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Cameron, E. Z., Linklater, W. L., Stafford, K. J., & Minot, E. O. (2000). Aging and improving reproductive success in horses: declining residual reproductive value or just older and wiser? Behav. Ecol. Sociobiol., 47(4), 243–249.
Abstract: In many mammalian species, female success in raising offspring improves as they age. The residual reproductive value hypothesis predicts that each individual offspring will be more valuable to the mother as she ages because there is less conflict between the current and potential future offspring. Therefore, as mothers age, their investment into individual offspring should increase. Empirical evidence for an influence of declining residual reproductive value on maternal investment is unconvincing. Older mothers may not invest more, but may be more successful due to greater experience, allowing them to target their investment more appropriately (targeted reproductive effort hypothesis). Most studies do not preclude either hypothesis. Mare age significantly influenced maternal investment in feral horses living on the North Island of New Zealand. Older mares, that were more successful at raising foals, were more protective for the first 20 days of life, but less diligent thereafter. Total maternal input by older mothers did not seem to be any greater, but was better targeted at the most critical period for foal survival and a similar pattern was observed in mares that had lost a foal in the previous year. In addition, older mothers were more likely to foal in consecutive years, supporting the hypothesis that they are investing less than younger mares in individual offspring. Therefore, older mothers seem to become more successful by targeting their investment better due to experience, not by investing more in their offspring.
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Linklater, W. L., Cameron, E. Z., Minot, E. O., & Stafford, K. J. (2004). Feral horse demography and population growth in the Kaimanawa Ranges, New Zealand. Wildl. Res., 31(2), 119–128.
Abstract: Although feral horses are a common management problem in numerous countries, detailed and long-term demographic studies are rare. We measured the age and sex structure, and pregnancy, birth and death rates in a population of 413 feral horses in New Zealand during 1994–98 and used them to construct a model simulating population growth. Survivorship increased with age (0–1 years old = 86.8%, 1–2 = 92.3%, 2–4 = 92.4%, ≥? 4 years old = females 94%, males 97% per annum). Birth sex ratio parity, a slight female bias in the adult sex ratio (92 males per 100 females) and higher adult male survivorship indicated lower average survivorship for young males than females that was not detectable in mortality statistics. Pregnancy and foaling rates for mares ≥? 2 years old averaged 79 and 49%, respectively. Foaling rates increased as mares matured (2–3-year-old mares = 1.9%, 3–4 = 20.0%, 4–5 = 42.1%, ≥? 5 = 61.5% per annum). Young mares had higher rates of foetal and neonatal mortality (95% of pregnancies failed and/or were lost as neonatal foals in 2–3-year-old mares, 70.6% in 3–4, 43.2% in 4–5, and 31% in mares ≥? 5 years old). Population growth was 9.6% per annum (9.5–9.8, 95% CI) without human-induced mortalities (i.e. r = 0.092). Our model, standardised aerial counts, and historical estimates of annual reproduction suggest that the historical sequence of counts since 1979 has overestimated growth by ~50% probably because of improvements in count effort and technique.</p>
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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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Cameron, E. Z., Setsaas, T. H., & Linklater, W. L. (2009). Social bonds between unrelated females increase reproductive success in feral horses. Proc. Natl. Acad. Sci. U.S.A., 106(33), 13850–13853.
Abstract: In many mammals, females form close social bonds with members of their group, usually between kin. Studies of social bonds and their fitness benefits have not been investigated outside primates, and are confounded by the relatedness between individuals in primate groups. Bonds may arise from kin selection and inclusive fitness rather than through direct benefits of association. However, female equids live in long-term social groups with unrelated members. We present 4 years of behavioral data, which demonstrate that social integration between unrelated females increases both foal birth rates and survival, independent of maternal habitat quality, social group type, dominance status, and age. Also, we show that such social integration reduces harassment by males. Consequently, social integration has strong direct fitness consequences between nonrelatives, suggesting that social bonds can evolve based on these direct benefits alone. Our results support recent studies highlighting the importance of direct benefits in maintaining cooperative behavior, while controlling for the confounding influence of kinship.
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