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Sundaresan, S. R., Fischhoff, I. R., Dushoff, J., & Rubenstein, D. I. (2007). Network metrics reveal differences in social organization between two fission-fusion species, Grevy's zebra and onager. Oecologia, 151(1), 140–149.
Abstract: For species in which group membership frequently changes, it has been a challenge to characterize variation in individual interactions and social structure. Quantifying this variation is necessary to test hypotheses about ecological determinants of social patterns and to make predictions about how group dynamics affect the development of cooperative relationships and transmission processes. Network models have recently become popular for analyzing individual contacts within a population context. We use network metrics to compare populations of Grevy's zebra (Equus grevyi) and onagers (Equus hemionus khur). These closely related equids, previously described as having the same social system, inhabit environments differing in the distribution of food, water, and predators. Grevy's zebra and onagers are one example of many sets of coarsely similar fission-fusion species and populations, observed elsewhere in other ungulates, primates, and cetaceans. Our analysis of the population association networks reveals contrasts consistent with their distinctive environments. Grevy's zebra individuals are more selective in their association choices. Grevy's zebra form stable cliques, while onager associations are more fluid. We find evidence that females associate assortatively by reproductive state in Grevy's zebra but not in onagers. The current approach demonstrates the utility of network metrics for identifying fine-grained variation among individuals and populations in association patterns. From our analysis, we can make testable predictions about behavioral mechanisms underlying social structure and its effects on transmission processes.
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De Stoppelaire, G. H., Gillespie, T. W., Brock, J. C., & Tobin, G. A. (2004). Use of remote sensing techniques to determine the effects of grazing on vegetation cover and dune elevation at Assateague Island National Seashore: impact of horses. Environ Manage, 34(5), 642–649.
Abstract: The effects of grazing by feral horses on vegetation and dune topography at Assateague Island National Seashore were investigated using color-infrared imagery, lidar surveys, and field measurements. Five pairs of fenced and unfenced plots (300 m2) established in 1993 on sand flats and small dunes with similar elevation, topography, and vegetation cover were used for this study. Color-infrared imagery from 1998 and field measurements from 2001 indicated that there was a significant difference in vegetation cover between the fenced and unfenced plot-pairs over the study period. Fenced plots contained a higher percentage of vegetation cover that was dominated by American beachgrass (Ammophila breviligulata). Lidar surveys from 1997, 1999, and 2000 showed that there were significant differences in elevation and topography between fenced and unfenced plot-pairs. Fenced plots were, on average, 0.63 m higher than unfenced plots, whereas unfenced plots had generally decreased in elevation after establishment in 1993. Results demonstrate that feral horse grazing has had a significant impact on dune formation and has contributed to the erosion of dunes at Assateague Island. The findings suggest that unless the size of the feral horse population is reduced, grazing will continue to foster unnaturally high rates of dune erosion into the future. In order to maintain the natural processes that historically occurred on barrier islands, much larger fenced exclosures would be required to prevent horse grazing.
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Hausberger, M., Bruderer, C., Le Scolan, N., & Pierre, J. - S. (2004). Interplay between environmental and genetic factors in temperament/personality traits in horses (Equus caballus). J Comp Psychol, 118(4), 434–446.
Abstract: The aim of the present study was to broach the question of the relative influence of different genetic and environmental factors on different temperament/personality traits of horses (Equus caballus). The researchers submitted 702 horses to standardized experimental tests and investigated 9 factors, either genetic or environmental. Genetic factors, such as sire or breed, seemed to influence more neophobic reactions, whereas environmental factors, such as the type of work, seemed to play a more dominant role in reactions to social separation or learning abilities. Additive effects were evident, showing how environmental factors may modulate behavioral traits. This study constitutes a first step toward understanding the relative weights of genetic factors and how the environment may intervene in determining individual behavioral characteristics.
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McCutcheon, L. J., & Geor, R. J. (2000). Influence of training on sweating responses during submaximal exercise in horses. J Appl Physiol, 89(6), 2463–2471.
Abstract: Sweating responses were examined in five horses during a standardized exercise test (SET) in hot conditions (32-34 degrees C, 45-55% relative humidity) during 8 wk of exercise training (5 days/wk) in moderate conditions (19-21 degrees C, 45-55% relative humidity). SETs consisting of 7 km at 50% maximal O(2) consumption, determined 1 wk before training day (TD) 0, were completed on a treadmill set at a 6 degrees incline on TD0, 14, 28, 42, and 56. Mean maximal O(2) consumption, measured 2 days before each SET, increased 19% [TD0 to 42: 135 +/- 5 (SE) to 161 +/- 4 ml. kg(-1). min(-1)]. Peak sweating rate (SR) during exercise increased on TD14, 28, 42, and 56 compared with TD0, whereas SRs and sweat losses in recovery decreased by TD28. By TD56, end-exercise rectal and pulmonary artery temperature decreased by 0.9 +/- 0.1 and 1.2 +/- 0.1 degrees C, respectively, and mean change in body mass during the SET decreased by 23% (TD0: 10.1 +/- 0.9; TD56: 7.7 +/- 0.3 kg). Sweat Na(+) concentration during exercise decreased, whereas sweat K(+) concentration increased, and values for Cl(-) concentration in sweat were unchanged. Moderate-intensity training in cool conditions resulted in a 1.6-fold increase in sweating sensitivity evident by 4 wk and a 0.7 +/- 0.1 degrees C decrease in sweating threshold after 8 wk during exercise in hot, dry conditions. Altered sweating responses contributed to improved heat dissipation during exercise and a lower end-exercise core temperature. Despite higher SRs for a given core temperature during exercise, decreases in recovery SRs result in an overall reduction in sweat fluid losses but no change in total sweat ion losses after training.
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Robert, N., Walzer, C., Ruegg, S. R., Kaczensky, P., Ganbaatar, O., & Stauffer, C. (2005). Pathologic findings in reintroduced Przewalski's horses (Equus caballus przewalskii) in southwestern Mongolia. J Zoo Wildl Med, 36(2), 273–285.
Abstract: The Przewalski's horse (Equus caballus przewalskii) was extinct in the wild by the mid 1960s. The species has survived because of captive breeding only. The Takhin Tal reintroduction project is run by the International Takhi Group; it is one of two projects reintroducing horses to the wild in Mongolia. In 1997 the first harem group was released. The first foals were successfully raised in the wild in 1999. Currently, 63 Przewalski's horses live in Takhin Tal. Little information exists on causes of mortality before the implementation of a disease-monitoring program in 1998. Since 1999, all dead horses recovered (n = 28) have been examined and samples collected and submitted for further investigation. Equine piroplasmosis, a tick-transmitted disease caused by Babesia caballi or Theileria equi, is endemic in Takhin Tal and was identified as the cause of death of four stallions and one stillborn foal. In December 2000, wolf predation was implicated in the loss of several Przewalski's horses. However, thorough clinical, pathologic, and bacteriologic investigations performed on dead and surviving horses of this group revealed lesions compatible with strangles. The extreme Mongolian winter of 2000-2001 is thought to have most probably weakened the horses, making them more susceptible to opportunistic infection and subsequent wolf predation. Other occasional causes of death since 1999 were trauma, exhaustion, wasting, urolithiasis, pneumonia, abortion, and stillbirth. The pathologic examination of the Przewalski's horses did not result in a definitive diagnosis in each case. Several disease factors were found to be important in the initial phase of the reintroduction, which could potentially jeopardize the establishment of a self-sustaining population.
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Nocera, J. J., Forbes, G. J., & Giraldeau, L. - A. (2006). Inadvertent social information in breeding site selection of natal dispersing birds. Proc Biol Sci, 273(1584), 349–355.
Abstract: Several species use the number of young produced as public information (PI) to assess breeding site quality. PI is inaccessible for synchronously breeding birds because nests are empty by the time the young can collect this information. We investigate if location cues are the next best source of inadvertent social information (ISI) used by young prospectors during breeding site choice. We experimentally deployed ISI as decoys and song playbacks of breeding males in suitable and sub-optimal habitats during pre- and post-breeding periods, and monitored territory establishment during the subsequent breeding season for a social, bobolink (Dolichonyx oryzivorus), and a more solitary species, Nelson's sharp-tailed sparrow (Ammodramus nelsoni). The sparrows did not respond to treatments, but bobolinks responded strongly to post-breeding location cues, irrespective of habitat quality. The following year, 17/20 sub-optimal plots to which bobolink males were recruited were defended for at least two weeks, indicating that song heard the previous year could exert a “carry-over attraction” effect on conspecifics the following year. Sixteen recruited males were natal dispersers, as expected when animals have little opportunity to directly sample their natal habitat quality. We suggest that differences in breeding synchronicity may induce an equivalent clinal distribution of ISI use.
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Huber, L., & Gajdon, G. K. (2006). Technical intelligence in animals: the kea model. Anim. Cogn., 9(4), 295–305.
Abstract: The ability to act on information flexibly is one of the cornerstones of intelligent behavior. As particularly informative example, tool-oriented behavior has been investigated to determine to which extent nonhuman animals understand means-end relations, object affordances, and have specific motor skills. Even planning with foresight, goal-directed problem solving and immediate causal inference have been a focus of research. However, these cognitive abilities may not be restricted to tool-using animals but may be found also in animals that show high levels of curiosity, object exploration and manipulation, and extractive foraging behavior. The kea, a New Zealand parrot, is a particularly good example. We here review findings from laboratory experiments and field observations of keas revealing surprising cognitive capacities in the physical domain. In an experiment with captive keas, the success rate of individuals that were allowed to observe a trained conspecific was significantly higher than that of naive control subjects due to their acquisition of some functional understanding of the task through observation. In a further experiment using the string-pulling task, a well-probed test for means-end comprehension, we found the keas finding an immediate solution that could not be improved upon in nine further trials. We interpreted their performance as insightful in the sense of being sensitive of the relevant functional properties of the task and thereby producing a new adaptive response without trial-and-error learning. Together, these findings contribute to the ongoing debate on the distribution of higher cognitive skills in the animal kingdom by showing high levels of sensorimotor intelligence in animals that do not use tools. In conclusion, we suggest that the 'Technical intelligence hypothesis' (Byrne, Machiavellian intelligence II: extensions and evaluations, pp 289-211, 1997), which has been proposed to explain the origin of the ape/monkey grade-shift in intelligence by a selection pressure upon an increased efficiency in foraging behavior, should be extended, that is, applied to some birds as well.
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Danchin, E., Giraldeau, L. - A., Valone, T. J., & Wagner, R. H. (2004). Public information: from nosy neighbors to cultural evolution. Science, 305(5683), 487–491.
Abstract: Psychologists, economists, and advertising moguls have long known that human decision-making is strongly influenced by the behavior of others. A rapidly accumulating body of evidence suggests that the same is true in animals. Individuals can use information arising from cues inadvertently produced by the behavior of other individuals with similar requirements. Many of these cues provide public information about the quality of alternatives. The use of public information is taxonomically widespread and can enhance fitness. Public information can lead to cultural evolution, which we suggest may then affect biological evolution.
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Dubois, F., & Giraldeau, L. - A. (2003). The forager's dilemma: food sharing and food defense as risk-sensitive foraging options. Am Nat, 162(6), 768–779.
Abstract: Although many variants of the hawk-dove game predict the frequency at which group foraging animals should compete aggressively, none of them can explain why a large number of group foraging animals share food clumps without any overt aggression. One reason for this shortcoming is that hawk-dove games typically consider only a single contest, while most group foraging situations involve opponents that interact repeatedly over discovered food clumps. The present iterated hawk-dove game predicts that in situations that are analogous to a prisoner's dilemma, animals should share the resources without aggression, provided that the number of simultaneously available food clumps is sufficiently large and the number of competitors is relatively small. However, given that the expected gain of an aggressive animal is more variable than the gain expected by nonaggressive individuals, the predicted effect of the number of food items in a clump-clump richness-depends on whether only the mean or both the mean and variability associated with payoffs are considered. More precisely, the deterministic game predicts that aggression should increase with clump richness, whereas the stochastic risk-sensitive game predicts that the frequency of encounters resulting in aggression should peak at intermediate clump richnesses or decrease with increasing clump richness if animals show sensitivity to the variance or coefficient of variation, respectively.
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VanDierendonck, M. C. (2006). Social relationships in a group of horses without a mature stallion (Vol. Chapter 4). Universiteit Utrecht.
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.
Keywords: 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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