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Castley, J. G., & Knight, M. H. (1997). Population status of plains zebra, Equus burchelli, in South African National Parks. Scientific Services, National Parks Board, .
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Dublin, H. T., Sinclair, A. R. E., Boutin, S., Anderson, E., Jago, M., & Arcese, P. (1990). Does competition regulate ungulate populations? Further evidence from Serengeti, Tanzania. Oecologia, 82(2), 283–288.
Abstract: Changes in populations of several ungulate species in the Serengeti-Mara region of East Africa over the past 30 years suggest several hypotheses for their regulation and coexistence. Recent censuses in the 1980s have allowed us to test the hypotheses that: (1) there was competition between wildebeest (Connochaetes taurinus) and Thomson's gazelle (Gazella thomsoni). This predicted that gazelle numbers should have declined in the 1980s when wildebeest were food limited. Census figures show no change in gazelle numbers between 1978 and 1986, a result contrary to the interspecific competition hypothesis; (2) wildebeest and African buffalo (Syncerus caffer) populations were regulated by intraspecific competition for food. Since both populations reached food limitation in the 1970s, the hypothesis predicted that the populations should have been stable in the 1980s. The results confirm these predictions for wildebeest and the buffalo population in the Mara reserve. In the Serengeti the buffalo population declined 41% over the period 1976-1984. The decline was not evenly distributed over the park, some areas showing an 80-90% decline, others no change or an increase in numbers. The decline was associated with proximity to human habitation; (3) an outbreak of the viral disease, rinderpest, in 1982 may have been the cause of the drop in buffalo population. Blood serum samples to measure the prevalence of antibodies were collected from areas of decreasing, stable and increasing populations. If rinderpest was the cause of decrease there should be a negative relationship between the prevalence of rinderpest and the instantaneous rate of increase (r). The results showed no relationship. We conclude that rinderpest was not the major cause of the drop in buffalo numbers. Elephant (Loxodonta africana) numbers dropped 81% in Serengeti in the period 1977-1986. In the Mara there was little change. The evidence suggests that extensive poaching in northern and western Serengeti during 1979-1984 accounted for the drop in both elephant and buffalo numbers.
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Kirkpatrick, J. F., Kasman, L. H., Lasley,, B. L., & Turner, J. W. J. (1988). Pregnancy Determination in Uncaptured Feral Horses. J Wildl Manag, 52(2), 305–308.
Abstract: The urinary excretion of estrone sulfate ($\text{E}{1}\text{S}$) by 25 free-roaming feral horses (Equus caballus) was measured by radioimmunoassay applied to extracts of urine-soaked soil. Twelve of 15 mares having $\text{E}{1}\text{S}$ concentrations >1.0 mg/mg creatinine (x = 2.64 +- 1.02 [SD]) produced foals. All 10 mares with $\text{E}{1}\text{S}$ concentrations <1.0 mg/mg creatinine (x = 0.44 +- 0.26) did not foal. Extracting urine from soil and measuring $\text{E}{1}\text{S}$ and creatinine can be used to determine pregnancy in free-roaming feral horses without the stress of capture or immobilization.
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Hawkes, J., Hedges, M., Daniluk, P., Hintz, H. F., & Schryver, H. F. (1985). Feed preferences of ponies. Equine Vet J, 17(1), 20–22.
Abstract: Preference trials were conducted with mature ponies. In Trial 1, oats were compared with oats plus sucrose. Four of six pony geldings selected oats plus sucrose, but one pony demonstrated a dislike for sucrose and one selected from the bucket on the right side regardless of content. Oats, maize, barley, rye and wheat were compared in Trial 2 using six mature pony mares. Oats were the preferred grain, with maize and barley ranking second and third respectively. Wheat and rye were the least preferred. Even though the ponies demonstrated preference, the total intake at a given meal was not greatly depressed when only the less palatable grains were fed. In Trial 3, pony mares selected a diet containing 20 per cent dried distillers' grain and 80 per cent of a basal mixed diet of maize, oats, wheat bran, soybean meal, limestone and molasses over 100 per cent basal mixed diet, but selected the basal diet over diets containing 20 per cent blood meal, beet pulp or meat and bone meal and 80 per cent basal diet. They did not differentiate against diets containing 20 per cent alfalfa meal or 10 or 5 per cent meat and bone meal when the diets were compared to the basal mixed diet.
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Klingel, H. (1975). Social organization and reproduction in equids. J Reprod Fertil Suppl, (23), 7–11.
Abstract: There are two distinct types of social organization and, accordingly, two types of mating systems in equids. In the horse, Plains zebra and Mountain zebra, the adults live in non-territorial and cohesive one-male groups and in stallion groups. The family stallions have exclusive mating rights which are respected by all others. In Grevy's zebra and in the African and Asiatic wild asses, the stallions are permanently territorial and have exclusive mating rights within their territories. Ecological and evolutionary aspects are discussed.
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Talbot, L. M., & Talbot, M. H. (1963). The Wildebeest in Western Masailand.
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Sigurjónsdóttir, H., van Dierendonck, M. C., Snorrason, S., & Thórhallsdóttir, A. G. (2003). Social relationships in a group of horses without a mature stallion. Behaviour, 140(6), 783–804.
Abstract: 1. The social relationships in a group of Icelandic horses without a mature stallion were studied. The horses were all familiar to each other. Mutual grooming and play relationships, spatial associations, dominance-subordinate relations and the effect of kinship on these relationships were analysed.TAGSTARTBRTAGEND 2. The social structure was clearly dominated by the behaviour of the adult mares. The horses preferred to form bonds within their social class (sex/age) and they kept close proximity with their friends. The group was effectively divided into two social subgroups, adult mares as one group and adult geldings and sub-adults as another group. The sub-adults and adult geldings formed associations, which were based on mutual grooming and play, while the adult mares did not play. Differences between the sexes were evident. Males played more than the females, had more playing partners and were more popular as playmates.TAGSTARTBRTAGEND 3. Aggression rates were low. The dominance hierarchy was linear. Adult mares ranked higher than adult geldings, sub-adults and the foals. Rank was significantly correlated with age. The closer the adult mares were in rank, the more they groomed with each other. Such relationships were not found amongst the other social group.TAGSTARTBRTAGEND 4. Kinship was calculated between all pairs of animals for up to 4 or 5 generations. Allogrooming and play frequencies and proximity were all positively correlated with kinship. Adult mares, which were close in the dominance hierarchy, were on average more related than those further apart.TAGSTARTBRTAGEND 5. The social relationships in the Icelandic herd were, to some extent, different from relationships reported from unmanaged and feral horse-herds with mature stallions and bachelors. Our results suggest that adult mares groom more in groups without a stallion. Furthermore, they have more preferred partners than in natural harems and their partners are other adult mares, not their weaned offspring as seems to be the case in feral herds. The sub-adults also seem to be more socially active in the absence of stallions. Interestingly, in the Icelandic group, the adult mares showed stallion like behaviours, like mounting and protecting foals. Only by studying the behaviour and the nature of the relationships of horses in groups of different compositions, can we expect to gain a comprehensive understanding about individual social strategies and cognitive capabilities of the species. Such knowledge is valuable for management and welfare of the horse.
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VanDierendonck, M. C., de Vries, H., Schilder, M.B.H. (1995). An Analysis of Dominance, Its Behavioural Parameters and Possible Determinants in a Herd of Icelandic horses in captivity. Netherl. J. Zool., 45(3-4), 362–385.
Abstract: Feral horses are social animals, which have to rely on survival strategies centered on the formation of cohesive social bonds within their bands. Many problems in the husbandry of social animals such as horses, are due to the fact that the limits of their adaptive abilities are exceeded. Evidence suggests that the fundamental social characteristics of domestic horses have remained relatively unchanged. The social structure, social strategies and social interactions were investigated (3 non-consecutive years, 24 hr per day for several weeks) in long term established groups of domestic horses (mares and geldings of all ages) and a few small introduced groups, kept in (semi)natural environments. The general aim was to investigate the social needs of domestic horses. The social life of domestic horses was characterised by long lasting bonds with preferred partners which were established and maintained by allogrooming, play, proximity and dominance behaviours. Bonding partners were mainly found within the same sex-age group, but adult geldings also bonded with sub-adult mares and geldings. Adult mares were clustered in a group, while the other animals formed a second group. Among the adult mares, subgroups according to reproductive state were formed. Individuals regulated their social network by interfering with interactions between other members of the herd, which in itself is complex. An intervention is a behavioural action of one animal that actively interferes with an ongoing interaction between a dyad with the apparent aim of altering that interaction. This was verified by post-hoc analyses of disturbed and undisturbed interactions. Interventions in allogrooming or play were performed significantly more often when at least one member of the initial dyad was a preferred partner of, or familiar to (within the small introduced bands) the intervener. The stronger the preferred association in allogrooming between the intervener and member(s) of the initial dyad, the higher the probability the intervener would displace one initial member and continue allogrooming with the other. Just five behaviours were extracted which reliably reflected the dominance relations among horses. Aggression with the hind quarters was used both offensively and defensively and therefore not suitable as a reliable parameter. Individual dominance relationships were related to social experience. The implications of these findings for horse husbandry were assessed. It is argued that the execution of affiliative behaviours may be rewarding in itself, and therefore always will be a highly motivated behaviour. It is shown that social positive physical interactions (allogrooming, play) with other horses is an ethological need and therefore indispensable in modern husbandry systems. Ethological needs are so important for the animal that husbandry systems that lack the possibilities to execute such behaviours will cause chronic stress. It is concluded that all horses need physical social contact, and that horses, which lack appropriate social learning experiences during ontogeny, may be hampered in their social functioning later in life. Solutions for problems, including dominance problems, in individual social housing and group housing are presented.
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Moehlman, P. D., Kebede, F., & Yohannes, H. (1998). The African wild ass (Equus africanus): conservation status in the horn of Africa. Appl. Anim. Behav. Sci., 60(2-3), 115–124.
Abstract: From 1989 to 1996, surveys were made in most of the historic range of African wild asses in Somalia, Ethiopia, and Eritrea. From the 1970s to the mid 1990s populations of African wild asses (Equus africanus, Fitzinger, 1857) in Somalia and Ethiopia have declined from approximately 6 to 30 per 100 km2 to 1 or 2 per 100 km2. Given the current IUCN criteria, they are Critically Endangered (CR) and face extremely high risk of extinction in the wild in the immediate future, as their populations have been reduced by at least 80% over the last 10+ years (IUCN, 1994). Basic research is needed on this species as scientific information on its reproductive biology, behavior, ecology, and genetics is very limited. Improved support needs to be provided to existing parks and reserves and new multiple use reserves need to be established.
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Moehlman, P. D., Fowler, L. E., & Roe, J. H. (1998). Feral asses (Equus africanus) of Volcano Alcedo, Galapagos: behavioral ecology, spatial distribution, and social organization. Appl. Anim. Behav. Sci., 60(2-3), 197–210.
Abstract: Feral asses were studied on Volcano Alcedo, Galapagos Islands, Ecuador, during the wet season of 1980. On the volcano rim during March/April, two stable groups were observed to have a `female (harem) defense' polygynous mating system [Emlen, S.T., Oring, S.W., 1977. Ecology, sexual selection, and the evolution of mating systems. Science 197 (4300), pp. 215-223] and social behavior patterns and feeding ecology similar to feral asses living in a habitat where forage and climate are similar, e.g., Ossabaw Island, Georgia [Moehlman, P.D., 1979. Behavior and ecology of feral asses (Equus asinus). Nat. Geogr. Soc. Res. Rep., 1970, pp. 405-411; Moehlman, P.D., 1997. Feral asses (Equus africanus): intraspecific variation in social organization in arid and mesic habitats. J. Appl. Anim. Behav. Sci., this issue; McCort, W.D., 1980. The feral asses (Equus asinus) of Ossabaw Island, Georgia., PhD Dissertation, Pennsylvania State University, University Park, 219 pp.].
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