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Linklater, W. L., & Cameron, E. Z. (2000). Tests for cooperative behaviour between stallions. Anim. Behav., 60(6), 731–743.
Abstract: Breeding groups with multiple stallions occur sympatrically with single-stallion breeding groups in feral horse, Equus caballus, populations. Mutualism and reciprocal altruism between stallions have been proposed to explain the origin and functioning of multistallion bands. However, empirical support for these hypotheses is contradictory and incomplete. Furthermore, there are no explicit tests of the predictions that each hypothesis makes about stallion behaviour and social structure. We compared nine multistallion and 18 single-stallion bands in the Kaimanawa Ranges, New Zealand. Compared with agonistic behaviours, affiliative behaviours were relatively unimportant in the relationships between stallions within bands. The number of stallions in the band did not have a positive influence on mare group size, stability, home range quality or reproductive success in bands. Furthermore, there was a positive relationship between aggression ('intolerance') by the dominant towards subordinate stallions and the subordinates' effort in mare group defence ('helping') but a negative relationship between helping effort by subordinates and their proximity to, and mating with, the bands' mares. Therefore, the predictions of the mutualism and reciprocal altruism hypotheses were not supported. Indeed, for some of the predictions we found the opposite outcomes to be true. Multistallion bands had significantly poorer reproductive success, and dominant stallions were less tolerant of subordinates that helped most and reduced their access to mares. Nevertheless, in all other respects Kaimanawa stallions in multistallion bands behaved like those described elsewhere. Thus, we reject cooperative hypotheses for multimale breeding groups in horses and discuss the mate parasitism and consort hypotheses as better alternatives. Copyright 2000 The Association for the Study of Animal Behaviour.
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Cameron EZ, L. W. L. (2000). Individual mares bias investment in sons and daughters in relation to their condition. Anim. Behav., 60(3), 359–367.
Abstract: The Trivers-Willard hypothesis (TWH) predicts that a mother will treat a son or daughter differently depending on her ability to invest and the impact of her investment on offspring reproductive success. Although many studies have investigated the hypothesis, few have definitively supported or refuted it because of confounding factors or an inappropriate level of analysis. We studied maternal investment in sons and daughters in feral horses, Equus caballus, which meet the assumptions of the TWH with a minimum of confounding variables. Population level analyses revealed no differences in maternal behaviour towards sons and daughters. When we incorporated mare condition, we found that sons were more costly to mares in good condition, whereas daughters were more costly to mares in poor condition, although no differences in maternal behaviour were found. However, since the TWH makes predictions about individual mothers, we examined investment by mares who reared both a son and a daughter in different years of the study. Mares in good condition invested more in their sons in terms of maternal care patterns, costs to maternal body condition and costs to future reproduction. Conversely, mares in poor condition invested more in daughters. Therefore, with an appropriate level of analysis in a species in which confounding variables are minimal, the predictions of the Trivers-Willard hypothesis are supported. Copyright 2000 The Association for the Study of Animal Behaviour.
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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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Adams, E. S. (2005). Bayesian analysis of linear dominance hierarchies. Anim. Behav., 69(5), 1191–1201.
Abstract: Studies on social animals often seek to identify dominance hierarchies, in which individuals are ranked according to competitive abilities based on counts of wins and losses in pairwise encounters. I illustrate Bayesian approaches, based on the method of paired comparisons, for determining ranks and for estimating relationships between dominance ability and other attributes. Bayesian inference combines prior probability distributions for each unknown parameter with likelihood functions to produce the joint posterior probability distribution for the quantities of interest. In contrast to nonparametric techniques for inferring ranks, Bayesian models yield measures of certainty for each inference and allow rigorous estimates of correlations between ranks and covariates even when there is considerable uncertainty as to the ranks themselves. A possible objection to the Bayesian approach is that it appears to entail more restrictive assumptions than do simpler methods. However, simulations show that Bayesian inferences are more robust to deviations from these assumptions than are the results of nonparametric methods.
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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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de Vries, H., Stevens, J. M. G., & Vervaecke, H. (2006). Measuring and testing the steepness of dominance hierarchies. Anim. Behav., 71(3), 585–592.
Abstract: In the analysis of social dominance in groups of animals, linearity has been used by many researchers as the main structural characteristic of a dominance hierarchy. In this paper we propose, alongside linearity, a quantitative measure for another property of a dominance hierarchy, namely its steepness. Steepness of a hierarchy is defined here as the absolute slope of the straight line fitted to the normalized David's scores (calculated on the basis of a dyadic dominance index corrected for chance) plotted against the subjects' ranks. This correction for chance is an improvement of an earlier proposal by de Vries (appendix 2 in de Vries, Animal Behaviour, 1998, 55, 827-843). In addition, we present a randomization procedure for determining the statistical significance of a hierarchy's steepness, which can be used to test the observed steepness against the steepness expected under the null hypothesis of random win chances for all pairs of individuals. Whereas linearity depends on the number of established binary dominance relationships and the degree of transitivity in these relationships, steepness measures the degree to which individuals differ from each other in winning dominance encounters. Linearity and steepness are complementary measures to characterize a dominance hierarchy.
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Poisbleau, M., Fritz, H., Valeix, M., Perroi, P. - Y., Dalloyau, S., & Lambrechts, M. M. (2006). Social dominance correlates and family status in wintering dark-bellied brent geese, Branta bernicla bernicla. Anim. Behav., 71(6), 1351–1358.
Abstract: In many gregarious species, including ducks and geese, being dominant provides more benefits than costs, because dominants have better access to resources essential for survival or reproduction. In geese, being in better body condition during migration towards the breeding grounds positively influences reproductive success. However, underlying proximate mechanisms linking prebreeding body condition on the wintering grounds to breeding success remain poorly understood. We investigated social dominance correlates and family status, in three consecutive winters, in a free-ranging, migrating, dark-bellied brent goose population. Families with juveniles dominated pairs, and pairs dominated singletons. Dominance rank did not increase with the number of juveniles per family. Males were dominant over females. Social dominance and reproductive status for a given winter were significantly correlated with body mass, body size and body condition during the previous winter, suggesting that body condition in winter also affects subsequent breeding success and hence also dominance. Levels of testosterone and triiodothyronine were not correlated with immediate or later dominance or reproductive status. We discuss the role of family status as a signal of social status in determining reproductive strategies.
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