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Lansade, L., & Bouissou, M. - F. (2008). Reactivity to humans: A temperament trait of horses which is stable across time and situations. Appl. Anim. Behav. Sci., 114(3-4), 492–508.
Abstract: The aim of this study was to test the existence of a “reactivity-to-humans” trait which is stable over time and across situations. For this purpose, four test situations were repeated on the same animals at 8 months, 1.5 years (N = 110 horses) and 2.5 years of age (N = 54 horses). These situations involved a “familiar passive human test” during which a familiar person stayed motionless in the test pen, a “familiar active human test” and an “unfamiliar active human test” during which a familiar or unfamiliar person attempted to touch the horse, and a “halter-fitting and heart-rate measurement test”, during which the horse was haltered and equipped with a surcingle and then had its heart-rate measured. Many behavioural variables were recorded during the tests, and correlations between ages and variables measured during other situations were investigated. When correlations were found, we concluded that the variables corresponded to temperament traits which are stable over time and across situations. The horses tested were divided into four groups according to breed (Anglo-Arab and Welsh) and year of birth (2001 and 2002). Data for each group were analysed separately with Spearman rank correlations. Regarding stability over time, there was a strong correlation between ages in the frequency of licking/nibbling the passive human, the time taken by a familiar or unfamiliar person to touch the horse, and the time taken to fit a halter (e.g. the time taken to touch the horse's muzzle was significantly correlated between ages in 7 out of the 8 possible cases: 0.40 <= R <= 0.67). Regarding stability across situations, results indicate that the more frequently a horse sniffed, licked or nibbled a passive human, the easier it was for either a familiar or unfamiliar active human to touch and halter it. Moreover, a horse which could be touched easily by a familiar human could also be touched and haltered easily by an unfamiliar human. These behaviours, which are linked directly to humans, therefore indicate good stability over time and across situations. In conclusion, this stability suggests the existence of a “reactivity-to-humans” temperament trait, whether the human is passive or active, familiar or unfamiliar. When the animals do not have frequent contact with humans, this reactivity is stable over time and can be measured as early as 8 months of age using the behavioural variables cited above.
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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.
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King, S. R. B. (2002). Home range and habitat use of free-ranging Przewalski horses at Hustai National Park, Mongolia. Appl. Anim. Behav. Sci., 78(2-4), 103–113.
Abstract: Przewalski horses (Equus ferus przewalskii), also known as takhi, were first re-introduced to the wild in Hustai National Park, Mongolia, in 1994. Since then the number of free harems increased to a maximum of seven; there are currently six (October 2000). The size of the home range of each of the harems changed among years and among seasons. The horses tended to settle in a home range close to where they were released although they explored the surrounding area. The use of the habitat within each home range changed through the day, with the horses grazing in the valleys during the morning and evening, and moving to higher places to stand rest and use as a refuge from heat and flies during the middle of the day. Range establishment and area, as well as habitat use are discussed.
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Feh, C., & Munkhtuya, B. (2008). Male infanticide and paternity analyses in a socially natural herd of Przewalski`s horses: Sexual selection? Behav. Process., 78(3), 335–339.
Abstract: The sexual selection hypothesis explains infanticide by males in many mammals. In our 11-year study, we investigated this hypothesis in a herd of Przewalski's horses where we had witnessed infanticidal attacks. Infanticide was highly conditional and not simply linked to takeovers. Attacks occurred in only five of 39 cases following a takeover, and DNA paternity revealed that, although infanticidal stallions were not the genetic fathers in four cases out of five, stallions present at birth did not significantly attempt to kill unrelated foals. Infanticide did not reduce birth intervals; only in one case out of five was the infanticidal stallion, the father of the next foal; mothers whose foals were attacked subsequently avoided associating with infanticidal stallions. Therefore, evidence for the sexual selection hypothesis was weak. The “human disturbance” hypothesis received some support, as only zoo bred stallions which grew up in unnatural social groups attacked foals of mares which were pregnant during takeovers.
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Ransom, J. I., & Cade, B. S. (2008). Influences of immunocontraception on intraband social behavior in free-ranging feral horses, Equus caballus. In IESM 2008.
Abstract: Contraception is often considered for population control of wildlife that is otherwise protected from more traditional management strategies, though little is known about the behavioral consequences of contraception in wild populations of socially complex animals. Feral horses, Equus caballus, in the western United States are ideal candidates for contraceptive management due to broad scale federal protection, reported herd growth rates of 15-25% per year, and finite public land allocated for them. We investigated influences of the immunocontraceptive porcine zona pellucida (PZP) on social behavior within bands of feral horses in three discrete populations for 4 years. Each band consisted of a single stallion and a harem of adult females (>1 years old) and their foals, as well as associated yearling males and females. Four mutually exclusive behaviors (herding, reproductive, harem tending, and agonism) were analyzed to investigate the difference between behaviors initiated by band stallions toward control mares and contracepted mares within the bands. Additionally, spatial relationships between each stallion and each harem female were analyzed to assess possible passive interactions. A candidate set of 22 hierarchical mixed effects models, using the discrete populations as a random effect on various fixed effects, was analyzed by restricted maximum likelihood estimation. The most supported models were selected by corrected Akaike"s Information Criteria (AICc). Analyses were done on 3 female age cohorts based on distinct fecundity rates: 2 to 5 year olds, 6 to 14 year olds, and > 14 year olds. Stallions herded control mares in the 6 to 14 year cohort more than contracepted mares (n=128, P=0.037) with treatment being the most supported effect, but foal presence (dependent foal) also contributed significantly to the model. Contracepted mares received more reproductive behaviors than control mares in the 6 to 14 year cohort (n = 151, P=0.020). No differences were detected in herding or reproductive behaviors in
the least fecund groups, the 2 to 5 year olds and ≥ 15 year olds. The only independent variable in the most supported model for reproductive behavior was treatment, and the covariates of foal presence, band residence (resident or transient female), band size, and body condition did not contribute. There were no differences in any age cohort for harem tending or agonism. Spatially, stallions maintained closer relationships with 2 to 5 year old contracepted females than with the same age control females (n=136, P < 0.001) while the group was feeding (at its most dispersed structure), but there were no differences while the band was resting or in locomotion. There were no spatial differences detected in the other age cohorts. Given the polyestrous nature and high fecundity of feral horses, the observed difference in reproductive behaviors between treatment groups was not surprising; however, the difference detected in herding rates was an unexpected behavioral modification. This change in herding behavior suggests that further investigation is needed to determine if PZP immunocontraception has an underlying influence on mare social rank, band structure, and band stability, as well as the scope and long-term importance of these behavioral dynamics.
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Lansade, L., Bouissou, M. - F., & Erhard, H. W. (2008). Reactivity to isolation and association with conspecifics: A temperament trait stable across time and situations. Appl. Anim. Behav. Sci., 106(2-4), 355–373.
Abstract: A temperament trait is generally defined as individual differences in behaviour that are present early in life and relatively stable across situations and over time. The aim of this study was to test the existence of a trait <<gregariousness>> in horses, by testing the stability across situations and over time of the responses to different social events. Sixty-six Welsh ponies and 44 Anglo-Arab horses were successively tested at 8 months and 1.5 years of age. Among them, 33 ponies and 21 horses were also tested at 2.5 years of age. They were submitted to four test situations: isolation and separation from, attraction towards and passing conspecifics. We carried out the analysis using each of four test groups as a unit (e.g. 33 Welsh ponies born in 2001, tested in isolation). Isolation and separation stood out as tests that showed a high consistency within test, across tests and across time. The most interesting behavioural parameter was the frequency of neighing, which was well correlated with other parameters measured in the same tests, such as defecation, locomotion and vigilance, as well as across the 3 years (e.g. for separation test: 0.41 < R < 0.61). Therefore, the behaviour of neighing observed in separation or isolation tests as early as 8 months of age appears to be a good indicator of similar behaviour in similar situations later in life, but also of other behaviours which can render the horse difficult to use. No parameter recorded during the attraction test presented stability across situations and time: the reactions to this test were not the expression of a stable characteristic of the individual and did not reflect the same characteristic as the three other tests. Of the different parameters recorded during the passing conspecifics test, the time to cross the arrival line near conspecifics showed good stability across years (0.35 < R < 0.68). This parameter was also correlated with many others recorded during the same test, and also, to a certain extent, to the frequency of neighing in the isolation and separation tests. This stability across responses expressed in various social contexts, and this stability over time, from 8 months to 2.5 years of age suggest the existence of a trait of gregariousness in the horse. From a practical point of view, that means it is possible to estimate the level of gregariousness of a horse as early as 8 months of age. Furthermore, additional analysis shows that gregariousness decreases with age.
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Moehlman, P. D., Kebede, F., & Yohannes, H. (1998). The African wild ass (Equus africanus): conservation status in the horn of Africa. Appl. Anim. Behav. Sci., 60(2-3), 115–124.
Abstract: From 1989 to 1996, surveys were made in most of the historic range of African wild asses in Somalia, Ethiopia, and Eritrea. From the 1970s to the mid 1990s populations of African wild asses (Equus africanus, Fitzinger, 1857) in Somalia and Ethiopia have declined from approximately 6 to 30 per 100 km2 to 1 or 2 per 100 km2. Given the current IUCN criteria, they are Critically Endangered (CR) and face extremely high risk of extinction in the wild in the immediate future, as their populations have been reduced by at least 80% over the last 10+ years (IUCN, 1994). Basic research is needed on this species as scientific information on its reproductive biology, behavior, ecology, and genetics is very limited. Improved support needs to be provided to existing parks and reserves and new multiple use reserves need to be established.
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Feh, C., Munkhtuya, B., Enkhbold, S., & Sukhbaatar, T. (2001). Ecology and social structure of the Gobi khulan Equus hemionus subsp. in the Gobi B National Park, Mongolia. Biol. Conserva., 101(1), 51–61.
Abstract: The status of the Gobi khulan Equus hemionus subsp. is recorded as “insufficiently known” in the Species Survival Commission's Equid Action Plan. Recent counts confirm that Mongolia holds the most important population of the whole species. Since 1953, the animals have benefited from a protected status, but this is now challenged. A 5-year study in the B part of the Gobi National Park on one subpopulation showed that it has remained stable over the past 15 years with an adequate mean reproductive rate of 15% and a 50% survival rate over the first year. Age/sex related mortality and prey analysis indicate that wolf predation probably has some impact on the population, in particular for 4-6-year-olds of both sexes at the start of reproduction. Desert and mountain steppes are the khulan's year-round preferred habitat, but `oases', play an important role at the beginning of lactation. Anthropogenic factors affect both home range and habitat use through direct intervention or permanent occupation of the scarce water sources. Khulans of this subpopulation, unlike other Asian and African wild asses, form year-round stable, non-territorial families. These families and all-male groups join together into “bands” in winter, and herds of several hundred animals, where reproductive rate is highest, form throughout the year. The existence of such herds may thus be critical for the breeding success of the population. Our study provides the first detailed quantitative data for this subspecies, which will help to monitor changes in the future.
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Pluhacek, J., Bartos, L., & Culik, L. (2006). High-ranking mares of captive plains zebra Equus burchelli have greater reproductive success than low-ranking mares. Appl. Anim. Behav. Sci., 99(3-4), 315–329.
Abstract: Plains zebra live in harems that include one to six adult mares. Between these mares is a strong order of social hierarchy. The social rank of an equid mare is typically correlated with her age. Further, high-ranking captive plains zebra mares produce more surviving offspring than low-ranking mares. The objectives of this study were to, first, examined the factors that influence social rank of captive plains zebra mares, and second, test if high-ranking mares conceive earlier and if they have shorter inter-birth intervals than low-ranking ones. We observed three herds of captive plains zebra (a total of 18 mares) at the Dvur Kralove Zoo, Czech Republic. During the 831 h of observation, we recorded 1713 aggressive interactions (biting and offensive kicking) between the mares. These data were used to determine, for each mare, the total number of mares that dominated her in each period of social stability. The GLMM model revealed that older mares were dominated by a lower number of mares than the younger mares. We also found that the probability that a mare would conceive declined with the increasing number of dominant mares. Further, we tested the relationship between the number of dominant mares and the inter-birth interval using 29 intervals for 15 mares. These inter-birth intervals were divided into two groups. When a stallion was continuously present in the herd, the intervals lasted from one birth to the next birth (natural intervals). When a herd was without a stallion, the intervals lasted from the release of the stallion into the herd to the birth of foal (stallion-influenced intervals). The analysis revealed that the inter-birth intervals decreased with an increasing number of dominant mares and the natural intervals decreased with an increasing number of offspring successfully reared by a mare. This finding is the first one in equids and contributes to the previous findings that suggest that social status influences reproductive success.
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Vervaecke, H., Stevens, J., Vandemoortele, H., Sigurjönsdöttir, H., & De Vries, H. (2007). Aggression and dominance in matched groups of subadult Icelandic horses (Equus caballus). J. Ethol., 25(3), 239–248.
Abstract: Abstract We studied sex differences in the nature of aggression and dominance behaviour in two newly formed groups of 1-year-old Icelandic horses. One herd contained nine geldings, the other nine mares. The groups were matched with regard to dominance-determining traits such as age, weaning age, composition of native herd, social experience, genetic origin, body condition and maternal dominance status. High-ranking individuals of both sexes were more aggressive, high-ranking males were older, and high-ranking females had a better body condition. Frequencies of aggressions were similar in both groups. The mares reacted significantly more by showing submission upon an aggression rather than by not responding or by escalating the aggression. For the geldings, this difference was not observed due to a lower tendency to submit. A linear dominance hierarchy was found in both groups. David`s scores provided additional information regarding cardinal rank distances and were used to calculate steepness of hierarchies. The female hierarchy was somewhat steeper compared to the male hierarchy, suggesting somewhat lower despotism among males. This was mainly a consequence of the lower unidirectionality in male submission. Male contests occurred predominantly between dyads at top and mid positions, suggesting a low degree of acceptance of the hierarchy.
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