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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.,, Linklater, W. L.,, Stafford, K. J.,, & Minot, E. O.,. (2003). Social grouping and maternal behaviour in feral horses (Equus caballus): the influence of males on maternal protectiveness. Behav. Ecol. Sociobiol., 53(2), 92–101.
Abstract: The risk of infant injury or mortality influences maternal behaviour, particularly protectiveness. Mares are found in bands with a single stallion or bands with more than one stallion in which paternity is less certain. We investigated maternal behaviour in relation to band type. Mares in bands with more than one stallion were more protective of their foals, particularly when stallions and foals approached one another. The rate of aggression between the stallion and foal was a significant predictor of maternal protectiveness, and mare protectiveness was significantly correlated with reduced reproductive success in the subsequent year. Mares that changed band types with a foal at foot, or had their band type experimentally altered, were more protective of their foal in multi-stallion bands than they were in single-stallion bands. Equids are unusual amongst ungulates in that infanticide and feticide have been reported. Both occur where paternity has been uncertain, and equid social structure is similar to other species in which infanticide has been reported. Stallions benefit from infanticide as the mare has greater reproductive success in the subsequent year. Stallion aggression is a significant modifier of mare behaviour and maternal effort, probably due to the risk of infanticide.
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Linklater, W. L., Cameron, E. Z., Stafford, K. J., & Veltman, C. J. (2000). Social and spatial structure and range use by Kaimanawa wild horses (Equus caballus: Equidae). New Zealand J. Ecol., 24(2), 139–152.
Abstract: We measured horse density, social structure, habitat use, home ranges and altitudinal micro-climates in the south-western Kaimanawa ranges east of Waiouru, New Zealand. Horse density in the Auahitotara ecological sector averaged 3.6 horses.km-2 and ranged from 0.9 to 5.2 horses.km-2 within different zones. The population's social structure was like that of other feral horse populations with an even adult sex ratio, year round breeding groups (bands) with stable adult membership consisting of 1 to 11 mares, 1 to 4 stallions, and their predispersal offspring, and bachelor groups with unstable membership. Bands and bachelor males were loyal to undefended home ranges with central core use areas. Band home range sizes varied positively with adult band size. Home ranges overlapped entirely with other home ranges. Horses were more likely to occupy north facing aspects, short tussock vegetation and flush zones and avoid high altitudes, southern aspects, steeper slopes, bare ground and forest remnants. Horses were more likely to be on north facing aspects, steeper slopes, in exotic and red tussock grasslands and flush zones during winter and at lower altitudes and on gentler slopes in spring and summer. Seasonal shifts by bands to river basin and stream valley floors in spring and higher altitudes in autumn and winter are attributed to the beginning of foaling and mating in spring and formation of frost inversion layers in winter. Given horse habitat selectivity and the presence of other ungulate herbivores, results from present exclosures are likely to exaggerate the size of horse impacts on range vegetation. Proposals to manage the population by relocation and confinement are likely to modify current social structure and range use behaviour and may lead to the need for more intensive management in the longer term.
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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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Linklater, W. L., Henderson, K. M., Cameron, E. Z., Stafford, K. J., & Minot, E. O. (2000). The robustness of faecal steroid determination for pregnancy testing Kaimanawa feral mares under field conditions. N Z Vet J, 48(4), 93–98.
Abstract: AIMS: To investigate the utility of faecal oestrone sulphate (OS) concentrations for detecting pregnancy in mares during behavioural studies of feral horses, in which the collection and preservation of samples is not immediate. METHODS: Oestrone sulphate concentrations were measured in fresh dung samples collected from 153 free-roaming Kaimanawa mares throughout the year. In addition, multiple samples were taken from the same pile to investigate the reliability of diagnosis from a single sample, as well as the influence of time until preservation on OS concentrations. Samples were also taken before and after a 10mm simulated rainfall event to test for dilution of OS concentrations by rain. Oestrone sulphate concentrations in all samples were measured using an enzyme immunoassay. RESULTS: From approximately 150 to 250 days of gestation, OS concentrations were consistently >80 ng/g in mares which subsequently foaled. Mares which did not foal and had low faecal OS concentrations in multiple samples throughout the year had faecal OS concentrations of 31+/-13 ng/g (mean+/-s.d.) with an upper 95% confidence limit of 57 ng/g. Mares sampled from 1 week before to 1 month after behavioural oestrus, and that did not foal in the previous and subsequent seasons, had OS concentrations of 37+/-32 ng/g (mean+/-s.d.) with an upper 95% confidence limit of 100 ng/g. The standard error of oestrone sulphate concentrations in multiple samples from the same dung pile ranged from 1 to 37% of the mean. This large within-pile variation, however, did not result in incorrect diagnoses from single samples unless mares were within 18 days of parturition. Keeping samples at ambient temperatures for up to 16 hours did not affect OS concentrations. Simulated rainfall caused a 17% mean reduction in OS concentrations, but did not change pregnancy diagnoses. CONCLUSIONS: Faecal OS concentrations >100 ng/g were indicative of pregnancy in Kaimanawa mares. For mares more than 150 days post-mating, OS concentrations <57 ng/g were indicative of non-pregnancy, while concentrations between 57 and 100 ng/g provided an inconclusive diagnosis. A single sample from each dung pile collected within 16 hours of defecation was sufficient to accurately diagnose pregnancy in mares 150-250 days post conception. CLINICAL RELEVANCE: Measurement of OS concentrations in dung samples was a reliable and robust indicator of pregnancy status in feral mares 150-250 days post mating. This corresponds approximately to the period from May to August, given the seasonal breeding pattern in this population. This method of determining pregnancy status is suitable for field use in behavioural and demographic studies of wild horse populations.
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Linklater, W. L., Cameron, E. Z., Stafford, K. J., & Austin, T. (1998). Chemical immobilisation and temporary confinement of two Kaimanawa feral stallions (Vol. 46).
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Chamove, A. S., Crawley-Hartrick, O. J. E., & Stafford, K. J. (2002). Horse reactions to human attitudes and behavior. Anthrozoos, 15(4), 323–331.
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