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Tyler, S. J. (1972). The behaviour and social organisation of the new Forest ponies. Anim. Behav. Monogr., 5(2), 85–196.
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Tomasello, M., Hare, B., & Fogleman, T. (2001). The ontogeny of gaze following in chimpanzees, Pan troglodytes, and rhesus macaques, Macaca mulatta. Anim. Behav., 61(2), 335–343.
Abstract: Primates follow the gaze direction of conspecifics to outside objects. We followed the ontogeny of this social-cognitive skill for two species: rhesus macaques and chimpanzees. In the first two experiments, using both a cross-sectional and a longitudinal design, we exposed individuals of different ages to a human looking in a specified direction. Rhesus infants first began reliably to follow the direction of this gaze at the end of the early infancy period, at about 5.5 months of age. Chimpanzees did not reliably follow human gaze until 3-4 years; this corresponds to the latter part of the late infancy period for this species. In the third experiment we exposed individuals of the same two species to a human repeatedly looking to the same location (with no special object at that location) to see if subjects would learn to ignore the looks. Only adults of the two species diminished their gaze-following behaviour over trials. This suggests that in the period between infancy and adulthood individuals of both species come to integrate their gaze-following skills with their more general social-cognitive knowledge about other animate beings and their behaviour, and so become able to deploy their gaze-following skills in a more flexible manner.
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Tomasello, M., Hare, B., & Agnetta, B. (1999). Chimpanzees, Pan troglodytes, follow gaze direction geometrically. Anim. Behav., 58(4), 769–777.
Abstract: Two experiments on chimpanzee gaze following are reported. In the first, chimpanzee subjects watched as a human experimenter looked around various types of barriers. The subjects looked around each of the barriers more when the human had done so than in a control condition (in which the human looked in another direction). In the second experiment, chimpanzees watched as a human looked towards the back of their cage. As they turned to follow the human's gaze a distractor object was presented. The chimpanzees looked at the distractor while still following the human's gaze to the back of the cage. These two experiments effectively disconfirm the low-level model of chimpanzee gaze following in which it is claimed that upon seeing another animate being's gaze direction chimpanzees simply turn in that direction and look around for something interesting. Rather, they support the hypothesis that chimpanzees follow the gaze direction of other animate beings geometrically to specific locations, in much the same way as human infants. The degree to which chimpanzees have a mentalistic interpretation of the gaze and/or visual experience of others is still an open question.
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Tomasello, M., Call, J., & Hare, B. (1998). Five primate species follow the visual gaze of conspecifics. Anim. Behav., 55(4), 1063–1069.
Abstract: Individuals from five primate species were tested experimentally for their ability to follow the visual gaze of conspecifics to an outside object. Subjects were from captive social groups of chimpanzees,Pan troglodytes, sooty mangabeys,Cercocebus atys torquatus, rhesus macaques,Macaca mulatta, stumptail macaques,M. arctoides, and pigtail macaques,M. nemestrina. Experimental trials consisted of an experimenter inducing one individual to look at food being displayed, and then observing the reaction of another individual (the subject) that was looking at that individual (not the food). Control trials consisted of an experimenter displaying the food in an identical manner when the subject was alone. Individuals from all species reliably followed the gaze of conspecifics, looking to the food about 80% of the time in experimental trials, compared with about 20% of the time in control trials. Results are discussed in terms of both the proximate mechanisms that might be involved and the adaptive functions that might be served by gaze-following.
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Todd, I. A., & Kacelnik, A. (1993). Psychological mechanisms and the Marginal Value Theorem: dynamics of scalar memory for travel time. Anim. Behav., 46(4), 765–775.
Abstract: Abstract. The relation between memory for travel time and foraging decisions was studied experimentally. The temporal properties of two environments with patchily distributed food were simulated in the laboratory using pigeons, Columba livia, as subjects. The two environments differed in mean travel time, while the coefficient of variation of travel time and the decelerated function relating cumulative food gain to time in the patch were held constant within and between environments. Each environment contained a uniform mixture of five travel times experienced in a random order. Two of the five travel times were common in both environments. Effects of travel time were studied by comparing prey collected per patch visit (PPV) after various travel times within each environment, and by comparing patch exploitation after equal travel times between environments. Within the environment with long mean travel time (LMT) PPV was positively correlated with the last and the penultimate travel times but not with travel times before that. The increase in PPV per second of last travel time was six times greater than the increase per second of penultimate travel time, implying very steep memory discounting. In the environment with short mean travel time (SMT), there was no correlation between PPV and previous travel times. However, comparisons between environments of visits following travel times common to both environments (thus removing the effect of the last travel time) showed that substantially more prey were taken after equal travel times in the LMT than in the SMT environment. This difference cannot be accounted for by the within-environment effect of penultimate travel time, implying that there is a different, less steeply devalued, effect of the mixture of travel times. A model of information processing based on combining Scalar Expectancy Theory with the predictions of rate maximization under the Marginal Value Theorem is presented. The model can approximate the results obtained in this and previous experiments and provides a framework for further analysis of memory mechanisms of foraging behaviour.
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Tilson, R. L., Sweeny, K. A., Binczik, G. A., & Reindl, N. J. (1988). Buddies and bullies: Social structure of a bachelor group of Przewalski horses. Appl. Anim. Behav. Sci., 21(1-2), 169–185.
Abstract: The Minnesota Zoological Garden established a bachelor (i.e. all-male) group of 8 Przewalski horses (Equus przewalskii) in 1984. Possible methods for mixing males emerged through preliminary introductions and review of the relevant literature. These included: (1) grouping colts (<=3 years old) after removing them from their natal bands; (2) introducing adult stallions to colts in spacious, controlled environments to reduce the risk of combat-related injury or death associated with largely unpredictable adult temperament; (3) introducing more than 1 animal at a time to established groups. Based on the frequency and orientation of agonistic interactions observed among the bachelors, a linear dominance hierarchy was discerned among the 4 adult stallions; all 4 colts were determined to be of equally low or seemingly non-ranking status. Social rank among the adults was most influenced by individual temperament, and contention for rank was limited almost exclusively to the 3 lower-ranking stallions; the alpha male participated in 73% of all agonistic interactions observed, but was identified as the recipient in less than 3% of these interactions. Intra-band associations were detected in the bachelor group. The most prevalent of these was among the 4 colts, found together in over 70% of the observations. Other affinities commonly observed were between the colts (as a sub-group) and different stallions. The dominant stallion, the colts' most frequent companion, actively discouraged the 3 lower-ranking stallions from associating with the colts. Further, the colts were subjected to significantly fewer agonistic behaviors than the non-alpha stallions (per colt=1.05+/-0.24 h-1; per non-alpha stallion=2.01+/-0.04 h-1). These findings support the hypothesis developed by other workers for ungulates that young bachelors are sheltered from aggression by association with an alpha male. With one exception, adult associations were rarely noted. Although combat-related injuries (superficial wounds and temporary lameness) were common among the bachelors, few required medical attention. No permanent disabilities or deaths have occurred since the group's inception. The existence of social organization and intra-band associations in the bachelor group support the suggestion that such groups were once a natural part of this species' social milieu. Bachelor groups are therefore recommended as an ethologically sound, space- and cost-effective means of managing “surplus” (i.e. non-harem) male Przewalski horses.
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Thouless, C. R., & Guinness, F. E. (1986). Conflict between red deer hinds: the winner always wins. Anim. Behav., 34(4), 1166–1171.
Abstract: Dominance relations between free-living, female red deer (hinds) (Cervus elaphus L.) on the Isle of Rhum, Scotland, were investigated. Most interactions were won by the older hind of the pair and this was the case even when both individuals had reached full body size. The younger hind was more likely to be the winner if the conflict was escalated or if the two hinds were strangers, in which case escalation was more frequent than usual. When outside their normal home range, older hinds were much more likely to lose, and younger ones more likely to win, than usual. These results can be best explained by the hinds using previous experience as a cue for conventional resolution of conflict, with the result that dominance relationships established early in life are perpetuated. No such cue is available if the hinds have not previously met.
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Thouless, C. R. (1990). Feeding competition between grazing red deer hinds. Anim. Behav., 40(1), 105–111.
Abstract: The effect of social rank on the feeding behaviour of female red deer, Cervus elaphus L., on the Isle of Rhum, Scotland, was investigated. Hinds were less likely to approach and more likely to leave the vicinity of other individuals if these hinds were dominant to them. Movements away by subordinates were more likely to involve a break from feeding. Feeding rate, as measured by bite rate, increased with distance from dominant neighbours, but was unaffected by the distance to subordinates. It appears that aggressive interactions had little direct effect on access to food. Instead, it is suggested that feeding competition in red deer hinds is largely a passive process, operating through the avoidance of conflict by subordinates.
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Thorne, J. B., Goodwin, D., Kennedy, M. J., Davidson, H. P. B., & Harris, P. (2005). Foraging enrichment for individually housed horses: Practicality and effects on behaviour. Appl. Anim. Behav. Sci., 94(1-2), 149–164.
Abstract: The stabled (UK) or stalled (USA) horse is commonly fed a restricted-forage diet in contrast to the varied ad libitum high-fibre diet it evolved to consume. A low-forage diet has been linked to the performance of stereotypical behaviour and health problems including gastric ulceration and impaction colic (in cases where horses are bedded on straw). Provision of a diet closer to that which the horse is adapted to and which enables more natural feeding behaviour warrants investigation. This trial aimed to establish whether the behavioural effects observed in short-term trials when stabled horses were provided with a multiple forage diet persist over longer periods. It also aimed to develop a practical methodology for maintaining stabled horses under forage-enriched conditions. Nine horses (aged 5-20 years, various breeds), acting as their own controls, participated in an 18-day, cross-over, Latin Square designed trial, in which they received comparable weights of two dietary treatments: a Single Forage (SF, hay) diet and a Multiple Forage (MF) diet (three long-chop and three short-chop commercially available forages). Following a 2-day acclimatisation, horses were maintained on the forage treatments for 7 days. Horses were observed on alternate days, morning and afternoon, during the 25 min following forage presentation. Horses then crossed over onto their second treatment and, following a further 2 days' acclimatisation, the same protocol was followed for a further 7 days. Observations from video were made using The Observer 3.0(R) and SPPS (version 11). Horses on the MF treatment performed foraging behaviour significantly more frequently and for significantly longer periods than horses on the SF treatment. On the MF treatment horses sampled all forages during observations. However, there were significant differences in the frequency and duration of foraging on individual forages, indicating that horses demonstrated individual preferences for particular forages. Stereotypic weaving behaviour only occurred on the SF treatment. The results indicate that the potentially beneficial behavioural effects of short-term multiple forage provision do persist when horses are managed on a MF diet for a 7-day period. They suggest that a MF diet provides a means of enriching the stabled horse's environment, by offering variety and enabling patch foraging behaviour. The methodology proved practical for maintaining horses under forage-enriched conditions and could easily be adopted by horse owners to facilitate foraging behaviour.
Keywords: Horse; Foraging behaviour; Eating; Feeding; Enrichment; Welfare
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Templeton, J. J., & Giraldeau, L. - A. (1996). Vicarious sampling: the use of personal and public information by starlings foraging in a simple patchy environment. Behav. Ecol. Sociobiol., 38(2), 105–114.
Abstract: Group foragers may be able to assess patch quality more efficiently by paying attention to the sampling activities of conspecifics foraging in the same patch. In a previous field experiment, we showed that starlings foraging on patches of hidden food could use the successful foraging activities of others to help them assess patch quality. In order to determine whether a starling could also use another individual's lack of foraging success to assess and depart from empty patches more quickly, we carried out two experimental studies which compared the behaviour of captive starlings sampling artificial patches both when alone and when in pairs. Solitary starlings were first trained to assess patch quality in our experimental two-patch system, and were then tested on an empty patch both alone and with two types of partner bird. One partner sampled very few holes and thus provided a low amount of public information; the other sampled numerous holes and thus provided a high amount of public information. In experiment 1, we found no evidence of vicarious sampling. Subjects sampled a similar number of empty holes when alone as when with the low and high information partners; thus they continued to rely on their own personal information to make their patch departure decisions. In experiment 2, we modified the experimental patches, increasing the ease with which a bird could watch another's sampling activities, and increasing the difficulty of acquiring accurate personal sampling information. This time, subjects apparently did use public information, sampling fewer empty holes before departure when with the high-information partner than when with the low-information partner, and sampling fewer holes when with the low-information partner than when alone. We suggest that the degree to which personal and public information are used is likely to depend both on a forager's ability to remember where it has already sampled and on the type of environment in which foraging takes place.
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