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Jameson, K. A., Appleby, M. C., & Freeman, L. C. (1999). Finding an appropriate order for a hierarchy based on probabilistic dominance. Anim. Behav., 57(5), 991–998.
Abstract: Methods of ranking individuals in a dominance hierarchy that use transitivity of relationships may obscure irregularities. Furthermore, these methods use only a small proportion of the information available from dominance encounters. This paper presents an intuitively appealing and easily implemented alternative to existing methods for ordering dominance data, developed from the work of Batchelder et al. (1992Journal of Mathematical Psychology36, 185-212). The procedure presented here is based on a mathematical model of paired comparisons and it involves only simple estimation procedures. We illustrate its use with data on dominance among red deerCervus elaphus, stags. The results indicate that dominance relationships are well characterized by the scale values that the model provides, and, because the method provides predictions for all pairings of animals, dominance predictions also exist for pairs of animals that have yet to be observed. Moreover, the dominance outcomes predicted by the model using the order scale are highly correlated with actual dominance observations at all levels. Overall, the procedure described provides a solution to the problem of identifying an appropriate order for a near-linear dominance hierarchy.
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Gammell, M. P., de Vries, H., Jennings, D. J., Carlin, C. M., & Hayden, T. J. (2003). David's score: a more appropriate dominance ranking method than Clutton-Brock et al.'s index. Anim. Behav., 66(3), 601–605.
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RHO, J. R., SRYGLEY, R. B., & CHOE, J. C. (2004). Behavioral ecology of the Jeju pony (Equus caballus): Effects of maternal age, maternal dominance hierarchy and foal age on mare aggression. Ecol. Res., 19(1), 55–63.
Abstract: On Jeju Island, Korea, dominance hierarchy and maternal care according to maternal age were studied in a herd of Jeju ponies (Equus caballus), consisting of 73 mares, their foals and one stallion. Dominance ranks were nearly linear and increased significantly with the age of mares. Most aggressive encounters involved mares under 5 years old. Mares under the age of 5 years have apparently not established their rank. The mean frequency of aggressive actions of mares per hour increased significantly as the day of parturition approached. Aggressive actions of mares with foals decreased significantly as their foals aged. The overall frequency of aggression of mares with foals also decreased significantly with the age of the mares. Our results suggest that the cost of maternal care is lower for older, more dominant mares than for subordinate ones.
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Feh, C. (1999). Alliances and reproductive success in Camargue stallions. Anim. Behav., 57(3), 705–713.
Abstract: A study of a herd of Camargue horsesEquus caballus, showed that while the majority of high-ranking stallions held single-male harems, some sons of low-ranking mares, being low ranking themselves, formed alliances that could last a lifetime. The two stallions were each other's closest associate and preferential grooming partner. Alliances were based on coalitions in which either both partners confronted an intruder synchronously or the dominant of the pair tended the female(s) while the subordinate simultaneously displayed towards the rival. Alliance partners were of similar age but were not more closely related to each other than to other stallions in the herd. Long-term paternity data revealed that subordinates sired close to a quarter of the foals born into the alliance group, and significantly more foals than low-ranking stallions in the herd adopting a `sneak'-mating strategy. The dominant appeared to benefit from the presence of his subordinate partner. Fights occurred all year round, and the subordinate stallion of each alliance pair fought outside competitors more than twice as often as the dominant. Forming short-term alliances before defending mares on their own may enhance long-term reproductive success for both partners. Other benefits to both partners include higher survivorship of their foals and increased access to proven reproductive mares. These results suggest that the relationship between alliance partners is based on mutualism, but several conditions for reciprocity seem to be fulfilled: the benefit to the dominant (assistance in fights), and the benefit to the subordinate (access to reproduction), are both costly to the other partner and delayed in time.
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Moons, C., Heleski, C. R., Leece, C. M., & Zanella, A. J. (2002). Conflicting Results in the Association Between Plasma and Salivary Cortisol Levels in Foals. Retrieved June 4, 2024, from http://www3.vet.upenn.edu/labs/equinebehavior/hvnwkshp/hv02/zanella.htm
Abstract: Introduction
Glucocorticoids are present in many biological fluids as a free fraction or bound to Corticoid
Binding Globulins (CBG) (Matteri et al, 2000). There are conflicting claims regarding the validity of
saliva as a biological fluid to measure cortisol in horses (Lebelt et al, 1996; McGreevy and Pell, 1998;
van der Kolk et al, 2001). Measuring changes in salivary cortisol levels in normal horses and horses
with Cushing`s disease van der Kolk and collaborators (2001) demonstrated the validity of saliva to
assess adrenal function. Puzzling results were reported by McGreevy and Pell (1998) who suggested
that plasma and salivary cortisol concentrations in horses showing oral stereotypies were correlated
but this association was non-existent in control animals. Investigating the responses of foals to
branding and foot-trimming Zanella et al (unpublished results) were unable to identify a relationship
between plasma and salivary cortisol levels in foals. In several species, levels of cortisol in plasma and
saliva are tightly correlated (Fenske, 1996). Cortisol found in blood consists of a fraction bound to
corticoid binding globulin (CBG) and a free, unbound fraction. Free cortisol represents the
biologically active fraction of this steroid hormone. Salivary cortisol reflects the unbound fraction
found in plasma or serum and it passes readily through the parotid membrane (Riad-Fahmy, 1983;
Horning Walker et al,1977). Unbound steroids transfer rapidly between plasma and saliva
(Walker,1989; Scott et al 1990). Saliva flow-rate does not appear to influence saliva cortisol levels in
different species (Hubert and de Jong-Meyer, 1989; Walker 1989, Scott et a, 1990). In horses, Lebelt
et al (1996) reported that salivary and plasma total cortisol in stallions were correlated. We
hypothesized that changes in salivary cortisol in foals would show a pattern that is correlated to that of
plasma free and plasma total cortisol concentrations in foals. In addition, we anticipated that the lack
of good sampling techniques provides an explanation for the failure in determining the association
between salivary and plasma cortisol in foals.
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Feh, C. (2005). Relationships and Communication in Socially Natural Horse Herds. In D. S. Mills, & S. M. McDonnell (Eds.), The domestic horse : the origins, development, and management of its behaviour. Cambridge: Cambridge University Press 2005.
Abstract: Horses are quite unique. In most mammals, sexes segregate and maintain bonds only during the breeding season (Clutton-Brock, 1989). Some canids, a few rodents and primate species such as gorillas, hamadryas baboons and red howler monkeys are the exception, where the same males stay with the same females all year round and over many breeding seasons. Typically, both sexes disperse at puberty in these species. In horses, it was clearly shown that the causes for female dispersal were incest avoidance and not intra-specific competition (Monard, 1996). As a rule, this is confirmed for mammal species where tenure length by males exceeds the age at first reproduction in females (Clutton-Brock, 1989). When horses are allowed to choose their mating partner freely, the inbreeding coefficient of the offspring is lower than expected should they mate randomly (Duncan et al, 1984).
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Nicol, C. J. (2000). Equine Stereotypies. In: Houpt K.A. (Ed.),. In Recent Advances in Companion Animal Behavior Problems. International Veterinary Information Service.
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Feh, C. (2001). Alliances between stallions are more than just multimale groups: reply to Linklater & Cameron (2000). Anim. Behav., 61, F27–F30.
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Caro, T. M., Graham, C. M., Stoner, C. J., & Vargas, J. K. (2004). Adaptive significance of antipredator behaviour in artiodactyls. Anim. Behav., 67(2), 205–228.
Abstract: We used comparative data to test functional hypotheses for 17 antipredator behaviour patterns in artiodactyls. We examined the literature for hypotheses about auditory and visual signals, defensive behaviour and group-related antipredator behaviour in this taxon and derived a series of predictions for each hypothesis. Next, we documented occurrences of these behaviour patterns and morphological, ecological and behavioural variables for 200 species and coded them in binary format. We then pitted presence of an antipredator behaviour against presence of an independent variable for cervids, bovids and all artiodactyls together using nonparametric tests. Finally, we reanalysed the data using Maddison's (1990, Evolution, 44, 539-557) concentrated-changes tests and a consensus molecular and taxonomic phylogeny. We found evidence that snorting is both a warning signal to conspecifics and a pursuit-deterrent signal, lack of evidence that whistling alerts conspecifics and indications that foot stamping is a visual signal to warn group members. Evidence suggested that tail flagging was a signal to both conspecifics and predators, that bounding, leaping and stotting were used both as a signal and to clear obstacles and that prancing functioned similarly to foot stamping. Analyses of tail flicking, zigzagging and tacking were equivocal. We confirmed that inspection occurs in large groups, freezing enhances crypticity, and species seeking refuge in cliffs tend to be small. Entering water and attacks on predators had few correlates. Finally, group living, a putative antipredator adaptation, was associated with large body size and species living in open habitats, confirming Jarman's (1974, Behaviour, 48, 215-267) classic hypothesis. Bunching and group attack apparently deter predators. Despite limitations, comparative and systematic analyses can bolster adaptive hypotheses and raise new functional explanations for antipredator behaviour patterns in general.
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FitzGibbon, C. D. (1994). The costs and benefits of predator inspection behaviour in Thomson's gazelles. Behav. Ecol. Sociobiol., 34(2), 139–148.
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