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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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Weeks, J. W., Crowell-Davis, S. L., & Heusner, G. (2002). Preliminary study of the development of the Flehmen response in Equus caballus. Appl. Anim. Behav. Sci., 78(2-4), 329–335.
Abstract: The flehmen response is commonly seen in most ungulates as well as in several other species (e.g. felids). The behavior is most often thought to be part of the sexual behavioral repertoire of males. One reigning hypothesis suggests that this behavior allows the male to determine the estrous state of a female through the chemosensory functions of the vomeronasal organ. However, females and young of both sexes also exhibit this behavior. Horse foals most frequently show the flehmen response during their first month of life with colts showing the behavior more often than fillies. This study tested the flehmen response on male and female foals throughout their pre-pubertal period. Foals were separately presented estrous and non-estrous urine weekly during the first month of life and then monthly until they were approximately 7 months of age. No significant differences were found between male and female foals for the following variables: latency to flehmen, duration of flehmen, frequency of flehmen and sniffs.
Keywords: Horse behavior; Flehmen; Foal development
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Ellard, M. - E., & Crowell-Davis, S. L. (1989). Evaluating equine dominance in draft mares. Appl. Anim. Behav. Sci., 24(1), 55–75.
Abstract: The social hierarchy of a herd of 12 draft mares was assessed using agonism in the field, paired-feeding tests and a group-feeding test. Results from the paired-feeding test correlated significantly, but imperfectly, with those from the field. Differential motivation among subjects for the feed and disruption of ambiguous relationships among mares reduced the reliability of the paired-feeding test as a measure of social dominance. Results from the group-feeding test did not correlate significantly with the field hierarchy and only a few mares ever ate from the bucket. Height, weight and age each correlated significantly with rank; a mare's tendency to remain alone did not. Total aggressive scores during the paired-feeding test correlated with rank. However, a high-ranking mare was no more aggressive to each of her subordinates than was a low-ranking mare. Rather, all mares aggressed more against individuals close in rank to themselves and with preferred field associates. In the field, mares associated most with other mares of similar rank.
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Keiper, R. R., & Sambraus, H. H. (1986). The stability of equine dominance hierarchies and the effects of kinship, proximity and foaling status on hierarchy rank. Appl. Anim. Behav. Sci., 16(2), 121–130.
Abstract: Dominance hierarchies were determined in four bands of feral horses living on Assateague Island. The bands varied in size from 10 to 16 horses, and consisted of one stallion, several mares and their offspring. The animals ranged in age from less than 1 to over 18 years. Field observation of all social interactions during the summer of 1981 was used to determine dominance. 1981 hierarchies for three of the bands were compared with hierarchies determined for the same bands in 1978, and showed that hierarchies change over time. Age was significantly correlated with rank. Mares with foals did not rank any higher in the hierarchies than mares without foals. Kinship did not appear to have an effect on dominance rank either, since neither juvenile nor adult offspring ranks correlated with the ranks of their mothers. The band stallion was not the highest-ranking animal of any band, but the location of the stallion peripheral to the main body of the band, the nature of his interactions with band members, and his length of residence in the band may have contributed to his low rank.
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Rendall, D., Seyfarth, R. M., Cheney, D. L., & Owren, M. J. (1999). The meaning and function of grunt variants in baboons. Anim. Behav., 57(3), 583–592.
Abstract: Wild baboons Papio cynocephalus ursinus, give tonal, harmonically rich vocalizations, termed grunts, in at least two distinct, behavioural contexts: when about to embark on a move across an open area ('move' grunts); and when approaching mothers and attempting to inspect or handle their young infants ('infant' grunts). Grunts in these two contexts elicit different responses from receivers and appear to be acoustically distinct (Owren et al. 1997 Journal of the Acoustical Society of America101 2951-2963). Differences in responses to grunts in the two contexts may, then, be due to acoustic differences, reflecting at least a rudimentary capacity for referential signalling. Alternatively, responses may differ simply due to differences in the contexts in which the grunts are being produced. We conducted playback experiments to test between these hypotheses. Experiments were designed to control systematically the effects of both context and acoustic features so as to evaluate the role of each in determining responses to grunts. In playback trials, subjects differentiated between putative move and infant grunts. Their responses based only on the acoustic features of grunts were functionally distinct and mirrored their behaviour to naturally occurring move and infant grunts. However, subjects' responses were in some cases also affected by the context in which grunts were presented, and by an interaction between the context and the acoustic features of the grunts. Furthermore, responses to grunts were affected by the relative rank difference between the caller and the subject. These results indicate that baboon grunts can function in rudimentary referential fashion, but that the context in which grunts are produced and the social identity of callers can also affect recipients' responses. Copyright 1999 The Association for the Study of Animal Behaviour.
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Palombit, R. A., Seyfarth, R. M., & Cheney, D. L. (1997). The adaptive value of 'friendships' to female baboons: experimental and observational evidence. Anim. Behav., 54(3), 599–614.
Abstract: Lactating female baboons, Papio cynocephalusoften maintain close associations with particular males. There are at least three proposed benefits of 'friendships' to females: (1) male protection against potentially infanticidal males; (2) male protection against harassment by dominant females; (3) male attachment to an infant that develops into future care of juveniles. These hypotheses were examined in a population of chacma baboons, P. c. ursinusin which male infanticide accounted for at least 38% of infant mortality. Almost all mothers of young infants formed strong bonds with one or two males with whom they had copulated during the cycle in which they conceived their infants. Females were primarily responsible for maintaining friendships during lactation, but they terminated these relationships if their infants died. In playbacks of females' screams, male friends responded more strongly than control males. They also responded more strongly to the screams of female friends than to the screams of control females. Following an infant's death, however, male friends responded less strongly than control males to the same females' screams. Finally, male friends responded more strongly than control males to playback sequences in which female screams were combined with the threat vocalizations of a potentially infanticidal alpha male, but not when female screams were combined with the threat calls of a non-infanticidal male or the alpha female. Both observations and experiments suggest that the benefits of friendships to females derive from the protection of their infants against infanticide.1997The Association for the Study of Animal Behaviour
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Nicol, C. J., & Pope, S. J. (1999). The effects of demonstrator social status and prior foraging success on social learning in laying hens. Anim. Behav., 57(1), 163–171.
Abstract: Opportunities for social learning within a group of animals are likely to be influenced by the social dynamics of that group. Some individuals may be more influential demonstrators than others even when there are no differences in their skill level or performance. In this study of domestic hens,Gallus gallus domesticus, differences in demonstrator salience were examined. From 24 separate flocks we selected as demonstrators a dominant cockerel, a dominant hen, a mid-ranking hen or a subordinate hen. Demonstrators were pretrained to perform an operant discrimination task to obtain food. Six observers from each flock individually watched the demonstrator perform the task for four 5-min sessions held on consecutive days. On the fifth day observers were tested individually in the operant chamber. We analysed data from 19 flocks, where there were no quantitative differences in demonstrator performance. Observer hens of relatively high social status performed more correct operant pecks than observer hens of relatively low social status. Demonstrator category also had a significant effect on subsequent observer behaviour. Hens that had observed cockerels performed very few general pecks or operant pecks. Hens that had observed dominant hens performed more operant pecks, but hens that had observed sub-ordinate hens performed more general pecks in the chamber. The results suggested either that there was an interaction between dominance and gender in demonstrator salience or that dominant hens might have been influential because of some factor imperfectly associated with their dominance status. A possible candidate was the foraging ability of the dominant hens. In a second experiment using the same protocol, we manipulated the prior foraging success of dominant hens from four additional flocks but this had no significant effect on their subsequent influence as demonstrators.
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Rivera, E., Benjamin, S., Nielsen, B., Shelle, J., & Zanella, A. J. (2002). Behavioral and physiological responses of horses to initial training: the comparison between pastured versus stalled horses. Appl. Anim. Behav. Sci., 78(2-4), 235–252.
Abstract: Horses kept in stalls are deprived of opportunities for social interactions, and the performance of natural behaviors is limited. Inadequate environmental conditions may compromise behavioral development. Initial training is a complex process and it is likely that the responses of horses may be affected by housing conditions. Sixteen 2-year-old Arabian horses were kept on pasture (P) (n=8) or in individual stalls (S) (n=8). Twelve horses (six P and six S) were subjected to a standardized training procedure, carried out by two trainers in a round pen, and 4 horses (two P and two S) were introduced to the round pen but were not trained (C; control). On sample collection day 0, 7, 21 and 28, behavior observations were carried out, blood samples were drawn and heart rates were monitored. Total training time for the stalled horses was significantly higher than total time for the pastured horses (S: 26.4+/-1.5 min; P: 19.7+/-1.1; P=0.032). The stalled group required more time to habituate to the activities occurring from the start of training to mounting (S: 11.4+/-0.96; P: 7.3+/-0.75 min; P=0.007). Frequency of unwanted behavior was higher in the stalled horses (S: 8.0+/-2.0; P: 2.2+/-1.0; P=0.020). Pastured horses tended to have higher basal heart rates on day 0 (S: 74.7+/-4.8; P: 81.8+/-5.3 bpm; P=0.0771). While the physiological data failed to identify differences between housing groups, the behavioral data suggest that pasture-kept horses adapt more easily to training than stalled horses.
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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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Sondergaard, E., & Ladewig, J. (2004). Group housing exerts a positive effect on the behaviour of young horses during training. Appl. Anim. Behav. Sci., 87(1-2), 105–118.
Abstract: In an experiment on the effects of social environment and training on the human-animal relationship, 20 horses were handled according to a defined schedule. Eight horses were housed singly and 12 horses were housed in four groups of 3 horses. Horses were handled three times per week in 10 min sessions from an age of 6 months until 2 years of age during two winter periods. A total of 50 and 70 sessions were given in the first and second period, respectively. Five randomly allocated people performed the training. The training scheme involved leading, tying up, touching, lifting feet, etc. in 43 stages. The horse had to fulfil the performance criteria of each stage in order to get to the next stage. In the first winter period, horses were led to the stable when they had “passed” a stage or after 10 min of training. In the second winter period, horses would start off at stage 1 again, and when they “passed” a stage they went on to the next stage within the same training session. Because of the change in training procedure results were analysed separately for the two winter periods. There was a significant difference between trainers in the number of times they allowed a horse to “pass” a stage within each winter period (χ32, P<0.05; χ32, P<0.001 for the first and the second winter period, respectively). Group housed horses “passed” more stages than single housed horses (17 versus 14; 27 versus 18 in the first and second winter period, respectively; P<0.05 for the interaction). Singly housed horses bit the trainer more frequently than did group housed horses (P<0.01). The responses of group housed horses to training clearly demonstrate the benefits of raising young horses in groups.
Keywords: Horses; Housing; Human-animal relationship; Social environment; Training; Learning
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