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Linklater, W. L., Cameron, E. Z., Minot, E. O., & Stafford, K. J. (1999). Stallion harassment and the mating system of horses. Anim. Behav., 58(2), 295–306.
Abstract: Feral horse, Equus caballus, breeding groups, called bands, usually include one but sometimes up to five stallions. We found that mares were loyal to single-stallion (SS) or multistallion (MS) bands or were social dispersers (maverick mares, Mv). The spacing and social behaviour of mares and stallions in single- and multistallion bands was measured. Indices of mare well-being were also measured including activity budgets (feeding: MS>SS=Mv; resting: MS<SS=Mv), band and mare travel (MS>SS), maternal effort in maintaining contact with foals (MS=Mv>SS), parasite levels in faeces (MS>Mv>SS), body condition (MS=Mv<SS), fecundity (Mv<MS<SS) and offspring mortality (Mv<MS<SS). We present evidence suggesting that the poorer well-being of maverick mares and multistallion band mares results from greater harassment by stallions. Stallion and mare behaviour and poor reproductive success in multistallion bands were not consistent with explanations for the existence of such bands based on cooperation or alternative mating strategies. We suggest an alternative explanation. Stable relationships between mares and a single stallion may enhance reproductive success by reducing aggression between individuals. Therefore, we propose that there is strong selection pressure for stable, long-term stallion-mare relationships, called consort relationships. We propose the consort hypothesis, that multistallion bands are an artefact of selection for stable relationships that occasionally result in more than one such relationship forming, because mares solicit more than one stallion and stallion dominance changes during band formation. Copyright 1999 The Association for the Study of Animal Behaviour.
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Cameron, E. Z., Stafford, K. J., Linklater, W. L., & Veltman, C. J. (1999). Suckling behaviour does not measure milk intake in horses, Equus caballus. Anim. Behav., 57(3), 673–678.
Abstract: Studies of parental investment in mammals have frequently used suckling behaviour to estimate energy transfer from mother to offspring, and consequently to measure maternal input. Such studies assume that the more an offspring sucks, the more milk it will receive. This assumption has been questioned, and a review of the literature found little support for it. To test if suckling behaviour provided an accurate index of milk or energy intake we used a radioactive isotope technique to label the milk of thoroughbred mares and to measure milk transfer to foals. We found no significant linear relationship between usual measures of suckling behaviour and milk or energy intake. No behaviours associated with suckling nor with characteristics of mares and foals improved the relationship; only the number of butts associated with each suck episode even approached significance. If we had used suckling behaviour to test theories on differential maternal investment our conclusions would have been in error. For example, female foals tended to suck for longer than males did but there was no difference in the amount of milk transferred. Consequently, we show that measures of suckling behaviour do not adequately predict milk intake in the domestic horse and we suggest that conclusions about differential maternal investment in mammals based on suckling behaviour are likely to be in error. Copyright 1999 The Association for the Study of Animal Behaviour.
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Cameron, E. Z. (1998). Is suckling behaviour a useful predictor of milk intake? A review. Anim. Behav., 56(3), 521–532.
Abstract: In studies on mammalian parental investment, time spent suckling is often used as a predictor of the milk transferred from mother to infant. It is assumed that the rate of milk transfer is positively correlated with the time spent suckling. However, this assumption has not been tested and empirical studies show conflicting results. Nevertheless, in species in which suckling can readily be observed, time spent suckling is still used to measure milk transfer, although an increasing number of workers recognize that the measure is potentially inaccurate. A meta-analysis on studies that have correlated measures of time spent suckling with milk intake estimates based on weight gain revealed a weak positive relationship and significant heterogeneity between studies. Isotope-labelling techniques for the measurement of milk transfer independent of behaviour have been in use since the 1970s, particularly in studies of species in which suckling is difficult to observe. Only one study has attempted to correlate behavioural measures with independent isotope measures, and it found no relationship between the two measures. I suggest that researchers have avoided such a test as it is unlikely that a strong relationship will be found between milk transfer and suckling behaviour, and I discuss the various factors that confound the relationship and contribute to high heterogeneity between studies. Consequently, the assumption that milk transfer can be measured by time spent suckling has inadequate empirical foundation, and needs to be tested using isotope-labelling methods. Copyright 1998 The Association for the Study of Animal Behaviour
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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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