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Cameron, E.Z.; Setsaas, T.H.; Linklater, W.L. |
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Social bonds between unrelated females increase reproductive success in feral horses |
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2009 |
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Proceedings of the National Academy of Sciences of the United States of America |
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Proc. Natl. Acad. Sci. U.S.A. |
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106 |
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33 |
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13850-13853 |
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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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10.1073/pnas.0900639106 |
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Equine Behaviour @ team @ |
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5152 |
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Clutton-Brock, T.H.; Russell, A.F.; Sharpe, L.L.; Brotherton, P.N.; McIlrath, G.M.; White, S.; Cameron, E.Z. |
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Effects of helpers on juvenile development and survival in meerkats |
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2001 |
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Science (New York, N.Y.) |
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Science |
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293 |
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5539 |
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2446-2449 |
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Animals; Breeding; Carnivora/growth & development/*physiology; *Cooperative Behavior; Feeding Behavior; Female; Male; Survival Rate; *Weight Gain |
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Although breeding success is known to increase with group size in several cooperative mammals, the mechanisms underlying these relationships are uncertain. We show that in wild groups of cooperative meerkats, Suricata suricatta, reductions in the ratio of helpers to pups depress the daily weight gain and growth of pups and the daily weight gain of helpers. Increases in the daily weight gain of pups are associated with heavier weights at independence and at 1 year of age, as well as with improved foraging success as juveniles and higher survival rates through the first year of life. These results suggest that the effects of helpers on the fitness of pups extend beyond weaning and that helpers may gain direct as well as indirect benefits by feeding pups. |
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Department of Zoology, University of Cambridge, Downing Street, Cambridge CB2 3EJ, UK. thcb@hermes.cam.ac.uk |
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0036-8075 |
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PMID:11577235 |
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refbase @ user @ |
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414 |
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Cameron, E.Z. |
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Facultative adjustment of mammalian sex ratios in support of the Trivers-Willard hypothesis: evidence for a mechanism |
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2004 |
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Proceedings. Biological sciences / The Royal Society |
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Proc Biol Sci |
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271 |
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1549 |
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1723-1728 |
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Age Factors; Animals; Body Constitution; *Evolution; Female; Glucose/metabolism/physiology; Litter Size; Male; Mammals/*physiology; *Models, Biological; Reproduction/physiology; Seasons; Sex Factors; *Sex Ratio; Time Factors |
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Evolutionary theory predicts that mothers of different condition should adjust the birth sex ratio of their offspring in relation to future reproductive benefits. Published studies addressing variation in mammalian sex ratios have produced surprisingly contradictory results. Explaining the source of such variation has been a challenge for sex-ratio theory, not least because no mechanism for sex-ratio adjustment is known. I conducted a meta-analysis of previous mammalian sex-ratio studies to determine if there are any overall patterns in sex-ratio variation. The contradictory nature of previous results was confirmed. However, studies that investigated indices of condition around conception show almost unanimous support for the prediction that mothers in good condition bias their litters towards sons. Recent research on the role of glucose in reproductive functioning have shown that excess glucose favours the development of male blastocysts, providing a potential mechanism for sex-ratio variation in relation to maternal condition around conception. Furthermore, many of the conflicting results from studies on sex-ratio adjustment would be explained if glucose levels in utero during early cell division contributed to the determination of offspring sex ratios. |
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Mammal Research Institute, Department of Zoology and Entomology, University of Pretoria, Pretoria 0002, South Africa. ezcameron@zoology.up.ac.za |
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0962-8452 |
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PMID:15306293 |
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refbase @ user @ |
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413 |
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Cameron, E.Z.; Linklater, W.L.; Stafford, K.J.; Minot, E.O. |
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Title |
Maternal investment results in better foal condition through increased play behaviour in horses |
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Journal Article |
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Year |
2008 |
Publication |
Animal Behaviour. |
Abbreviated Journal |
Anim. Behav. |
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76 |
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5 |
Pages |
1511-1518 |
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Equus caballus; feral horse; maternal investment; play |
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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. |
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0003-3472 |
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Equine Behaviour @ team @ |
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4709 |
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Linklater, W.L.; Cameron, E.Z. |
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Title |
Tests for cooperative behaviour between stallions |
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Journal Article |
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Year |
2000 |
Publication |
Animal Behaviour. |
Abbreviated Journal |
Anim. Behav. |
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60 |
Issue |
6 |
Pages |
731-743 |
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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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Ecology Group, Institute of Natural Resources, Massey University |
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0003-3472 |
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PMID:11124871 |
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refbase @ user @ |
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415 |
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Cameron, E.Z.; Stafford, K.J.; Linklater, W.L.; Veltman, C.J. |
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Title |
Suckling behaviour does not measure milk intake in horses, Equus caballus |
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Journal Article |
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Year |
1999 |
Publication |
Animal Behaviour. |
Abbreviated Journal |
Anim. Behav. |
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57 |
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3 |
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673-678 |
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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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Ecology Group, Institute of Natural Resources, Massey University |
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0003-3472 |
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PMID:10196058 |
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refbase @ user @ |
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418 |
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Author |
Cameron, E.Z. |
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Title |
Is suckling behaviour a useful predictor of milk intake? A review |
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Journal Article |
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1998 |
Publication |
Animal Behaviour. |
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Anim. Behav. |
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56 |
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3 |
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521-532 |
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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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Department of Ecology, Massey University |
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0003-3472 |
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PMID:9784199 |
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refbase @ user @ |
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419 |
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Linklater, W.L.; Cameron, E.Z.; Minot, E.O.; Stafford, K.J. |
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Title |
Stallion harassment and the mating system of horses |
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Journal Article |
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Year |
1999 |
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Animal Behaviour. |
Abbreviated Journal |
Anim. Behav. |
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58 |
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2 |
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295-306 |
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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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Ecology Group, Institute of Natural Resources, Massey University |
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0003-3472 |
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PMID:10458881 |
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refbase @ user @ |
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417 |
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Cameron, E.Z.; Linklater, W.L.; Stafford, K.J.; Minot, E.O. |
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Title |
Aging and improving reproductive success in horses: declining residual reproductive value or just older and wiser? |
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Journal Article |
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2000 |
Publication |
Behavioral Ecology and Sociobiology |
Abbreviated Journal |
Behav. Ecol. Sociobiol. |
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47 |
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4 |
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243-249 |
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Maternal investment – Equidae – Equus caballus |
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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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2019 |
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Linklater, W.L.; Cameron, E.Z.; Stafford, K.J.; Veltman, C.J. |
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Social and spatial structure and range use by Kaimanawa wild horses (Equus caballus: Equidae) |
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Journal Article |
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2000 |
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New Zealand Journal of Ecology |
Abbreviated Journal |
New Zealand J. Ecol. |
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24 |
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2 |
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139-152 |
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Bachelor male; Band; Density; Habitat use; Home range; Management proposals; Micro-climate; Vegetation monitoring; habitat use; home range; mammal; social structure; spatial distribution; New Zealand; Equus caballus |
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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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Ecology Group, Institute of Natural Resources, Massey University, Private Bag 11-222, Palmerston North, New Zealand |
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01106465 (Issn) |
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Cited By (since 1996): 12; Export Date: 21 April 2007; Source: Scopus; Language of Original Document: English; Correspondence Address: Linklater, W.L.; Ecology Group; Institute of Natural Resources; Massey University; Private Bag 11-222 Palmerston North, New Zealand; email: wlinklater@hotmail.com |
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