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Christensen, J. W., Zharkikh, T., Ladewig, J., & Yasinetskaya, N. (2002). Social behaviour in stallion groups (Equus przewalskii and Equus caballus) kept under natural and domestic conditions. Appl. Anim. Behav. Sci., 76(1), 11–20.
Abstract: The aim of this study was to investigate social behaviour in differently reared stallions in their respective environments; one group of stallions was reared under typical domestic conditions whereas the other group was reared and lives under natural conditions. The domestic group consisted of 19, 2-year-old stallions (Equus caballus), which were all weaned at 4 months of age and experienced either individual or group housing facilities before being pastured with the other similarly aged stallions. The natural living and mixed age group of Przewalski stallions (E. przewalskii) consisted of 13 stallions, most of which were juveniles (n=11, <=4 years; n=2, >9 years). The domestic group was studied in a 4-ha enclosure at the Danish Institute of Agricultural Sciences and the Przewalski group under free-ranging conditions in a 75-ha enclosure in the Askania Nova Biosphere Reserve, Ukraine. Behavioural data was collected during 168 h of direct observation. The occurrence of 14 types of social interactions was recorded and group spacing behaviour was studied using nearest neighbour recordings. In spite of very different environments, reflecting domestic and natural rearing conditions, many similarities in behaviour was found. Play and play fight behaviour was very similar in the two stallion groups. Quantitative differences were found in social grooming since Przewalski stallions groomed more frequently (P=0.004), and in investigative behaviours, since domestic stallions showed more nasal (P=0.005) and body sniffing (P<0.001), whereas Przewalski stallions directed more sniffing towards the genital region (P<0.001). These differences may, however, be attributed to environmental factors and in the period of time the stallions were together prior to the study period. Quantitative differences appeared in some agonistic behaviours (kick threat, P<0.001; and kick, P<0.001), but data do not support earlier findings of Przewalski horses being significantly more aggressive than domestic horses. In general, Przewalski stallions engaged in more social interactions, and they showed less group spacing, i.e. maintained a significantly shorter distance between neighbours (P<0.001). The study indicates that also domestic horses, which have been reared under typical domestic conditions and allowed a period on pasture, show social behaviour, which is very similar to that shown by their non-domestic relatives.
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Proops, L., Walton, M., & McComb, K. (2010). The use of human-given cues by domestic horses, Equus caballus, during an object choice task. Anim. Behav., 79(6), 1205–1209.
Abstract: Selection pressures during domestication are thought to lead to an enhanced ability to use human-given cues. Horses fulfil a wide variety of roles for humans and have been domesticated for at least 5000 years but their ability to read human cues has not been widely studied. We tested the ability of 28 horses to attend to human-given cues in an object choice task. We included five different cues: distal sustained pointing, momentary tapping, marker placement, body orientation and gaze (head) alternation. Horses were able to use the pointing and marker placement cues spontaneously but not the tapping, body orientation and gaze alternation cues. The overall pattern of responding suggests that horses may use cues that provide stimulus enhancement at the time of choice and do not have an understanding of the communicative nature of the cues given. As such, their proficiency at this task appears to be inferior to that of domestic dogs, Canis lupus familiaris, but similar to that of domestic goats, Caprus hircus.
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Kavar, T., & Dovc, P. (2008). Domestication of the horse: Genetic relationships between domestic and wild horses. Livestock Science, 116(1-3), 1–14.
Abstract: To date, a large amount of equine genetic data has been obtained regarding (i) extant domestic horses of various breeds from all over the world, (ii) ancient domestic horses, (iii) the extant Przewalski's wild horse, and (iv) the late Pleistocene wild horse from Eurasia and North America. Here, a review of mtDNA and Y chromosome marker analyses is presented in the context of horse domestication. High matrilineal (mtDNA) diversity, which can be found in both extant and ancient (domestic and wild) horses, has suggested that a high number of wild (and tamed) mares were domesticated. Alternatively, Y chromosome marker analysis revealed a single haplotype in all domestic horses analyzed; interestingly even a small population of extant Przewalski's wild horses showed two different Y chromosome haplotypes. It seems that an extreme male population bottleneck occurred due to domestication, while reduction in the female population was only moderate, leaving about 100 distinct haplotypes. For this reason, we speculate that domestication might have started when the appropriate stallion was found or was obtained by selection. Perhaps it had some unusual but special characteristics which could have accelerated the process of domestication. We doubt that only a single Y chromosome haplotype will be found in present-day domestic horses if there are no important differences between the founder stallion/s and the other stallions that were not included in the domestication. In the Eneolithic, tamed and wild mares have probably been spread all over Eurasia, although the number of animals was most likely very low and the populations were limited to a restricted area (e.g., taming centers). Only two subspecies of wild horses (Tarpan and Przewalski's wild horse) have survived up to recently. During the further process of domestication, mares (tamed or wild) were preferentially crossed to stallions having more desirable characteristics. We assume that mares from different regions varied in their morphology due to adaptation to their local environmental conditions. These data might explain rapid expansion of horse populations, as well as their rapid differentiation into various phenotypes during the early phase of domestication.
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Nuñez, C. M. V., Adelman, J. S., Smith, J., Gesquiere, L. R., & Rubenstein, D. I. (2014). Linking social environment and stress physiology in feral mares (Equus caballus): Group transfers elevate fecal cortisol levels. General and Comparative Endocrinology, 196, 26–33.
Abstract: Abstract Feral horses (Equus caballus) have a complex social structure, the stability of which is important to their overall health. Behavioral and demographic research has shown that decreases in group (or band) stability reduce female fitness, but the potential effects on the physiological stress response have not been demonstrated. To fully understand how band stability affects group-member fitness, we need to understand not only behavioral and demographic, but also physiological consequences of decreases to that stability. We studied group changes in feral mares (an activity that induces instability, including both male and female aggression) on Shackleford Banks, NC. We found that mares in the midst of changing groups exhibit increased fecal cortisol levels. In addition, mares making more group transfers show higher levels of cortisol two weeks post-behavior. These results offer insights into how social instability is integrated into an animal’s physiological phenotype. In addition, our results have important implications for feral horse management. On Shackleford Banks, mares contracepted with porcine zona pellucida (PZP) make approximately 10 times as many group changes as do untreated mares. Such animals may therefore be at higher risk of chronic stress. These results support the growing consensus that links between behavior and physiological stress must be taken into account when managing for healthy, functional populations.
Keywords: Equus caballus; Fecal cortisol; Feral mare; Group transfer; Stress; Social instability
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Cameron, E. Z., Linklater, W. L., Stafford, K. J., & Minot, E. O. (2008). Maternal investment results in better foal condition through increased play behaviour in horses. Anim. Behav., 76(5), 1511–1518.
Abstract: Play behaviour is widespread in mammals, but benefits to play have been difficult to demonstrate. Physical training is one of the many proposed hypotheses, suggesting that males and females should play differently, that increased maternal investment should lead to increases in play, and that increases in play should result in physical advantages. In a population of feral horses, Equus caballus, males and females did not differ in their play behaviour except that males initiated more of their play bouts than females. Maternal condition influenced play behaviour only in males, with sons of mothers in good condition playing more. However, when we controlled for maternal effects by comparing a son and a daughter of the same mother, daughters played more when their mother was in poor condition and sons played more when their mother was in good condition. Mothers of foals that played more lost more condition. Therefore, the difference in play behaviour could not be explained by offspring sex or maternal condition alone, but play behaviour mirrored variation in maternal investment. In addition, those individuals that played more survived better and had better body condition as yearlings despite weaning earlier. Since increased activity has been linked to enhanced musculoskeletal development in domestic horses, we suggest that play provides a link between increased maternal investment, increased body condition and future reproductive success in feral horses, and probably in other species.
Keywords: Equus caballus; feral horse; maternal investment; play
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Ransom, J. I., Cade, B. S., & Hobbs, N. T. (2010). Influences of immunocontraception on time budgets, social behavior, and body condition in feral horses. Appl. Anim. Behav. Sci., 124(1-2), 51–60.
Abstract: Managers concerned with shrinking habitats and limited resources for wildlife seek effective tools for limiting population growth in some species. Fertility control is one such tool, yet little is known about its impacts on the behavioral ecology of wild, free-roaming animals. We investigated influences of the immunocontraceptive porcine zona pellucida (PZP) on individual and social behavior in bands of feral horses (Equus caballus) in three discrete populations and used 14 hierarchical mixed effect models to gain insight into the influences of PZP treatment on feral horse behavior. A model of body condition was the strongest predictor of feeding, resting, maintenance, and social behaviors, with treated females allocating their time similarly to control females. Time spent feeding declined 11.4% from low condition to high condition females (F1,154 = 26.427, P < 0.001) and was partially reciprocated by a 6.0% increase in resting (F1,154 = 7.629, P = 0.006), 0.9% increase in maintenance (F1,154 = 7.028, P = 0.009), and 1.8% increase in social behavior (F1,154 = 15.064, P < 0.001). There was no difference detected in body condition of treated versus control females (F1,154 = 0.033, P = 0.856), but females with a dependent foal had lower body condition than those without a foal (F1,154 = 4.512, P = 0.038). Herding behavior was best explained by a model of treatment and the interaction of band fidelity and foal presence (AICc weight = 0.660) which estimated no difference in rate of herding behavior directed toward control versus treated females (F1,102 = 0.196, P = 0.659), but resident females without a dependent foal were herded 50.9% more than resident females with a foal (F3,102 = 8.269, P < 0.001). Treated females received 54.5% more reproductive behaviors from stallions than control mares (F1,105 = 5.155, P = 0.025), with the model containing only treatment being the most-supported (AICc weight = 0.530). Treated and control females received harem-tending behaviors from stallions equally (F1,105 = 0.001, P = 0.969) and agonistic behaviors from stallions equally (F1,105 < 0.001, P = 0.986). Direct effects of PZP treatment on the behavior of feral horses appear to be limited primarily to reproductive behaviors and most other differences detected were attributed to the effects of body condition, band fidelity, or foal presence. PZP is a promising alternative to traditional hormone-based contraceptives and appears to contribute few short-term behavioral modifications in feral horses.
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Nuñez, C. M. V., Adelman, J. S., Mason, C., & Rubenstein, D. I. (2009). Immunocontraception decreases group fidelity in a feral horse population during the non-breeding season. Appl. Anim. Behav. Sci., 117(1-2), 74–83.
Abstract: The behavioral effects of the immunocontraceptive agent porcine zona pellucida (PZP) have not been adequately studied. Important managerial decisions for several species, including the wild horse (Equus caballus), have been based on this limited research. We studied 30 horses on Shackleford Banks, North Carolina, USA to determine the effects of PZP contraception on female fidelity to the harem male. We examined two classes of females: contracepts, recipients of the PZP vaccine (n = 22); and controls, females that have never received PZP (n = 8). We conducted the study during the non-breeding season from December 2005 to February 2006, totaling 102.2 h of observation. Contracepted mares changed groups more often than control mares (P = 0.04). Contracepts also visited more harem groups than did control mares (P = 0.02) and exhibited more reproductive interest (P = 0.05). For both contracepted and control females, the number of group changes (P = 0.01) and number of groups visited (P = 0.003) decreased with the proportion of years mares were pregnant. Our study shows that the application of PZP has significant consequences for the social behavior of Shackleford Banks horses. In gregarious species such as the horse, PZP application may disrupt social ties among individuals and inhibit normal social functioning at the population level.
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Sankey, C., Richard-Yris, M. - A., Leroy, H., Henry, S., & Hausberger, M. (2010). Positive interactions lead to lasting positive memories in horses, Equus caballus. Anim. Behav., 79(4), 869–875.
Abstract: Social relationships are important in social species. These relationships, based on repeated interactions, define each partner's expectations during the following encounters. The creation of a relationship implies high social cognitive abilities which require that each partner is able to associate the positive or negative content of an interaction with a specific partner and to recall this association. In this study, we tested the effects of repeated interactions on the memory kept by 23 young horses about humans, after 6 and 8 months of separation. The association of a reward with a learning task in an interactional context induced positive reactions towards humans during training. It also increased contact and interest, not only just after training, but also several months later, despite no further interaction with humans. In addition, this ‘positive memory’ of humans extended to novel persons. Overall, positive reinforcement enhanced learning and memorization of the task itself. These findings suggest remarkable social cognitive abilities that can be transposed from intraspecific to interspecific social contexts.
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Schneider, G., & Krueger, K. (2012). Third-party interventions keep social partners from exchanging affiliative interactions with others. Anim. Behav., 83(2), 377–387.
Abstract: Third-party interventions are defined as the interruption of dyadic interactions by third animals through direct physical contact, interposing or threats. Previous studies focused on the analysis of interventions against agonistic encounters. However, there have been no evaluations of interventions against affiliative behaviours, particularly in relation to the intervening animal�s social relationships and its social and spatial position. Horses, Equus caballus, are an interesting model species, as interventions against affiliative interactions occur more frequently than against agonistic interactions. In this study, 64 feral horses displayed 67 interventions in affiliative interactions and eight interventions in agonistic interactions within the observation period. We analysed the interventions in affiliative encounters, and found that it was mainly higher-ranking females that intervened in the affiliative interactions of group mates in the stable horse harems. The intervening animals took an active part in affiliative and agonistic encounters within the group, but did not occupy particular social roles or spatial positions. They intervened in affiliative interactions in which group mates with which they had social bonds interacted with other members of the group. They targeted the nonbonded animal and approached the one with which they were socially bonded. We suggest some species use third-party interventions in affiliative interactions to prevent competition for preferred social interaction partners from escalating into more costly agonistic encounters.
Keywords: Equus caballus; horse; rank; social bond; social network; third-party intervention
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R. A. Hopkins. CALIFORNIA WILDLIFE HABITAT RELATIONSHIPS SYSTEM (Vol. M174).
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