (1997). Individual and social behavior of lowland gorillas in outdoor exhibits compared with indoor holding areas. Appl. Anim. Behav. Sci., 54(4), 359–370.
Abstract: The behavior of nine lowland gorillas (Gorilla gorilla gorilla) living in three social groups at Zoo Atlanta was compared in an indoor holding area versus an outdoor exhibit. Focal animal data were collected for each animal during 15 min observation sessions, alternating between indoors and outdoors. A variety of solitary and social behaviors differed in the two conditions. All individual and social behaviors that showed a difference, except eating, occurred more indoors than outdoors. These included aggressive displays, reclining, self manipulation, and social examination of others. Additionally, the gorillas spent more time closer together in the indoor condition. A variety of other behaviors measured did not change between the two environments. There was a clear effect on behavior of the different housing conditions in which the gorillas were kept. It is suggested that the differences in aggressive behavior may be related to environmental complexity. It is further suggested that zoos should be aware that differences in behavior reported by caretaking staff, researchers and visitors may be a reflection of the differing environmental circumstances in which the animals are observed.
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Heitor, F., & Vicente, L. (2010). Dominance relationships and patterns of aggression in a bachelor group of Sorraia horses (Equus caballus). J. Ethol., 28(1), 35–44.
Abstract: Abstract The influence of individual factors on dominance rank and the relationship between rank distance and patterns of aggression predicted by models of evolutionarily stable strategies (ESS) of animal conflict were investigated in a managed bachelor group of Sorraia horses, Equus caballus. The group was composed of four to six stallions 3- to 12-years-old during the study period. The dominance hierarchy was significantly linear and rank was not related to age, weight, height or aggressiveness. Frequency and intensity of agonistic interactions were low, but higher-ranking stallions did not receive lower aggressiveness than lower-ranking stallions. There was some evidence that dominance relationships were more contested among close-ranking stallions, as predicted. Agonistic-related interactions among close-ranking stallions served similar functions to those among distant-ranking stallions, but the latter interacted more frequently than expected for access to resting sites and/or resting partners. Therefore, we found some evidence that agonistic-related interactions among distant-ranking stallions play a larger role in providing access to valuable and defendable resources than those among close-ranking stallions. Nevertheless, the fact that space to escape from aggression was limited and breeding access was independent from dominance rank may have reduced the benefits relative to costs of aggression and therefore limited the occurrence of contests over dominance and resources.
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Beaugrand, J. P. (1997). Relative importance of initial individual differences, victory and defeat experiences, and assessment accuracy during hierarchy formation: A simulation study (Vol. 41). Elsevier.
Abstract: This simulation study explores some conditions leading to transitivity within dominance orders. Combinations of three parameters were varied to study their consequences upon hierarchy formation and upon the degree of linearity of resultant structures. The factors studied were (i) the importance of initial Resource Holding Potentials (RHPs) , (ii) changes brought in RHPs by successive victories and defeats, and (iii) accuracy of RHP assessment made by opponents. Results show that initial differences in RHP always lead to perfectly transitive chains whose rank order reflects the importance of initial differences. Even when simulated animals make important errors while assessing each other during round robin tournaments, emerging dominance structures are perfectly linear and ranks obtained in the structure are highly correlated with initial values in RHPs. Moreover, accumulated experiences of victory and/or defeat alone always lead to perfectly linear hierarchies. Their combination with initial individual differences in RHP led to the same conclusion. Even when assessment was far from being perfect, not only perfect chains were formed but initial values in RHPs significantly influenced rank order when the contribution of victory and defeat to RHP was relatively unimportant. The higher the importance of victory and defeat to RHP as compared to that of initial RHP values, the lower was the correlation between initial RHP values and the ranks order reached by individuals in the resultant hierarchies. In general also, the lower the variation within initial RHPs, the lower was the correlation between initial RHPs and ranks in the hierarchy. At a given level of initial RHP dispersion, increasing the contribution of victory and defeat to RHP diminished the correlation between initial RHP values and obtained ranks. In addition, inaccurate assessment reduced the overall correlation, especially when dispersion of initial RHP values was low and the contribution of victory and defeat was high. These results shed some light on the controversy about the respective roles of initial individual attributes and that of patterns of resolution in the formation of animal hierarchies. We present the emergence of social order within closed systems as those simulated here as a case of self-organization.
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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.
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